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8707
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dbpedia
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2
| 4
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https://www.imdb.com/name/nm0336829/bio/
|
en
|
Michael Gray
|
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[] |
[
"Michael Gray",
"Biography"
] | null |
[
"IMDb"
] | null |
Michael Gray. Actor: Shazam!. Michael Gray was born on 2 September 1951 in Chicago, Illinois, USA. He is an actor, known for Shazam! (1974), Shazam! Fury of the Gods (2023) and Surge of Power: Where There's Smoke (2024). He has been married to Stacy Benon since 1994.
|
en
|
IMDb
|
https://www.imdb.com/name/nm0336829/bio/
|
Michael Gray was born on September 2, 1951 in Chicago, Illinois, USA. He is an actor, known for Shazam! (1974), Shazam! Fury of the Gods (2023) and Surge of Power: Where There's Smoke (2024). He has been married to Stacy Benon since 1994.
|
|||||
8707
|
dbpedia
|
2
| 59
|
https://www.newcomerstlouis.com/obituaries/michael-gray
|
en
|
Michael J. Gray Obituary 2023
|
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[
"Newcomer St. Louis"
] |
2024-06-25T20:14:19
|
Michael Joseph Gray, age 76, of Saint Charles, Missouri passed away in Lake St. Louis, Missouri on Wednesday, May 31, 2023 after his battle with pancreatic cancer. He was born...
|
en
|
https://cdn.filestackcontent.com/6PuEJTwxS0WlUWtsf6jF
|
Newcomer St. Louis
|
https://www.newcomerstlouis.com/obituaries/michael-gray
|
Guestbook
Visits: 1
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8707
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dbpedia
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0
| 40
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https://www.beatport.com/track/most-precious-love/15425545
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en
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Blaze, Barbara Tucker, UDAUFL - Most Precious Love (Michael Gray Remix) [King Street Sounds]
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Download & Stream Blaze, Barbara Tucker, UDAUFL - Most Precious Love (Michael Gray Remix) [King Street Sounds] in highest quality | Find the latest releases here | #1 source for DJ Sets and more
|
en
|
/images/favicon-48x48.png
|
https://www.beatport.com/track/most-precious-love/15425545
|
August/September Top 10 House Charts
Yuichi's August 2021 beatport Top 10 Chart
Get On The Floor Charts (July 2021)
|
|||||
8707
|
dbpedia
|
0
| 56
|
https://www.tiktok.com/%40sofutureclub/video/7265283726077087008
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en
|
Make Your Day
|
[] |
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[] |
[
""
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en
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8707
|
dbpedia
|
3
| 54
|
https://www.ncaa.com/news/basketball-men/article/2022-03-07/michael-jordan-college-stats-best-games-quotes-moments
|
en
|
Michael Jordan: College stats, best games, quotes, moments
|
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[
"Andy Wittry | NCAA.com",
"NCAA.com"
] |
2022-03-07T00:00:00
|
These are the essential facts from Michael Jordan's college basketball career at North Carolina.
|
en
|
/apple-touch-icon.png
|
NCAA.com
|
https://www.ncaa.com/news/basketball-men/article/2022-03-07/michael-jordan-college-stats-best-games-quotes-moments
|
Michael Jordan needs no introduction to basketball fans but for the uninitiated, Jordan is arguably the greatest basketball player of all time. His legendary career includes six NBA championships with the Chicago Bulls but his national championship-winning ways started as a freshman at North Carolina, when he hit the most important shot in the 1982 NCAA title game.
Here's everything you need to know about Michael Jordan's college career at North Carolina.
The vitals on Michael Jordan
School: North Carolina
Position: Guard
Height: 6-6
Weight: 195 pounds
Years active: 1981-84
NCAA tournament record: 8-2
Career averages: 17.7 points per game, 5.0 rebounds per game, 54.0% shooting
season Games FG FGa FG% rebounds points 1981-82 34 5.6 10.5 .534 4.4 13.5 1982-83 36 7.8 14.6 .535 5.5 20.0 1983-84 31 8.0 14.5 .551 5.3 19.6 Career 101 7.1 13.2 .540 5.0 17.7
How many years did Michael Jordan play in college?
Michael Jordan played at North Carolina for three years, from the 1981-82 season to the 1983-84 season. Jordan was just the fourth player in North Carolina history to start his first game as a freshman.
What was Michael Jordan's record in college?
North Carolina went 88-13 in three seasons with Michael Jordan.
What kind of prospect was Michael Jordan coming out of high school?
Michael Jordan ranked fourth on Knoxville newspaper reporter Ken Mink's top-100 rankings of the best high school players in the country — behind Adrian Branch (Maryland), Stuart Gray (UCLA) and Bobby Lee Hurt (Alabama), and one spot ahead of Patrick Ewing (Georgetown). The Tar Heels had landed just six McDonald's All-Americans ever before Jordan enrolled at North Carolina.
How many national championships did Michael Jordan win in college?
Michael Jordan won one national championship in three seasons with North Carolina. The Tar Heels won the 1982 national championship when Jordan was a freshman, thanks to his game-winning jumper against Georgetown.
What was Michael Jordan's game like?
Michael Jordan did it all on the basketball court. That's not to say he was yet the best player in the world when he was a freshman at North Carolina but he was a starter from day one, which was reserved for the very best players, and he averaged 13.5 points per game in his first season with the Tar Heels.
After a 22-point performance against Tulsa, the reigning NIT champion, in early December of his freshman season, The Charlotte Observer's Kevin Quirk wrote, "It was Michael Jordan's kind of game — lots of opportunities in the open court to flash his offensive prowess ... Good things — and some great — were interspersed with those errors. Things such as a potent fast break, superb inside passing, intimidating defense and spectacular individual efforts. Most of those belonged to Jordan, who scored on jump shots, drives, offensive rebounds and a couple of dunks."
Jordan became the team's leading scorer as a sophomore. He averaged 20 points per game, which was roughly a 50 percent increase from his freshman season. He was second in rebounding and averaged more than two steals per game.
He shot 55 percent inside the arc that season and almost 45 percent behind it.
"He truly has no flaws, or so few they aren't worth mentioning, " the Los Angeles Times reported. "He has great jumping ability and a classic jump shot. He can handle the ball on the break. He can do it all."
The Charlotte Observer's Kevin Quirk wrote this in a "Who's up, who's down for ACC tournament" story from March 1984, Jordan's junior season,
"Up: "Michael Jordan's shooting, Michael Jordan's defense, Michael Jordan's moves, Michael Jordan's dunks.
"Down: Anyone who has to guard Michael Jordan."
What were some of Michael Jordan's best games?
Michael Jordan sank the game-winning basket in the 1982 national championship game against Georgetown to give legendary coach Dean Smith his first national championship. It added to the growing legend that was Jordan himself. Jordan scored 16 points on 7-of-13 shooting with a team-high nine rebounds, two assists and two steals in the win. James Worthy led the Tar Heels in that game with a career-high 28.
You can watch the full replay of the game below. Jordan's shot comes at around the 1:19 mark.
Jordan told reporters after the game he had a pregame vision of making the game-winning shot. Jordan said he didn't see the actual shot go in.
"To tell the truth," Jordan said, "I didn't see it go in. I didn't want to look." That shot closed Jordan's freshman season in a full circle, with him scoring the first and the last baskets for the Tar Heels in the 1981-82 campaign.
Jordan scored in double figures in four of North Carolina's five NCAA tournament games that season, including 18 points against Houston in the Final Four. His two greatest scoring outbursts in the NCAA tournament came in the 1983 regional finals, when he scored 26 points to go along with six rebounds before fouling out.
In the Tar Heels' first game of the 1984 NCAA tournament, Jordan's last, he had 27 points on 11-of-15 shooting and six rebounds in a win against Temple. You can watch a compilation of MJ's best NCAA tournament moments below.
Michael Jordan wasn't Michael Jordan from the moment he set foot on campus in Chapel Hill but he was very good. In the fall semester of his freshman year of college, No. 1 North Carolina defeated No. 2 Kentucky by 13 in a game in which Jordan scored 19 points, including him making eight of his last 10 shots.
There's no bigger moment than making a national championship-winning jump shot but in case it wasn't clear, Jordan had a knack for coming up in the clutch in crucial moments. When the defending national champion Tar Heels were in danger of falling to 0-3 to start the 1982-83 season, Jordan stole the ball against Tulane and made a spinning 24-footer to force overtime and North Carolina eventually won in triple-OT.
Two and a half months later, No. 1 North Carolina overcame a 16-point, second-half deficit against No. 3 Virginia (the Tar Heels trailed by 10 with 4:12 left) thanks to Jordan's offensive rebound and putback to bring North Carolina within one, then he forced a steal and dunked home the winning basket. In Jordan's final regular-season ACC game, he scored 25 points to help the Tar Heels complete an undefeated regular season record in conference play as North Carolina held off rival Duke 96-83 in double overtime.
What awards did Michael Jordan win in college?
Here are some of the awards and honors that Michael Jordan received at North Carolina.
1982 ACC Rookie of the Year
1982 national champion
1982 All-Tournament Team
1982 Freshman All-American
1983 First Team All-ACC
1983 East Regional Team
1983 consensus All-American
1983 National Player of the Year (The Sporting News)
1984 First Team All-ACC
1984 ACC Player of the Year
1984 ACC Athlete of the Year
1984 consensus All-American
1984 consensus National Player of the Year
1980s NCAA Tournament All-Decade Team
NCAA Tournament All-Time Team
Named the No. 1 male athlete in ACC history
Named a top-15 player in the 75 Years of March Madness Celebration
What records did Michael Jordan set in college and where does he rank among historical greats?
Here are some of the records and all-time rankings for Michael Jordan in college.
Most points by a sophomore in North Carolina history (721 points)
North Carolina's leader in points, steals in 1982-83 and 1983-84
Tied for fourth in North Carolina history in steals in a season (78 steals)
Sixth in North Carolina history in field goals made in a season (282 field goals made)
Seventh, 34th in North Carolina history in points in a season (721 points)
11th-most points scored by a North Carolina freshman in a season (460 points)
12th in North Carolina history in career scoring average (17.7 points per game)
14th on North Carolina's all-time scoring list (1,788 points)
Tied for 18th in North Carolina history for most points scored in a game (39 points)
Tied for 25th in North Carolina history for highest scoring average in a season (20.0 points per game)
RELATED: The complete results from the best #MarchMadnessMoments bracket
What did people say about Michael Jordan?
North Carolina coach Dean Smith: "Michael Jordan has adapted very well to our type of basketball. He's different than Walter Davis or Al Wood or David Thompson so I think those are unfair comparisons. He's got a lot to learn, but he can be an outstanding player." (Dec. 3, 1981)
Smith: "He has talent. He's a very quick learner. He's a bright student, which passes on to the court and a remarkable young man who has fit in extremely well." (Dec. 26, 1981)
Smith: "Michael is a late bloomer. He improves every month. Look at his defense. When he was a freshman, he had a lot of work to do. Last year, he won the defensive award 12 times. He's going to be one of our great defensive players, although I don't think I'll tell him that until he graduates. But you can just watch him get better and better. When he was a freshman, he ran the 40 in 4.8 (seconds). This year, he ran it in 4.3." (Nov. 22, 1983)
North Carolina teammate Matt Doherty: "He's a perfect player. When I grew up, I wanted to be 6-7 and I wanted to be able to do anything on the court. I wanted to grow up to be what Michael Jordan grew up to be."
North Carolina teammate Buzz Peterson: "I remember telling people that he'd be the Dr. J of our time."
Buzz Peterson: "It's not just basketball. He hates to lose at anything. If we're playing cards of backgammon, we'll have to play all night until he wins."
Former North Carolina teammate James Worthy: "He'll be a star the instant he steps into the NBA. There's no question that he'll be great. We always knew that.'
Jordan's father, James Jordan: "I think Michael got so good because (his brother) Larry used to beat him all the time. He took it hard. Michael didn't start beating him till he started to really grow."
James Jordan: "When he comes home to Wilmington, people treat him like he's a god."
Fort Worth Star-Telegram's Gil Lebreton: "On this night, there was a Michael Jordan, too. With teammate Sam Perkins having a relatively sub-par night, it took Jordan, another freshman, to shoulder the load. It was Jordan who made the interception that led to (James) Worthy's final stuff, and it was Jordan who scored Carolina's memorable final two baskets, a rainmaker, arching jumper over (Patrick) Ewing with 3:26 to play, and a dead-solid perfect 16-footer from the corner with just :15 left. He finished with 16 points, a team-high nine rebounds, and a two-fisted hug from coach Dean Smith. For Carolina, Jordan is a good peek into the future as it could possibly find." (March 30, 1982)
OTHER GREATS: UCLA's Bill Walton | Michigan State's Magic Johnson | Indiana State's Larry Bird
What are some quotes from Michael Jordan?
Michael Jordan on height: "I always wanted to be tall. I thought 6-8 would be nice."
Jordan on his meteoric rise from high school to college: "I wonder sometimes myself if it isn't all a dream. I expect to wake up sometime."
Jordan on his college recruitment: "I never thought I'd be able to play at a Division I school. Nobody from my high school ever had before. It really shocked me when North Carolina started recruiting me. I never thought that could happen."
Jordan on competing at his first basketball camp: "I was so nervous my hands were sweating. I saw all these All-Americans and I thought I was just the lowest thing on the totem pole. Here I was, a country boy from Wilmington. But the more I played, the more confident I became. I thought to myself, 'Maybe I can play with these guys.'"
Jordan on what people back home were saying when he got to North Carolina: "The people back home, stardom was the last thing they saw for me. People said I'd go there and sit the bench and never get to play. I kind of believed 'em myself, but it was a challenge for me."
Jordan, after coach Dean Smith tried to get Jordan to keep his tongue in his mouth when he played: "I haven't bitten it off yet."
In November of his last season in college, Jordan on winning: "After my freshman year, when we won it, I figured that was the way it was supposed to be. Everyone else was going crazy and I was just acting normal. I didn't know any better. I didn't know the impact of it, what we'd done, or how special it was. I was a little boy in a man's body. But when we lost last year, I realized how much winning meant. I realized how hard I had had to work and how hard the team had had to work. I think it makes this season more special."
Jordan on winning: "I hate to lose. I guess that's it as much as anything."
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https://treemily.com/blog/the-shelbys-family-tree-and-the-truth-about-the-peaky-blinders-leader/
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en
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Peaky Blinders Family Tree: The Shelby Family
|
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2023-09-15T06:42:10+00:00
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Find out more about the Peaky Blinders’ leader Thomas Shelby and review Peaky Blinders Family Tree in the Treemily blog post!
|
en
|
Treemily
|
https://treemily.com/blog/the-shelbys-family-tree-and-the-truth-about-the-peaky-blinders-leader/
|
The Peaky Blinders series went out with a bang in 2022, concluding the thrilling story about England’s most feared crime syndicate of the early 20th century. If you’re keen on rewatching the series and would like to refresh your memory on all the prominent characters – you’re on the right webpage. Let’s take a closer look at the Shelby family tree (and more), with a spoiler alert, of course. Also, for your own family tree visualizations, don’t forget to sign up for our Family Tree Builder and try it for free.
What Is the Show About?
Peaky Blinders is a British crime drama series released in 2013. The show tells the story of the Shelby gangster family of Irish descent who lives in Birmingham, England. Their story begins in 1919 just after the First World War ends. The leader of the gang and the main character Thomas “Tommy” Shelby is portrayed by an Irish actor Cillian Murphy. Throughout the seasons, viewers watch how the family is working on expanding their business and influence not only in their hometown but also internationally.
Shelby Family Tree
The Shelbys family line begins with Mr. Shelby whose name is unknown and his wife Birdie Boswell. Birdie was a Gypsy Princess and a member of the largest and most powerful gypsy families in England at that time. Together with Mr. Shelby Birdie had two children, a son Arthur Shelby Sr. and a daughter Elizabeth “Polly” Gray (née Shelby). Take a look at the Shelby family tree created with the Treemily family tree chart maker.
Thomas (Tommy) Shelby
Thomas Michael Shelby was born in 1890 in Birmingham, England, and was a second child of Arthur Shelby Sr. He is the leader of the Peaky Blinder criminal gang, ahead of the Shelby family and Shelby Company Limited. During the First World War, Thomas received drastic experience while serving as a sapper and it has changed his life forever. As awful as it sounds, it was the war that gave rise to the Shelby family’s success and made Thomas a strong and fearless leader. His leadership qualities, as well as his strategic and commanding abilities, made him the head of the family instead of his elder brother Arthur who is too hot-headed and impulsive to make important decisions.
Grace Shelby (Late ex-wife)
Tommy’s wife Grace Helen Shelby (née Burgess) worked as an Irish barmaid when she met her future husband. Graсe was born in Galway, Ireland in 1894. Before marrying Thomas Shelby, she was married to an American banker Clive Macmillan. Grace and Tommy’s son Charles Shelby was born in 1922. Tommy Shelby also has a daughter named Ruby. Her mother, a former prostitute Lizzie Stark, was previously engaged to John Shelby but the engagement was called off.
Lizzie Shelby (Wife)
Elizabeth “Lizzie” Shelby (née Stark) is the second wife of Thomas Shelby. She was initially engaged to Thomas’ brother John, but the latter called off the wedding upon finding out that she wasn’t faithful to him. Soon afterwards Thomas made Lizzie his secretary and they developed a connection. Lizzie married Thomas, although they always had a very tense and complicated relationship – Lizzie often fell victim to abuse and unfair treatment by not only Thomas, but other Shelby family members. Towards the end of the show, not able to bear the abuse any longer, Lizzie left Thomas for good, cutting him out of her life.
Arthur Shelby Jr
Arthur William Shelby Jr. is the eldest son of Arthur Shelby Sr. born in 1887 and is a Deputy Vice President at Shelby Company Limited. Arthur is married to Linda Shelby (born in 1895). The couple has a son named Billy.
Linda Shelby (Wife)
Linda Shelby is the ex-wife of Arthur Shelby Jr.. As a devout Quaker, Linda made a lot of effort to change her husband for the better, often criticizing his illegal activities. Linda is highly intelligent, determined and confident, however, she also has her weaknesses – for instance, she is shown to be addicted to cocaine. Over the course of the show, their relationship aggravated more and more, which resulted in Linda trying to kill Arthur; however, the attempt failed. Soon thereafter, Linda divorced Arthur and left the Shelby family for good.
John Shelby
John Michael Shelby is the 3rd son of Arthur Shelby Sr. Also known as John Boy, he served as a soldier of the Warwickshire Yeomanry during the First World War alongside his brothers Arthur and Thomas. His first wife Martha Shelby died of unknown causes leaving John widowed with 4 kids to raise. His second wife Esme Martha Shelby (née Lee) is a member of the Lee gypsy family also from Birmingham. Together they have 3 kids.
Esme Shelby
Esme Martha Shelby (née Lee) is the wife of John Shelby. The marriage between her and John was supposed to be a sign of truce between the Shelby and the Lee families. Still, the two grew really close and eventually had 4 children. After John’s tragic death, Esme was absolutely devastated and made a decision to leave the Shelby family for good, taking all the children with her.
Finn Shelby
Finn Shelby is the youngest of the Shelby brothers born in 1908. Even though Finn is involved with Peaky Blinders and their business, his brothers always try to keep him away from any dangerous and illegal actions.
Ada Thorne (née Shelby)
Ada Thorne (née Shelby) is the only daughter of Arthur Shelby Sr. and his wife born in 1897. She is the only member of the family who is not involved with Shelby’s family business. Her husband Freddie Thorne is a communist and a former best friend of Thomas Shelby. Ada and Freddie have a son Karl Thorne named after Karl Marx.
Freddie Thorne (Husband)
Freddie Thorne is the husband of Ada Thorne, with whom he also has one son, Karl Thorne. Freddie is a street-smart, tough war veteran who spent his later years as a communist agitator (even his son was named after Karl Marx). At school, he was a close friend to Thomas Shelby, however, the two grew distant as years went by. Regardless, they both had respect for each other and may have grown closer again, but Freddie met an unfortunate end off-screen – he died during the Spanish Influenza pandemic.
The Gray Family
Polly Gray (née Shelby)
Elizabeth Pollyanna “Polly” Gray (née Shelby) is the matriarch of the Shelby Family and sister to Arthur Shelby Sr. She is also the mother of Michael and Anna Gray, and aunt of Thomas, Arthur, John, Finn and Ada Shelby. She is a highly intelligent woman who is both a great leader and a skilled accountant, taking charge of the Peaky Blinders while Thomas is away and being the treasurer of Shelby Company Limited.
Michael Gray
Michael Gray is the son of Polly Gray and Mr. Gray as well as the adopted son of Rosemary Johnson. When Michael was a young boy, his father died as a result of an accident involving excessive drinking; he was later taken away from his mother Polly and adopted by the Johnson family. When Michael first gets involved in the Peaky Blinders business, he’s shown to be very diligent and responsible, however, overtime his ambitions to take charge grow massively. This results in a confrontation with Thomas, leading to many deaths, including his own.
Gina Gray (Wife)
Gina Gray (née Nelson) is the wife of Michael Gray and the mother of Laurence Gray. Gina is a highly intelligent and manipulative character who always craves for more power and money. Throughout the later seasons of the show, Gina persistently pushes Michael to overthrow Thomas as the head of the Peaky Blinders, and the couple eventually plots the murder of the gang leader. The plot fails, however, with Michael getting shot in head, and Gina getting banished from the family for good.
Other Shelby Members
Arthur Shelby Sr
Arthur Shelby Sr. was married to a woman from the Strong family. The couple had five children together, four sons and one daughter. Arthur abandoned his kids after their mother’s death. All of the children were raised by Arthur’s sister Polly.
Uncle Charlie
Charles “Charlie” Strong (Uncle Charlie) is not a blood relative to the Shelby family, acting more like a father figure. He works for the Shelby Company Limited, often seen helping with stock and preparing shipments, as well as tending to horses.
Discover other famous people’s family trees both real and fiction by checking out the rest of the Treemily blog articles. Share them with your friends and family or order a printed version of a tree to make a great gift for your loved ones.
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8707
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dbpedia
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0
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https://musicbrainz.org/artist/cbb81bc6-fc8f-4d90-8a56-b5e0b357de3a
|
en
|
Michael Gray
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UK house DJ/producer, Type: Person, Gender: Male, Born: 1966-07-12 in Croydon, Area: United Kingdom
|
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|
/static/images/favicons/apple-touch-icon-57x57.png
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8707
|
dbpedia
|
2
| 63
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https://www.bchfh.com/memorials/michael-gray/2313708/obituary.php
|
en
|
Burpee, Carpenter & Hutchins Funeral Home
|
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ROCKLAND - Michael Sean Gray born June 17th 1976 unexpectedly passed away at his home at the age of 39. Michael had a long battle with Hodgkins Lymphoma. Michael is predeceased by his mother Alice Jeanette Gray and his father Calvin Albert Gray. He is survived by his sister Sheri A. Gray of Rockland and his brother John A. Gray of Warren, and many nieces, nephews, and cousins. Michael attended Thomaston schools and later managed his father’s business. Michael enjoyed hanging with his friends, taking road trips and spending time with his beloved girlfriend Jocelyn. Michael was diagnosed with lymphoma 6 years ago and fought so hard but stayed positive. We would like to give extended thanks to anyone that supported him. There is a memorial fund set up to aid in funeral costs if you would like to donate. The fund is set up through gofundme.com (Michael Gray Funeral Fund). A service will be held at a later date. To share a memory or condolence with his family, please visit his Book of Memories at www.bchfh.com. Arrangements are in the care of Burpee, Carpenter & Hutchins Funeral Home, 110 Limerock Street, Rockland.
|
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mike you will be sadly... (read more)
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8707
|
dbpedia
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3
| 78
|
https://www.espn.com/nfl/story/_/id/28066447/patriots-rb-jonas-gray-brush-fantasy-vs-reality-five-years-ago
|
en
|
Patriots RB Jonas Gray's brush with fantasy vs. reality, five years ago
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2019-11-13T13:00:00+00:00
|
Five years ago, an unknown Patriots running back named Jonas Gray put up a monster performance against the Colts. A few days later, an uncharged phone changed the course of his career.
|
en
|
ESPN.com
|
https://www.espn.com/nfl/story/_/id/28066447/patriots-rb-jonas-gray-brush-fantasy-vs-reality-five-years-ago
|
THE DAY JONAS GRAY destroyed your fantasy football team started out innocently enough, with a hot Epsom salts bath and breakfast at a forgettable downtown Indianapolis restaurant with his mom and brother. Had Gray known what would happen later that night, he might have tried to find his family better tickets for the Patriots-Colts game on Nov. 16, 2014. But Gray was an undrafted third-year running back living in a one-bedroom apartment outside of Boston. Nosebleed seats it was.
In the upper deck that night, Jerri Gray-Allen, a retired police officer who drove 300 miles from her Michigan home, tried to low-key it when her son scored his first touchdown. She clapped and high-fived and let out a small scream. But by the time Gray entered the end zone for touchdown No. 3, she had dropped to her knees. "My baby," she said to herself, "is finally getting his shot."
Up to that point, Gray's career had been defined by practice squads and wishful thinking. But that night in Indianapolis, the stars and, most important, Bill Belichick's game plan aligned. Gray erupted for 201 yards and four touchdowns, becoming the first player in the Super Bowl era to account for more than a quarter of the league's rushing scores in a week. Those four touchdowns were the first four of his career. That hadn't happened since 1921.
His underdog story, played out in front of millions of eyeballs on Sunday Night Football, was intoxicating and endearing. Gray jumped into the arms of Rob Gronkowski and playfully head-butted Tom Brady. After the game, Colts quarterback Andrew Luck found Gray and repeatedly, earnestly, told him how happy he was for him.
Gray has a degree in English from Notre Dame, but when NBC's Michele Tafoya put a microphone in front of his face after the game, he was so nervous he wasn't sure what would come out. He'd watched those interviews on his TV for years, wondering what it would be like to be the man -- the one who gets the game ball and has everyone riveted to what he has to say.
"I remember saying to myself, 'Wow, this is so cool,'" Gray says. "'I hope I don't mess up.'"
In five days, Gray's moment would be over, all because of a cellphone charger and one harsh reality of playing for the NFL's greatest dynasty: Game plans trump a good story every single day.
GRAY'S BREAKOUT EARNED him the cover of the Nov. 24 issue of Sports Illustrated, under a headline that read, "Jonas Gray ... Because of Course." Gray's star was already fading by the time the magazine hit newsstands, but there will always be permanence about that night.
If you're a hard-core fantasy football owner, Nov. 16 is eternally known as Jonas Gray Day, the anniversary of one of the most seismic shifts in a fantasy football weekend. Gray earned 43 points that night, the second-highest-scoring game by any player in 2014. Two Boston-area brothers, Rob and Dave Gomes, started Gray that weekend and won $1 million in a DraftKings contest. (Gray's 2014 salary, by the way, netted him less than half of that amount.)
Gray was such an unknown that he was active in just 1.3% of ESPN leagues that week, and, according to ESPN senior fantasy writer Tristan H. Cockcroft, was still available to be picked up, essentially for free, in 91.6% of them.
"When we celebrate national one-hit-wonder day, my timeline on Twitter gets flooded with pictures of Jonas Gray," says ESPN NFL insider Field Yates. "Patriots running backs are always, for fantasy football purposes, tantalizing. There's always going to be value there, but it's unpredictable because they've had such a cast of playmakers come through."
That unpredictability was on full display the following weekend: By then, Gray appeared in 75.7% of ESPN's fantasy leagues and was started in 32.9%.
He didn't play a snap.
EVEN BEFORE THE Colts game that made him one of Belichick's most famous one-hit wonders, Gray was well-versed on the vagaries of football. He had been a four-star recruit out of Detroit Country Day High School but found himself buried on the depth chart at Notre Dame, with just 75 carries in his first three seasons.
Back then, Gray leaned on his sense of humor to keep him going. He did some stand-up comedy in the South Bend area, once opening for Dustin Diamond, who played Screech on "Saved by the Bell." "If you ask anyone there," says former Irish teammate Mike Golic Jr., "Jonas was funnier than Dustin Diamond."
Despite the setbacks, Gray believed he could play with anyone. He cracked the starting lineup in his senior year and exploded for 791 yards and 12 touchdowns on 114 carries. His hopes of being drafted in the NFL were in sight. Then on Senior Day, he tore his ACL, MCL and LCL. His knee, and his dream, crumpled.
Gray stood on crutches in the locker room after the game, and his teammates gathered around him, awkwardly struggling with what to say. But Gray didn't want their sympathy. He recited a line from one of his favorite books, The Count of Monte Cristo: Life is a storm, my young friend. You will bask in the sunlight one moment, be shattered on the rocks the next. What makes you a man is what you do when that storm comes.
He was eventually signed by the Dolphins, then spent the 2013 season on the Baltimore Ravens' practice squad. He joined the Patriots on a futures contract in January 2014. Though he missed out on the 53-man roster heading into the season, Gray, who stood 5-foot-10 with a walloping 230-pound frame, impressed in training camp. Belichick, in those first weeks of the season, told him, "You're close." In mid-October, after a knee injury sidelined Stevan Ridley, Gray was promoted and amassed 131 yards on 32 carries in his first three games.
Nobody could've predicted that Gray would get 37 carries against the Colts. But Belichick is a master at modifying his personnel to each opponent, plugging in no-name players with particular skill sets that expose every weakness. His practice squads are fluid, and they weigh heavily into this game of chess. "They can move you up at any moment," says former Patriots guard Chris Barker, who spent six rotations on New England's practice squad. "They can literally call you the Friday night before the [team] flight and say, 'All right, you're activated.' You've got to be ready to play."
The Indianapolis game came after a bye week, which gave Belichick and offensive coordinator Josh McDaniels extra time to prepare. And fresh in their minds was the Colts-Pats matchup in the AFC divisional round 10 months earlier. In that game, a 43-22 Pats win, another bruising back named LeGarrette Blount had pounded his way for 166 yards and four touchdowns. (Blount, by the way, would have just five carries the next week in the AFC championship.)
Belichick had also noticed a weak spot in the Colts' defense, with run-stopper Arthur Jones out that week with an injury. So they stacked their offense with six linemen, two tight ends and a plan: Feed the 24-year-old Gray constantly. In the days before the game, Patriots owner Robert Kraft stopped Gray in the locker room and offered encouragement. "I think this is going to be a big week for you," Gray recalls Kraft saying.
All those years of waiting, and now Gray had a wall of blockers and the ball. It was his show. He ran up the middle and off left tackle. He scored his first touchdown with 8:37 remaining in the first quarter, bulldozing into the end zone before some of the Lucas Oil Stadium crowd could even settle into their seats.
"I was like a kid in a candy store," Gray says. "It was like nothing I could describe. To get to that point, the dedication it takes, the hard work, it's hard to even describe it to people. I mean, it's a blast."
After the game, a reporter in the locker room snapped a picture of Brady shaking Gray's hand and sent it to Gray's mother.
In that moment, Gray felt something he hadn't for years. He felt as if he belonged.
GRAY'S AGENT, Sean Stellato, is based in New England and has represented numerous Patriots over the years. He knows the Patriot Way, and the expendability implicit in that philosophy, but after watching that Colts game, Stellato was certain that Gray was going to be a star.
"When you have a client that has a huge game," he says, "it's like, 'Wow, this is the break he needed. This is the breakout.' But like I tell all my clients, you've got to stay paranoid. You obviously don't ever want to take the pedal off the gas when you have success. It can come and go, as we know, in the snap of a finger."
On the Patriots' plane ride back from Indianapolis, Gray thought of none of that. He could barely sleep, already thinking ahead -- if he could finish strong in the last six games, maybe he could make the Pro Bowl. Sure, it was premature, but anything felt possible.
The next day, LeGarrette Blount, by then with the Steelers, left the field during a Monday Night Football game at Tennessee after receiving zero carries. He was subsequently released, and the Patriots picked him up on Nov. 20, a Thursday. Blount was a proven veteran, and his addition seemingly made Gray redundant on the Patriots' roster. But Gray wasn't particularly worried. He figured that he could learn from Blount, that Blount would only help make him better.
The Patriots were playing Detroit that weekend, a team Gray had followed since he was a kid growing up in Michigan. Thursday night, Gray stayed up late watching film of the Lions on his cellphone and iPad, lying on his couch with ice packs on his legs.
He plugged his phone into a charger and fell asleep on the couch. Gray was too tired to notice that the charger was dangling precariously out of the wall socket.
ONE OF THE first things you learn in the House of Belichick is to never, ever be late. A tardy is the same thing as an absence. Bryan Stork, a rookie center for the New England Patriots back in 2014, once contemplated buying a snowplow blade just in case he got caught in a nor'easter on his way to a team meeting, but he eventually decided it would have been over the top.
The Patriots had an early team meeting on Friday, Nov. 21, and like always, Gray had set the alarm on his cellphone the night before. It never went off. He awoke to the sun, glanced at the kitchen clock and saw, to his horror, that it was 8:30 -- one hour after the meeting had started. He scrambled for his phone, which was dead. In the agonizing minutes it took to charge the battery, Gray was awash in panic. When the phone finally turned on, he saw a text from veteran nose tackle Vince Wilfork. "Are you OK?"
Kevin Anderson, who was the Patriots' football operations manager at the time, texted and stopped by to check on Gray. (The Patriots' fear, according to Gray, was that the instant success might've made him vulnerable -- maybe he was celebrating and had too much to drink. Gray says Anderson made note that he had not been drinking.) Anderson told him to stay home until he called him later. But the waiting was killing Gray.
He texted teammates, family and former coaches. He apologized for his mistake. Sometime around 5:30 p.m., Gray showed up at the Patriots' facility to talk to Belichick. He wanted to explain everything -- that he did his job but the phone charger didn't.
The coach didn't appear particularly angry, Gray says. Belichick was on a treadmill walking and reading notes. He repeatedly told him, "We just can't have it." He said there would be repercussions for the game against Detroit, though he didn't specify how much Gray would sit. (Belichick, through a Patriots spokesperson, declined to comment for this story.)
Two days later, Gray didn't play a single snap against his hometown Lions. When Belichick was asked in the postgame why Gray didn't play, he told reporters, "We do what we think's best, and that's what we did today."
At least outwardly, Belichick didn't blame Gray's absence on the missed meeting, or on the tweet that Gray sent the day before the game -- a comment about "how fast people can turn their back on you."
Gray had quickly deleted the tweet, which also mentioned that he would "keep it moving and grind harder." And it seemed as if Gray did.
"Whenever I saw him around, he was positive and happy," Chris Barker says. "I've never seen him mad about his opportunity to play."
But Gray believes that late wake-up forever changed his trajectory with the team. In the next game, against the Packers on Nov. 30, the Patriots used four running backs for 17 carries. Blount received 10 of them. Gray had one.
For the rest of the season, Gray recorded double-digit attempts only once -- 11, for 62 yards, against the Dolphins. In all, he had just 91 yards after his monster game against the Colts. He was a healthy scratch for Super Bowl XLIX.
EVEN THEN, GRAY believed he'd get another chance in 2015. He played well in the first game of the preseason, ripping off a 55-yard touchdown run against the Green Bay Packers. He spiked the ball and flexed his arms like he had in that extraordinary November game.
A few weeks later, Belichick and Nick Caserio, the director of player personnel, called him in for a meeting. Belichick told him they were going in a different direction, that they didn't need any bigger backs in 2015. He said it didn't diminish the role Gray played in their Super Bowl championship. "Still, to this day, it gets to me," Gray says, fighting back tears.
"I was pretty shocked."
He went to Miami but got caught in a numbers crunch. He had high hopes after signing with Jacksonville in December, but then tore his quad during training camp in 2016.
Now Gray is 29 years old and stuck in an existential limbo. Is he still a football player? Or a guy who continually works out because he can't let go?
Gray has three kids now and is back in the Boston area working for an energy company. He spends what little free time he has working out, and he says he's ready if football calls. He takes inspiration from his mom, who struggled to raise two boys alone before deciding, at 28, to enroll at a community college. Jerri Gray-Allen didn't abandon her dreams, and she went on to be a crime scene investigator with the Pontiac Police Department -- and she says she'd never tell her son to quit football.
"He loves the game. That's what he wants to do.
"Of course I was disappointed in the system," she says. "But I was never disappointed in Jonas. I know what type of person he is, and I know what type of player he is. I'm not just saying that because he's my son."
Because he hasn't played in a few years, Gray considers his football age to be closer to 25 than 29. He is filled with confidence. He watches the NFL and believes he's as good as, if not better than, some of the running backs who have opportunities he doesn't.
He wonders where he'd be right now if his phone had been charged on that November day.
"I don't think I'd still be with the Patriots," he says. "But I'd definitely be in the NFL. I probably would be somewhere with a large contract playing on a team. I probably would've left New England because they couldn't pay me."
EVERY NOVEMBER, THE memories rush back to that game in Indianapolis. He can't believe it's been five years. Gray says that he carries no ill will toward Belichick and that he "respects the hell out of him."
Gray attended an XFL summer showcase in July, and Stellato said his client is in the best shape of his life. But Gray wasn't selected in last month's draft. An XFL player personnel staffer, who spoke under the condition of anonymity, said that teams simply preferred younger running backs in the draft, though Gray could still be added to a roster before the league launches in 2020.
In the meantime, Gray waits for his chance. Every once in a while, he'll think of something funny and write it down for a comedy skit maybe somewhere down the road. But a comeback? That's no laughing matter to Gray. If an unknown kid can blow up the NFL, and fantasy football, on one November weekend, anything is possible.
"I think it's gonna happen," he says. "I really do. My story is still being written."
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8707
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dbpedia
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https://forum.ibiza-spotlight.com/threads/english-dj-looking-for-gigs-2005-season.15531/
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English DJ looking for gigs 2005 season
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"I ibiza_lou New Member",
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2005-02-15T11:27:25+01:00
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I know it is always best to get there to do this kind of thing but I will post up my details as well.
I am an English DJ currently working on the Costa...
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en
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Ibiza Spotlight forums
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https://forum.ibiza-spotlight.com/threads/english-dj-looking-for-gigs-2005-season.15531/
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I know it is always best to get there to do this kind of thing but I will post up my details as well.
I am an English DJ currently working on the Costa del Sol and am trying to get out to either Majorca or Ibiza this year. I play a variety of music, with my best selection and knowledge being from 90s to present house, commercial dance and RnB. I am a cd only DJ, lots of experience and I am also now starting to produce my own music. A full CV and mix cds can be provided, if you require any further info contact me direct on info@dj-dc.com or go to the website www.dj-dc.com. Below is my latest chart so people can get a feel for my music style
Artist, track, mix, label
1 Sunset Strippers Star To Fall (Unknown) White Label
2 Studio B I Ssee Girls (Tom Neville) Boss Records
3 Solitaire You Got The Love (Ext Club Mix) Su Su
4 Thomas/Falcon High Again (A1) Oxyd
5 Superchumbo Dirty Filthy (Orig) TWISTED
6 Deux Sun Rising Up (Vocal Mix) Urbana
7 Salif Keita vs Junior Jack Madam Samba (Unknown) White Label
8 LnM Project ft Bonnie Bailey Everywhere (Orig) Hed Kandi
9 Workidz Disco Sound (Orig) Secret service
10 Kings Of Tomorrow Thru (Junior Jack Mix) Defected
11 Lovefreekz Shine (Club Mix) Positiva
12 Armand Van Helden MyMyMy (DC Edit) Southern Fried
13 Dylan Rhymes Salty (Orig) KINGSIZE
14 Jupiter Ace 1000 Years (Orig) Oxyd
15 Techtonic Funk 980 (Orig) Gabor szanto
16 Michael Gray The Weekend (Extended Vocal) Eye Industries/Island
17 Workidz Fantastic Love (Orig) Disco Galaxy
18 EKA Something More (Mendes Full House Mix) Ekow Earmah
19 General Moders Cross The Sky (Extended Mix) SIZE RECORDS
20 Seamus Haji A DJ Saved My Life (ATFC Mix) Big love
Thank you
DC
too many dj's are trying to play out in ibiza that the bars/clubs take advantage of you guys by letting you play and NOT paying you... and they don't care if you walk away cause there's always someone else waiting to take ur spot...
so if i were you i'd try and get a paid DJing job sorted before u make a permanent move for the season otherwise u'll find urself in trouble...
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https://www.detroitchamber.com/bios/michael-gray/
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Michael Gray
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[
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2023-01-04T15:34:34+00:00
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Michael Gray serves as the Director of Operations and partner of Four Man Ladder, overseeing and collaborating with the leadership teams of Grey Ghost, Second Best, and, most recently, Basan, located on the ground floor of the recently renovated Eddystone in the District Detroit.
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Detroit Regional Chamber
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https://www.detroitchamber.com/bios/michael-gray/
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Michael Gray serves as the Director of Operations and partner of Four Man Ladder, overseeing and collaborating with the leadership teams of Grey Ghost, Second Best, and, most recently, Basan, located on the ground floor of the recently renovated Eddystone in the District Detroit.
Gray is a metro-Detroit native with more than 25 years of hospitality leadership experience, emphasizing exceptional guest experiences, business development, mentorship, and healthy workplace culture. He has opened 16 restaurants and bars in the Midwest throughout his career, highlighted by San Morello and Evening Bar in the Shinola Hotel.
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https://www.geol.umd.edu/~tholtz/G104/lectures/104thyreo.html
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GEOL 104 Thyreophora: Defense! Defense! Defense!
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Key Points:
•Among the primitive ornithischians are big-handed, deep-skulled Heterodontosauridae and slender Eocursor; the rest form the clade Genasauria
•Thyreophorans represent the armored dinosaurs, and are a clade of (predominantly) quadrupedal ornithischians.
•There are characterized by the presence of osteoderms (armor plates) in their skin. Different clades of thyreophorans express these osteoderms in different patterns.
•Beyond a few basal taxa, thyreophorans are divided into the plated Stegosauria and the tank-like Ankylosauria.
•Armor in thyreophorans seem to have functions beyond simple defense: they served as display structures and (in the case of the stegosaurs and the club-tailed ankylosaurine ankylosaurs) as active weapons.
As we saw in the last lecture, the only likely Triassic prionodontian is Pisanosaurus of the early Late Triassic Argentine Ischigualasto Formation. The fossil is incomplete, so many aspects of its anatomy are uncertain.
As previously discussed, Pisanosaurus plus Prionodontia is characterized by the following traits:
The predentary bone: a single midline bone joining the left and right dentaries:
Covered in life with a horny beak
Allowed slight rotation of dentaries to help them chop up food
Similar structures known in a few other animals, including paired edentulous (toothless) anterior dentary ends in silesaurids
Broad phyllodont dentition (that is, broad leaf-shaped teeth with large denticles)
Five or more sacral vertebrae
Inset tooth row and ridges along the maxilla and dentary suggesting that ornithischians had a muscular or skin cheek.
If so, this would help keep food in the mouth while munching. However, this hypothesis has its detractors, and some suggest the alternative of a large horny surface along the sides of the mouth instead.)
The latest and most comprehensive work shows that a large muscle from the temporal region to the dentary occupied this space, associated with chewing ability. Thus, this wasn't a cheek, but also there was some "meat" in this area.
Maxillary fenestra is reduced (and in many forms lost altogether)
Prionodontia is characterized by:
Opisthopuby (backwards-pointing pubis) (the "bird-like" aspect of their hips that gives this group its name)
Allows for greater gut capacity for digesting plants without making the body too wide
Backwards pubis evolves independently among different advanced saurischian groups, as we will see later
Epaxial (above the vertebrae) ossified (turned to bone) tendons, stiffening the back and (perhaps) acting as a "spring" to absorb and release energy while running
Primitive Ornithischian Groups: Heterodontosaurids & Eocursor:
Heterodontosauridae was a primitive group of ornithischians. Although one fragmentary specimen was thought to be from the Late Triassic, redating shows it was younger; the oldest heterodontosaurids are known from Early Jurassic, and persist into the mid-Early Cretaceous. They had skulls which are relatively deep and powerfully built, indicating that they ate fairly tough food. Advanced heterodontosaurids also had premaxillary tooth row that were ventral to the maxillary tooth row and jaw joints that were ventral to the dentary tooth row: the result were jaws that brought the teeth together all at once (like a nutcracker), and not slicing (as in scissors, or as in most dinosaurs). These latter jaw adaptations evolved convergently in Ornithopoda, and so for a long time Heterodontosauridae was considered a clade within Ornithopoda. However, primitive heterodontosaurs lack these convergent adaptations.
Most heterodontosaurids are quite small. Some are only about 1-1.5 m long, and Fruitadens of the Late Jurassic of western North America may have been no more than 80 cm long as an adult (most of which length is tail) and Manidens of the Middle Jurassic of Argentina only 65-75 cm; that makes these the smallest known ornithischians.
Interestingly, the early Late Jurassic Chinese heterodontosaurid Tianyulong had a fuzzy body covering over at least part of its body! If this is found to be homologous to the protofeathers of tetanurine theropod saurischians it would suggest that the concestor of all dinosaurs was fuzzy, and that dinosaurs were thus fuzzy ancestrally! At present, however, there is enough uncertainty to make the homology between Tianyulong's fuzz and tetanurine protofeathers suspicious. (But do not be terribly surprised if in the future we discover that most dinosaurs were fuzzy to some degree or another! All we need is a fuzzy primitive sauropodomorph, and it is basically a done deal!) (By the way, the initial reports placed Tianyulong in the Early Cretaceous, but the formation in which it was found has been redated to the earliest part of the Late Jurassic, around 160 Ma.) (A few years ago a radical new hypothesis for the position of Heterodontosauridae was proposed: that "heterodontosaurids" were a paraphyletic grade of Jurassic and Early Cretaceous pachycephalosaurs! However, the latest most comprehensive studies have not supported this hypothesis.)
The similarly aged neornithischian Kulindadromeus of Siberia also shows simple filimentous fuzz, as well as scales, plates, and additional bizarre tufted plates, showing that primitive ornithischians had a wide variety of integumental features.
Once thought to be from the Late Triassic, Eocursor of South Africa is from the earliest Early Jurassic.
Ornithischians more derived than Heterodontosauridae, Eocursor, and Laquintasaura had greatly reduced hands, losing most of their grasping ability. This suggests a switch to jaws-only while obtaining food.
The remaining ornithischians (Genasauria, the "cheeked reptiles") include the armored Thyreophora and the highly diverse Neornithischia (especially beaked Ornithopoda and ridge-headed Marginocephalia).
Simplified cladogram of Thyreophora
More detailed phylogeny of Genasauria, with emphasis on Thyreophora
MAJOR GROUPS OF THYREOPHORANS
Thyreophora (longshield bearers)
Oldest definite specimens from the Early Jurassic (some problematic specimens from Late Triassic), persist until very end of the Cretaceous
Global distribution (all continents)
Known from diverse environments
Thyreophora are united by various skeletal attributes, the most obvious of which is:
Osteoderms: bony armor in the skin, forming scutes covered by keratin
The derived thyreophorans are the plated Stegosauria (shingled lizards) and the heavily armored Ankylosauria (fused lizards). Both clades are present by the Middle Jurassic. There are several Early Jurassic taxa which fall outside either of the two advanced clades.
BASAL THYREOPHORANS
Lesothosaurus of the Early Jurassic of southern Africa and similarly-aged Laquintasaura appear to be the oldest and earliest branching thryeopohiorans. Laquintasaura is the oldest and most primitive ornithischian known from group assemblages, indicating that at least some of these lived in groups during life. These are united with the later thyreophorans on various subtled cranial and skeletal traits that are outside the scope of this course. Importantly, these are the only known unarmored thyreophorans. (But we would expect the oldest ones to have been unarmored.) To be fair, other analyses find one or both to be the oldest and most primitive neornithischian(s) or in their traditional positions as non-genasaurian ornithischians.
The oldest and most primitive definite (i.e., armored) thyreophoran is Scutellosaurus of the Early Jurassic of western North America. It was a 1.5 m long biped (possibly facultative biped) not very dissimilar to other primitive ornithischians like Lesothosaurus or Hexinlusaurus: small herbivores with small hands. The primary distinction of Scutellosaurus is the presence of a great many small osteoderms over the body. These would protect against small-bodied predators, but might not help against the new larger theropods that had begun to appear in the Early Jurassic.
In response, thyreophorans evolved larger size and heavier armor, as seen in Emausaurus and Scelidosaurus (new specimen shown here), both of Europe, and comparable-aged Yuxisaurus of China. The larger body size (3-4 m long) and proportionately larger osteoderms may have been more effective defense against attacking predators, but forced them onto all fours (at least for Scelidosaurus): in other words, they were obligate quadrupeds.
(Note that the hypothesis shown here is that Scutellosaurus, Emausaurus, and Scelidosaurus were progressive closer to the Stegosauria-Ankylosauria clade (Eurypoda). However, some paleontologists have considered Emausaurus to be a primitive stegosaur, and others that Scelidosaurus was the oldest and most primitive ankylosaur. However, eurypods share a number of transformation not found in these Early Jurassic taxa: these include:
Metacarpi and metatarsi connected only at the proximal and and spreading out distally (making for broad hands and feet, as seen in these ankylosaur tracks)
Femur shaft is straight
Part of the ilium anterior to the ilium is longer than the part posterior, and fares out from the midline
Spike osteoderms protruding from the scapular region
Reduction of the antorbital and infratemporal fenestrae
among others.)
A possible fly in the ointment of this scenario is recently (2022) described Jakapil of the early Late Cretaceous of Argentina. Known only from a single partial skeleton, this seems to be a late survivor of the non-eurypodan thyreophorans, even closer to Eurypoda than Scelidosaurus. However, the remains of the forelimb and hindlimb indicate it was almost certainly a biped! Obviously we need more of the skeleton to make certain, but Jakapil is a good reminder that Evolution is not a planned directional phenomena, and that some lineages will buck the trend that the rest of their clade are following if that is the most successful set of adaptations at that place and time. Also, there is the definite non-zero chance that Jakapil belongs to some entirely non-thyreophoran group that convergently evolved osteoderms. Indeed, its mandible is very much like that of a neornithischian, so perhaps it is the first discovery of a previously-unknown clade of Gondwanan marginocephalian!
STEGOSAURIA
From a Scelidosaurus-like ancestor, the stegosaurs evolved armor that was less covering and more concentrated. While they had some small osteoderms in their skin (particularly around the neck and the hips), most of their armor was specialized as:
Enormous spikes protruding from the scapular region (present in ankylosaurs, but proportionately much larger in primitive stegosaurs)
A pair of (initially symmetrical) parallel rows of osteoderms along the back, comprised of some mixture of:
Plates: mediolaterally flattened tall osteoderms Spikes: conical tall osteoderms
The thagomizer: two (or more?) pairs of spikes at the tail, used as a defensive weapon. (NOTE: named after a Far Side cartoon.) Unlike the traditional position of museum mounts and drawings, the spikes of the actual thagomizer were almost certainly laterally directed.
In general, the stegosaur armament suggests active defense: the dinosaur probably turned in response to attacking predators, trying to keep the tail towards the theropod so that it could use its thagomizer. Damaged thagomizer spikes and theropod bones with thagomizer-generated puncture wounds confirms their use in defense.
Early stegosaurs were only about 2.5-3 m long, but the most derived forms ranged up to 9 m or more. Their narrow snouts suggests that they were rather picky eaters (that is, instead of munching a lot of plants at once, they were selective as to which ones they chomped.) Biomechanical analysis shows that their bite was stronger than many herbivorous saurischians, but still weaker than many specialized ornithischians. Although they were obligate quadrupeds in terms of locomotion, they may have been able to rear on their hind legs in order to feed higher in trees.
Some tantalizing footprints suggest possible Early Jurassic stegosaurs, but these may be from a more basal Scelidosaurus-like thyreophoran instead. Primitive stegosaurs include Middle Jurassic Isaberrysaura of Argentina (first reported as being an ornithopod, and being Early Jurassic) and Bashanosaurus and Huayangosaurus of China; and Late Jurassic Chungkingosaurus, Gigantspinosaurus and Tuojiangosaurus (all three from China), western North American , Alcovasaurus [figures A, B, E & F], and Early Jurassic Paranthdon of South Africa. More derived stegosaurs form the clade Stegosauridae, and include Middle Jurassic Adratiklit of Morocco, long-necked Late Jurassic Dacentrurus (small specimens of which were once considered their own genus, Miragaia) of Europe, Kentrosaurus of eastern Africa, and the Stegosaurinae.
Stegosaurids had dorsoventrally stretched neural arches and disproportionately short forelimbs, giving them an odd profile.
The most derived stegosaurids (Stegosaurinae) lacked shoulder spines (also missing in Tuojiangosaurus), had only plates rather than spikes along the back (again, shared with Tuojiangosaurus), and had alternating rather than parallel plates. This advanced group includes Middle Jurassic Loricatosaurus of Europe, Late Jurassic Jiangjunosaurus of China, the two Late Jurassic western North American genera (Hesperosaurus and famous Stegosaurus [also known from Portugal]), and Early Cretaceous Wuerhosaurus of China, the last of the [distinctive] stegosaurs. (Some stegosaur fragments are known from Europe about the same age as Wuerhosaurus, but are not distinctive enough to place within the stegosaur phylogeny). (Note: some paleontologists consider Wuerhosaurus, and Hesperosaurus to belong within the genus Stegosaurus.)
Stegosaurs are relatively common in Middle and Late Jurassic formations (especially so in China), are present but rare in some Early Cretaceous Asian, African, and European assemblages, and vanish before the end of the Early Cretaceous (at present, Mongolostegus of Mongolia is the youngest known). Claims of later stegosaurs have so far turned out to be either mis-dated or misidentified.
ANKYLOSAURIA
While the stegosaurs evolved active defense, the ankylosaurs (at least at first) seem to have been selected for passive defense: the ability to stay put and absorb damage from an attack. They were even more extensively armored than Scelidosaurus, and were characterized by:
Osteoderms fused to the skull roof Rings of osteoderms over the neck and shoulders Extensive osteoderms (some large) over the rest of the dorsal surface and in some cases other parts, including the belly, the limbs, the cheeks, and even the eyelids!
Unusual for dinosaurs, the trunk of ankylosaurs were typically wider across mediolaterally than tall dorsoventrally
Additionally, in ankylosaurs the predentary bone is reduced, and the jaws arranged so that they would have more extreme rotation (and also be pulled back further) when chewing then in most ornithischians. This evolved convergently (and to a far more extreme form) in the advanced ornithopods. This motion allowed ankylosaurs to more effectively chew up their food.
The oldest ankylosaurs are from the Middle Jurassic: Tianchiasaurus of China, Sarcolestes of Britain, and newly discovered Spicomellus of Morocco. None of these are known from anything approaching a complete skeleton, and it is not inconceivable that one or more of these are actually more basal thyreophorans (in the Scelidosaurus-part of the tree). Alternatively, as mentioned previously, some analyses place Early Jurassic Scelidosaurus as the basalmost ankylosaur rather than a non-eurypod thyreophoran.
Ankylosaur systematics is at present quite complicated. Traditionally Ankylosauria was divided into the club-tailed Ankylosauridae and the club-less Nodosauridae. Almost all recent studies, however, show that the "nodosaurids" are a paraphyletic grade comprised of several subgroups. The problem is that the different analyses don't agree what genera belong to which subgroup, nor do they agree on the arrangement of these subgroups relative to each other or to Ankylosauridae. An attempt at consensus is shown here, but this particular arrangement doesn't confirm to any one particular study and will almost assuredly change with future studies.
Late Jurassic North American Mymoorapelta and Gargoyleosaurus have uncertain positions: possibly basal branches of Ankylosauria, or basal ankylosaurids, or basal polacanthids, or some combination thereof.
In late 2021, there was the announcement of a relatively complete skeleton of a Late Cretaceous ankylosaur from southern Chile, given the name Stegouros. This taxon showed traits which united it with Early Cretaceous Kunbarasaurus of Australia and poorly-known Late Cretaceous Antarctopelta of Antarctica and Patagopelta of Argentina formed a clade Parankylosauria, found to be the sister group to Euankylosauria (the remaining "nodosaurids" plus Ankylosauridae). Stegouros at least has an interesting slashing tail weapon: an expanded series of laterally-oriented blade-like osteoderms.
Polacanthidae includes Early Cretaceous Gastonia of North America and Polacanthus of Europe, and possibly Hylaeosaurus (the first discovered thyreophoran). A number of other Early and Late Cretaceous ankylosaurs appear to be polacanthids, including gigantic early Late Cretaceous Peloroplites and smaller mid-Late Cretaceous Niobrarasaurus, both of North America. They tend to have extremely large blade-like osteoderms in their shoulder regions and a shield over the sacral region formed by a series of large osteoderms surrounded by many smaller ones.
The North American Early Cretaceous Sauropelta and poorly known Tatankacephalus show some similarities to polacanthids, but also to the other "nodosaurid" clades.
The Struthiosauridae are a radiation of primarily European forms such as Europelta of the Early Cretaceous, Pawpawsaurus and Stegopelta of the Early-Late Cretaceous boundary (the only North American struthiosaurids), and Struthiosaurus and Hungarosaurus of the Late Cretaceous.
Struthiosaurids may form a clade with Panoplosauridae, a group of primarily North American Late Cretaceous forms. Edmontonia, Denversaurus, and Panoplosaurus are the best known examples. These have broader, smoother skulls than most "nodosaurs", and often have huge laterally-oriented spines along their sides in the shoulder region. Some of these are among the largest ankylosaurs. Early Late Cretaceous Nodosaurus itself may belong to this group (if so, then either "Panoplosauridae" or "Panopolosauridae + Struthiosauridae" would then properly be "Nodosauridae"!)
There remain, however, a number of "nodosaur"-grade ankylosaurids which are quite difficult to place. Early Late Cretaceous North American Animantarx and Cedarpelta do not have stable positions in these studies. Tiny Liaoningosaurus of Early Cretaceous China is known only by probable juveniles and only in lake deposits: some have speculated it was a semiaquatic fish eater!
ANKYLOSAURIDAE
The remaining ankylosaurs do seem to form the clade Ankylosauridae; however, that group is not limited to the club-tailed forms of the Late Cretaceous as once thought. Among the most primitive known member of Ankylosauridae is spectacularly-preserved Borealopelta. There is a newly discovered clade of shield-hipped ankylosaurids (in which the shield is formed of fused osteoderms of all the same size) which includes European Early Cretaceous Vectipelta and Early/Late Cretaceous Dongyangopelta and early Late Cretaceous Zhejiangosaurus of China.
The remaining ankylosaurids have distally-stiffened tails (although to be fair, complete tails are not recovered in most of the earlier forms.)
The best known part of Ankylosauridae is the more derived clade Ankylosaurinae there are extremely complex air chambers in the skull (convergently evolved with Gastonia)
Additionally, ankylosaurines are further transformed relative to other armored dinosaurs in having:
Small triangular horns on the rear of the skull
A distally interlocking rigid tail ending in
a powerful tail club
Thus, ankylosaurines had a more active defense (in the form of the tail club) than other ankylosaurs (other than the slashing tail of Stegouros). While these assuredly were used against attacking predators, like most animal weapons they were also used against other individuals of the same species. In fact, well-preserved ankylosaur specimens show impact wounds generated by tail club strikes.
Ankylosaurids ranged from about 3 m to 8-10 m (the latter size includes Tarchia of Mongolia and gigantic Ankylosaurus of western North America). Pinacosaurus of Asia is one of the best known primitive ankylosaurines. Other Asian forms from the Late Cretaceous include Zaraapelta, Saichania, and Talarurus. Euoplocephalus, Dyoplosaurus, Akainacephalus, Zuul, Ziapelta, Scolosaurus and Anodontosaurus of the Late Cretaceous of western North America are the best studied.
Ankylosaurines are only known from the late Early Cretaceous of Asia and the Late Cretaceous of Asia and western North America at present; similar distributions are known for various other dinosaurs, as we shall see.
Ankylosaurids are more than their armor! Recent work has demonstrated that the bones which control the tongue and throat are incredibly well-developed in these dinosaurs: far more so than all other dinosaurs (outside of some birds.) They may have had long, powerful tongues: possibly for insect eating, possibly for grabbing plants, and possible for both and more. However, even more recent discoveries show that these bones also supported a powerful larynx (voice box): this suggests that ankylosaurids (and perhaps other dinosaurs) might have had strong vocal communications.
EVOLUTIONARY PATTERNS IN THYREOPHORA
Defense:
Probably the most conspicuous aspect of their evolution. Thyreophorans begin with a set of small scutes; develop larger scutes at the expense of bipedalism (and speed); then split between specialists in active (Stegosauria) vs. passive (Ankylosauria) defenses. Within the ankylosaurs, though, Parankylosauria and Ankylosaurinae independently evolve an active defensive tail weapon.
Relative success
Stegosaurs and ankylosaurs, as sister taxa, have their origins at the same time. However, stegosaurs flourish first (in the Middle and Late Jurassic), only to peter out during the Early Cretaceous and disappear before its end. Ankylosaurs are typically rare in the Jurassic (although at one location Gargoyleosaurus is very common), but become abundant in the Cretaceous.
Feeding adaptations:
Narrow-snouted basal thyreophorans and stegosaurs differ from broader-snouted ankylosaurs. The latter have a greatly reduced predentary bone, which may have allowed for more complex motion of the dentaries (for additional munching power) while feeding. The powerful tongue of ankylosaurids hint at some specialized form of feeding.
Group behavior:
Thyreophorans are only rarely found in mass death assemblages, and so (unlike some sauropods, ornithopods, and marginocephalians) probably did not live in large groups. The presence of tail-club generated wounds on ankylosaurids, and the existence of lateral shoulder spines on many non-ankylosaurid ankylosaurs, strongly suggests within-species competition (i.e., dueling matches.)
Display structures:
The spikes, plates, and osteoderms of thyreophorans almost certainly had a defensive function. But many are very broad (particularly stegosaur plates), and the patterns tend to be specific to each species. They may have served an additional function: as visual displays of species recognition. There may also have been a sexual display function to them, but at present it is uncertain if the variation we see in some thyreophorans is from sexual dimorphism or if it is from multiple species of the same genus living together.
Some relevant videos: To Next Lecture.
To Previous Lecture.
To Lecture Schedule.
Last modified: 2 July 2024
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https://www.ubuy.ci/en/product/54CIBCX0M-vitae-zhejiangosaurus-dinosaur-model-toy-collectable-art-figure
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ZHEJIANGOSAURUS Dinosaur Collectable Art Figure Toy Cote dIvoire
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Discover the collectable art figure of Zhejiangosaurus, an extinct nodosaurid dinosaur. Shop now at Ubuy Cote dIvoire for a unique addition to your collection.
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en
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Ubuy Cote dIvoire
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https://www.ubuy.ci/en/product/54CIBCX0M-vitae-zhejiangosaurus-dinosaur-model-toy-collectable-art-figure
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622
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https://en.wikipedia.org/wiki/Nodosauridae
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Nodosauridae
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2006-03-05T14:00:31+00:00
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https://en.wikipedia.org/wiki/Nodosauridae
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Extinct family of armored dinosaurs
Nodosaurids
Gargoyleosaurus skeleton cast Scientific classification Domain: Eukaryota Kingdom: Animalia Phylum: Chordata Clade: Dinosauria Clade: †Ornithischia Clade: †Thyreophora Clade: †Ankylosauria Clade: †Euankylosauria Family: †Nodosauridae
Marsh, 1890 Subgroups
Acanthopholis
Anoplosaurus
Dongyangopelta
Gastonia
Gargoyleosaurus
Glyptodontopelta
Horshamosaurus
Hylaeosaurus?
Invictarx
Mymoorapelta?
Priconodon?
Propanoplosaurus
Rhadinosaurus
Sauroplites
Polacanthinae?
Nodosaurinae? Abel, 1919
Acantholipan
Ahshislepelta?
Borealopelta
Niobrarasaurus
Nodosaurus
Patagopelta
Peloroplites?
Sauropelta
Silvisaurus
Taohelong?
Tatankacephalus
Panoplosaurini?
Madzia et al., 2021
Animantarx
Denversaurus
Edmontonia
Panoplosaurus
Texasetes
Struthiosaurini?
Madzia et al., 2021
Ahshislepelta?
Europelta
Hungarosaurus
Niobrarasaurus?
Pawpawsaurus
Stegopelta
Struthiosaurus
Synonyms
Acanthopholididae Nopcsa, 1902
Acanthopholidae Nopcsa, 1917
?Hylaeosauridae Nopcsa, 1902
Palaeoscincidae Nopcsa, 1918
Panoplosauridae Nopcsa, 1929
Struthiosauridae Kuhn, 1966
Nodosauridae is a family of ankylosaurian dinosaurs known from the Late Jurassic to the Late Cretaceous periods in what is now Asia, Europe, North America, and possibly South America. While traditionally regarded as a monophyletic clade as the sister taxon to the Ankylosauridae, some analyses recover it as a paraphyletic grade leading to the ankylosaurids.
Description
[edit]
Nodosaurids, like their sister group the ankylosaurids, were heavily armored dinosaurs adorned with rows of bony armor nodules and spines (osteoderms), which were covered in keratin sheaths. Nodosaurids, like other ankylosaurians, were small- to large-sized, heavily built, quadrupedal, herbivorous dinosaurs, possessing small, leaf-shaped teeth. Unlike ankylosaurids, nodosaurids lacked mace-like tail clubs, instead having more flexible tail tips. Many nodosaurids had spikes projecting outward from their shoulders. One particularly well-preserved nodosaurid "mummy", the holotype of Borealopelta markmitchelli, preserves a nearly complete set of armor in life position, as well as the keratin covering and mineralized remains of the underlying skin, which indicate reddish pigments in a countershading pattern.[1][2]
Classification
[edit]
The family Nodosauridae was erected by Othniel Charles Marsh in 1890, and anchored on the genus Nodosaurus.[3][4]
The clade Nodosauridae was first informally defined by Paul Sereno in 1998 as "all ankylosaurs closer to Panoplosaurus than to Ankylosaurus," a definition followed by Vickaryous, Teresa Maryańska, and Weishampel in 2004. Vickaryous et al. considered two genera of nodosaurids to be of uncertain placement (incertae sedis): Struthiosaurus and Animantarx, and considered the most primitive member of the Nodosauridae to be Cedarpelta.[5]
Following the publication of the PhyloCode, Nodosauridae needed to be formally defined following certain parameters, including that the type genus Nodosaurus was required as an internal specifier. In formally defining Nodosauridae, Madzia and colleagues followed the previously established use for the clade, defining it as the largest clade including Nodosaurus textilis but not Ankylosaurus magniventris. As all phylogenies referenced included both Panoplosaurus and Nodosaurus within the same group relative to Ankylosaurus, the addition of another internal specifier was deemed unnecessary.
Nodosauridae is traditionally composed of the basal clade Polacanthinae (sometimes recovered outside of the Nodosauridae), as well as the Panoplosaurini and Struthiosaurini within the Nodosaurinae. Topology A below demonstrates these relationships, following the phylogenetic analyses of Rivera-Sylva and colleagues (2018), with clade names added by definition from Madzia et al. (2021).[6][7] However, in 2023, Raven and colleagues proposed an alternate phylogeny for nodosaurids; instead of the traditional dichotomous split between nodosaurids and ankylosaurids, their analyses resulted in a paraphyletic grade of these taxa comprising the monophyletic clades Panoplosauridae, Polacanthidae and Struthiosauridae. These results are displayed in Topology B below.[8][9] Corresponding clades are shown in matching colors for clarity, and ⊞ buttons can be clicked to expand nodes:
Nodosaurinae is defined as the largest clade containing Nodosaurus textilis but not Hylaeosaurus armatus, Mymoorapelta maysi, or Polacanthus foxii, and was formally defined in 2021 by Madzia and colleagues, who utilized the name of Othenio Abel in 1919, who created the term to unite Ankylosaurus, Hierosaurus and Stegopelta.[7][10] The name has been significantly refined in content since Abel first used it to unite all quadrupedal, plate-armoured ornithischians,[10] now including a significant number of taxa from the Early Cretaceous through Maastrichtian of Europe, North America, and Argentina. Previous informal definitions of the group described the clade as all taxa closer to Panoplosaurus, or Panoplosaurus and Nodosaurus, than to the early ankylosaurs Sarcolestes, Hylaeosaurus, Mymoorapelta or Polacanthus, which was reflected in the specifiers chosen by Madzia et al. when formalizing the clade following the PhyloCode. The 2018 phylogenetic analysis of Rivera-Sylva and colleagues was used as the primary reference for Panoplosaurini by Madzia et al., in addition to the supplemental analyses of Thompson et al. (2012), Arbour and Currie (2016), Arbour et al. (2016), and Brown et al. (2017).[7][11][12][6][13][1]
Panoplosaurini is defined as the largest clade containing Panoplosaurus mirus, but not Nodosaurus textilis or Struthiosaurus austriacus, and was named in 2021 by Madzia and colleagues for the group found in many previous analyses, both morphological and phylogenetic. Panoplosaurini includes not only the Late Cretaceous Panoplosaurus, Denversaurus and Edmontonia, but also the mid Cretaceous Animantarx and Texasetes, as well as Patagopelta. However, in the study describing it, its authors only placed it as a nodosaurine outside Panoplosaurini.[14] The approximately equivalent clade Panoplosaurinae, named in 1929 by Franz Nopcsa, but was not significantly used until Robert Bakker reused the name in 1988, alongside the new clades Edmontoniinae and Edmontoniidae, which were considered to unite Panoplosaurus, Denversaurus and Edmontonia to the exclusion of other ankylosaurs. As none of the clades were commonly used, or formally named following the PhyloCode, Madzia et al. named Panoplosaurini instead, as the group of taxa fell within the clade Nodosaurinae, and having the same -inae suffix on both parent and child taxon could be confusing in future.[7] The 2018 phylogenetic analysis of Rivera-Sylva and colleagues was used as the primary reference for Panoplosaurini by Madzia et al., in addition to the supplemental analyses of Arbour et al. (2016), Brown et al. (2017), and Zheng et al. (2018).[7][6][13][1][15]
Struthiosaurini is defined as the largest clade containing Struthiosaurus austriacus, but not Nodosaurus textilis or Panoplosaurus mirus, and was named in 2021 by Madzia and colleagues for the relatively stable group found in many previous analyses. Struthiosaurini includes not only the Late Cretaceous European Struthiosaurus, but also the Early Cretaceous European Europelta, the Late Cretaceous European Hungarosaurus, and Stegopelta and Pawpawsaurus from the mid Cretaceous of North America. The approximately equivalent clade Struthiosaurinae, named in 1923 by Franz Nopcsa was previously used to include European nodosaurids, but was never formally named following the PhyloCode, so Madzia et al. named Struthiosaurini instead, as the group of taxa fell within the clade Nodosaurinae, and having the same -inae suffix on both parent and child taxon could be confusing in future.[7] The 2018 phylogenetic analysis of Rivera-Sylva and colleagues was used as the primary reference for Struthiosaurini by Madzia et al., in addition to the supplemental analyses of Arbour et al. (2016), Brown et al. (2017), and Zheng et al. (2018).[7][6][13][1][15]
Biogeography
[edit]
Nodosaurids are known from diverse remains throughout Europe, Asia, and North America.[16]
Some Gondwanan ankylosaurs, including the Antarctican Antarctopelta and Argentinian Patagopelta, were originally regarded as belonging to the Nodosauridae, but later analyses provided support for them belonging to the Parankylosauria, a separate lineage of basal ankylosaurs restricted to the Southern Hemisphere.[17][18][19]
See also
[edit]
Dinosaurs portal
Timeline of ankylosaur research
References
[edit]
Further reading
[edit]
Carpenter, K. (2001). "Phylogenetic analysis of the Ankylosauria." In Carpenter, K., (ed.) 2001: The Armored Dinosaurs. Indiana University Press, Bloomington & Indianapolis, 2001, pp. xv-526
Thompson, R.S.; Parish, J.C.; Maidment, S.C.R.; Barrett, P.M. (2012). "Phylogeny of the ankylosaurian dinosaurs (Ornithischia: Thyreophora)". Journal of Systematic Palaeontology. 10 (2): 301–312. Bibcode:2012JSPal..10..301T. doi:10.1080/14772019.2011.569091. S2CID 86002282.
Arbour, V.M.; Zanno, L.E.; Gates, T. (2016). "Ankylosaurian dinosaur palaeoenvironmental associations were influenced by extirpation, sea-level fluctuation, and geodispersal". Palaeogeography, Palaeoclimatology, Palaeoecology. 449: 289–299. Bibcode:2016PPP...449..289A. doi:10.1016/j.palaeo.2016.02.033.
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The Princeton Field Guide to Dinosaurs: Second Edition [Second ed.] 9781400883141
|
[
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] |
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The best-selling Princeton Field Guide to Dinosaurs remains the must-have book for anyone who loves dinosaurs, from amat...
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en
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dokumen.pub
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https://dokumen.pub/the-princeton-field-guide-to-dinosaurs-second-edition-secondnbsped-9781400883141.html
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Table of contents :
CONTENTS
Preface
Acknowledgments
Introduction
History of Discovery and Research
What Is a Dinosaur?
Dating Dinosaurs
The Evolution of Dinosaurs and Their World
Extinction
After the Age of Dinosaurs
Biology
General Anatomy
Skin, Feathers, and Color
Respiration and Circulation
Digestive Tracts
Senses
Vocalization
Disease and Pathologies
Behavior
Brains, Nerves, and Intelligence
Social Activities
Reproduction
Growth
Energetics
Gigantism
Mesozoic Oxygen
The Evolution—and Loss—of Avian Flight
Dinosaur Safari
If Dinosaurs Had Survived
Dinosaur Conservation
Where Dinosaurs Are Found
Using the Group and Species Descriptions
Group and Species Accounts
Dinosaurs
Theropods
Sauropodomorphs
Ornithischians
Additional Reading
Index: Dinosaur Taxa
Formations
Citation preview
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https://www.chinadaily.com.cn/china/2007-08/16/content_6028905.htm
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New dinosaur species found in Zhejiang
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[
"http://chinadaily.allyes.com/main/adfshow?user=ChinaDailyNetwork|UltimatePage|Ultimate_top_banner&db=chinadaily"
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2007-08-16T00:00:00
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chinadaily
| null |
HANGZHOU: A team of Japanese and Chinese scientists has announced the discovery of a new dinosaur species that used to roam the southwestern part of present-day East China's Zhejiang Province 100 million years ago.
After studying dinosaur fossils uncovered in 2000 at a village near the city of Lishui during the construction of a freeway, scientists named the new species "Zhejiangosaurus Lishuiensis", according to an article in Acta Geologica Sinica, an English-language academic quarterly magazine published by the Geological Society of China.
Local cultural relic workers unearthed the well-preserved fossils of two rear limbs, hips and parts of a spine seven years ago. A six-member research team has been analyzing the fossilized bones at the Zhejiang Provincial Museum of Natural Sciences in Hangzhou.
The researchers concluded that the fossils showed the Zhejiangosaurus Lishuiensis to be an adult herbivorous nodosaur, measuring 6 m in length and more than 1 m in height, with a "mild temperament and ungainly build".
"This particular dinosaur had bony dermal plates covering the top of its body, two lines of sharp spikes that protruded from its back and a clubless tail," said Jin Xingsheng, deputy curator of Zhejiang Provincial Museum of Natural Sciences and also a member of the research team.
Nodosaur fossils are common archaeological finds in North America, but they are quite rare in China. Before the discovery in Lishui, archaeologists had only found similar nodosaur fossils in Luoyang, in Central China's Henan Province.
Jin and four other Chinese scientists were guided by Dr Yoichi Azuma, curator of Japan's Fukui Profectural Dinosaur Museum and the only foreign scientist to have worked on the research team.
"With guidance from Yoichi Azuma, we have devoted a lot of time over the last few years to repairing the fossils, trying to restore the original image of the Zhejiangosaurus Lishuiensis, and a life-size model of the dinosaur will be ready for public viewing in the museum later this year," said Jin.
Fossils of dinosaur eggs belonging to a larger bird-like dinosaur species known as Theropoda were also found in Tiantai, a county situated closer to the coast in Zhejiang, in March this year.
"Our findings regarding the Zhejiangosaurus Lishuiensis will shed light on the climatic conditions in this coastal province of Zhejiang dating back to the mid-Cretaceous Period," Jin said.
Xinhua
(China Daily 08/16/2007 page5)
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https://peerj.com/articles/12362/
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The phylogenetic nomenclature of ornithischian dinosaurs
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[] |
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[
""
] | null |
[
"Gauthier JA",
"Langer MC",
"Novas FE",
"Bittencourt J",
"Ezcurra MS",
"de Queiroz K",
"Cantino PD",
"Ghiselin MT",
"Gilmore CW",
"Glut DF"
] |
2021-05-04T00:00:00
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Ornithischians form a large clade of globally distributed Mesozoic dinosaurs, and represent one of their three major radiations. Throughout their evolutionary history, exceeding 134 million years, ornithischians evolved considerable morphological disparity, expressed especially through the cranial and osteodermal features of their most distinguishable representatives. The nearly two-century-long research history on ornithischians has resulted in the recognition of numerous diverse lineages, many of which have been named. Following the formative publications establishing the theoretical foundation of phylogenetic nomenclature throughout the 1980s and 1990s, many of the proposed names of ornithischian clades were provided with phylogenetic definitions. Some of these definitions have proven useful and have not been changed, beyond the way they were formulated, since their introduction. Some names, however, have multiple definitions, making their application ambiguous. Recent implementation of the International Code of Phylogenetic Nomenclature (ICPN, or PhyloCode) offers the opportunity to explore the utility of previously proposed definitions of established taxon names. Since the Articles of the ICPN are not to be applied retroactively, all phylogenetic definitions published prior to its implementation remain informal (and ineffective) in the light of the Code. Here, we revise the nomenclature of ornithischian dinosaur clades; we revisit 76 preexisting ornithischian clade names, review their recent and historical use, and formally establish their phylogenetic definitions. Additionally, we introduce five new clade names: two for robustly supported clades of later-diverging hadrosaurids and ceratopsians, one uniting heterodontosaurids and genasaurs, and two for clades of nodosaurids. Our study marks a key step towards a formal phylogenetic nomenclature of ornithischian dinosaurs.
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PeerJ
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https://peerj.com/articles/12362/
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Phylogenetic Nomenclature of Ornithischian Clades
For the sake of clarity, all clade names are provided in alphabetical order. The definitions are summarized in Table 1. The extent of all clade names is further depicted on Fig. 1 that shows the relationships of taxa included in the present study as specifiers (both, internal as well as external) and additionally on Figs. 2–4 that represent selected ornithischian-wide phylogenies published within recent years: Madzia, Boyd & Mazuch (2018: Fig. 4B), Dieudonné et al. (2020: Figs. 1 and 2), and Yang et al. (2020: Fig. 12).
Ankylopollexia Sereno, 1986 (converted clade name)
Registration number: 585
Definition. The smallest clade containing Camptosaurus dispar (Marsh, 1879) and Iguanodon bernissartensis Boulenger in Beneden, 1881. This is a minimum-clade definition. Abbreviated definition: min ∇ (Camptosaurus dispar (Marsh, 1879) & Iguanodon bernissartensis Boulenger in Beneden, 1881).
Reference phylogeny. Figure 12 of Madzia, Jagt & Mulder (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 3 of Madzia, Boyd & Mazuch (2018), Figure 20 of Verdú et al. (2018), Figure 9 of Verdú et al. (2020), Figure 11 of McDonald et al. (2021), and Figure 11 of Santos-Cubedo et al. (2021).
Composition. The clade Ankylopollexia comprises Camptosaurus dispar and members of the clade Styracosterna.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Ankylopollexia has been (informally) defined before by Sereno (1998: 62) who applied the minimum-clade definition and used Camptosaurus and Parasaurolophus as the internal specifiers. Since the name has traditionally been used in the exact sense, we apply it to the same clade, but prefer to use Iguanodon bernissartensis as the second internal specifier rather than P. walkeri because the name Ankylopollexia was formed after the stiff cone-shaped thumb that characterizes Iguanodon-grade ornithopods. The inclusion of a different internal specifier does not change the extent of Ankylopollexia under any of the published phylogeny inferences. Also, even though the name derives from an apomorphy, it was never used for an apomorphy-based clade.
Ankylosauria Osborn, 1923 (converted clade name)
Registration number: 588
Definition. The largest clade containing Ankylosaurus magniventris Brown, 1908 but not Stegosaurus stenops Marsh, 1887. This is a maximum-clade definition. Abbreviated definition: max ∇ (Ankylosaurus magniventris Brown, 1908 ~ Stegosaurus stenops Marsh, 1887).
Reference phylogeny. Figure 11 of Arbour & Currie (2016) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 3 of Thompson et al. (2012), Figure 1 of Arbour, Zanno & Gates (2016), Figure 3 of Brown et al. (2017), and Figure 26 of Wiersma & Irmis (2018).
Composition. Under the primary reference phylogeny, Ankylosauria comprises Minmi sp. (= Kunbarrasaurus ieversi), Mymoorapelta maysi, and members of the clades Ankylosauridae and Nodosauridae.
Synonyms. The name Ankylosauromorpha Carpenter, 2001 has been recently used under an alternative systematic scheme for the same branch as Ankylosauria, as defined herein (Norman, 2021; see ‘Discussion’). No other taxon names are currently in use for the same or approximate clade.
Comments. The name Ankylosauria has been (informally) defined before (Carpenter, 1997; Sereno, 1998; Sereno, 2005). These definitions were maximum-clade and used Ankylosaurus (Carpenter, 1997; Sereno, 1998) or Ankylosaurus magniventris (Sereno, 2005) as the internal specifier and Stegosaurus (Carpenter, 1997; Sereno, 1998) or Stegosaurus stenops (Sereno, 2005) as the external specifier. Since Ankylosauria has been ‘traditionally’ used in this sense (though, see also ‘Discussion’), we formalize this definition. Note that Norman (2021) recently provided two phylogenetic definitions for Ankylosauria, a maximum-clade and a minimum-clade. In the maximum-clade definition Norman (2021) used Euoplocephalus and Edmontonia as the internal specifiers and Scelidosaurus as the external specifier, while in the minimum-clade definition the use of the name was anchored on Euoplocephalus and Edmontonia. See ‘Discussion’ for additional comments. Note that the external specifier Stegosaurus stenops is not included in the primary reference phylogeny. From the taxa analyzed by Arbour & Currie (2016), S. stenops is most closely related to Huayangosaurus taibaii (see, e.g., Maidment et al., 2020).
Ankylosauridae Brown, 1908 (converted clade name)
Registration number: 589
Definition. The largest clade containing Ankylosaurus magniventris Brown, 1908 but not Nodosaurus textilis Marsh, 1889. This is a maximum-clade definition. Abbreviated definition: max ∇ (Ankylosaurus magniventris Brown, 1908 ~ Nodosaurus textilis Marsh, 1889).
Reference phylogeny. Figure 11 of Arbour & Currie (2016) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 3 of Thompson et al. (2012), Figure 1 of Arbour, Zanno & Gates (2016), Figure 3 of Brown et al. (2017), Figure 26 of Wiersma & Irmis (2018), and Figure 9 of Zheng et al. (2018).
Composition. Under the primary reference phylogeny, Ankylosauridae comprises Ahshislepelta minor, Aletopelta coombsi, Cedarpelta bilbeyhallorum, Chuanqilong chaoyangensis, Gastonia burgei, Liaoningosaurus paradoxus, and members of the clades Shamosaurinae and Ankylosaurinae.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Ankylosauridae has been (informally) defined before by Sereno (1998, 2005) who applied a maximum-clade definition and used Ankylosaurus magniventris as the internal specifier and Panoplosaurus mirus as the external specifier. Considering that Ankylosauridae has been traditionally used as a sister taxon to Nodosauridae (see, e.g., Thompson et al., 2012 for details), we use a definition that incorporates Nodosaurus textilis as the external specifier. Note that N. textilis is not included in the primary reference phylogeny. Both, A. magniventris and N. textilis were analyzed by, and their relationship is indicated in, Rivera-Sylva et al. (2018a).
Ankylosaurinae Nopcsa, 1918 (converted clade name)
Registration number: 590
Definition. The largest clade containing Ankylosaurus magniventris Brown, 1908 but not Shamosaurus scutatus Tumanova, 1983. This is a maximum-clade definition. Abbreviated definition: max ∇ (Ankylosaurus magniventris Brown, 1908 ~ Shamosaurus scutatus Tumanova, 1983).
Reference phylogeny. Figure 11 of Arbour & Currie (2016) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 3 of Thompson et al. (2012), Figure 1 of Arbour, Zanno & Gates (2016), Figure 8 of Arbour & Evans (2017), Figure 26 of Wiersma & Irmis (2018), and Figure 9 of Zheng et al. (2018).
Composition. Under the primary reference phylogeny, Ankylosaurinae comprises Crichtonpelta benxiensis, Pinacosaurus spp., Saichania chulsanensis, Tarchia kielanae, Tsagantegia longicranialis, Zaraapelta nomadis, ‘Zhejiangosaurus luoyangensis’, and members of the clade Ankylosaurini.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Ankylosaurinae was (informally) defined before (Sereno, 1998; Sereno, 2005; Vickaryous, Maryanska & Weishampel, 2004). All these definitions were maximum-clade and used Ankylosaurus (Sereno, 1998) or Ankylosaurus magniventris (Sereno, 2005; Vickaryous, Maryanska & Weishampel, 2004) as the internal specifiers and Minmi paravertebra and Shamosaurus scutatus (Sereno, 1998), Gargoyleosaurus parkpinorum, Minmi paravertebra, and Shamosaurus scutatus (Sereno, 2005) or only Shamosaurus scutatus (Vickaryous, Maryanska & Weishampel, 2004) as the external specifiers. Owing to the dubious taxonomic status of ‘M. paravertebra’ (Arbour & Currie, 2016) and non-ankylosaurid affinities of G. parkpinorum (e.g., Arbour & Currie, 2016; Rivera-Sylva et al., 2018a; Wiersma & Irmis, 2018; Zheng et al., 2018), we formalize the definition of Vickaryous, Maryanska & Weishampel (2004) in that we use a single external specifier (Shamosaurus scutatus).
Ankylosaurini Arbour & Currie, 2016 (converted clade name)
Registration number: 592
Definition. The largest clade containing Ankylosaurus magniventris Brown, 1908 but not Pinacosaurus grangeri Gilmore, 1933 and Saichania chulsanensis Maryańska, 1977. This is a maximum-clade definition. Abbreviated definition: max ∇ (Ankylosaurus magniventris Brown, 1908 ~ Pinacosaurus grangeri Gilmore, 1933 & Saichania chulsanensis Maryańska, 1977).
Reference phylogeny. Figure 11 of Arbour & Currie (2016) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 1 of Arbour, Zanno & Gates (2016), Figure 8 of Arbour & Evans (2017), Figure 26 of Wiersma & Irmis (2018), and Figure 9 of Zheng et al. (2018).
Composition. Under the primary reference phylogeny, Ankylosaurini comprises Ankylosaurus magniventris, Anodontosaurus lambei, Dyoplosaurus acutosquameus, Euoplocephalus tutus, Nodocephalosaurus kirtlandensis, Scolosaurus cutleri, Talarurus plicatospineus, and Ziapelta sanjuanensis.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Ankylosaurini was first (informally) defined by Arbour & Currie (2016) who applied the maximum-clade definition and used Ankylosaurus magniventris as the internal specifier and Pinacosaurus grangeri and Saichania chulsanensis as the external specifiers. The name was used for a clade that largely includes later-diverging North American ankylosaurines, many of which were previously synonymized with Euoplocephalus tutus (Arbour & Currie, 2013), although under some topologies the name may be more restricted in its use (Thompson et al., 2012).
Aralosaurini Prieto-Márquez et al., 2013 (converted clade name)
Registration number: 593
Definition. The largest clade containing Aralosaurus tuberiferus Rozhdestvensky, 1968 and Canardia garonnensis Prieto-Márquez et al., 2013 but not Lambeosaurus lambei Parks, 1923, Parasaurolophus walkeri Parks, 1922, and Tsintaosaurus spinorhinus Young, 1958. This is a maximum-clade definition. Abbreviated definition: max ∇ (Aralosaurus tuberiferus Rozhdestvensky, 1968 & Canardia garonnensis Prieto-Márquez et al., 2013 ~ Lambeosaurus lambei Parks, 1923 & Parasaurolophus walkeri Parks, 1922 & Tsintaosaurus spinorhinus Young, 1958).
Reference phylogeny. Figure 25 of Prieto-Márquez et al. (2013) is treated here as the primary reference phylogeny. Additional reference phylogeny includes Figure 11 of McDonald et al. (2021).
Composition. Under the primary reference phylogeny, Aralosaurini comprises Aralosaurus tuberiferus and Canardia garonnensis.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name was first (informally) defined by Prieto-Márquez et al. (2013) who applied the minimum-clade definition and used Aralosaurus tuberiferus and Canardia garonnensis as the internal specifiers. Following such definition, however, Aralosaurini would cover the entire lambeosaurine branch under some topologies that include both of the internal specifiers (Kobayashi et al., 2019; Prieto-Márquez et al., 2019; Zhang et al., 2019; Gates, Evans & Sertich, 2021; Kobayashi et al., 2021; Longrich et al., 2021), or would even comprise the same contents as Euhadrosauria (Ramírez-Velasco et al., 2021). Recently, however, McDonald et al. (2021) inferred Aralosaurini as delimited by Prieto-Márquez et al. (2013). Therefore, we define the name but make it inapplicable under a subset of recent phylogenies.
Brachylophosaurini Gates et al., 2011 (converted clade name)
Registration number: 594
Definition. The largest clade containing Brachylophosaurus canadensis Sternberg, 1953 but not Edmontosaurus regalis Lambe, 1917, Hadrosaurus foulkii Leidy, 1858, Kritosaurus navajovius Brown, 1910, and Saurolophus osborni Brown, 1912. This is a maximum-clade definition. Abbreviated definition: max ∇ (Brachylophosaurus canadensis Sternberg, 1953 ~ Edmontosaurus regalis Lambe, 1917 & Hadrosaurus foulkii Leidy, 1858 & Kritosaurus navajovius Brown, 1910 & Saurolophus osborni Brown, 1912).
Reference phylogeny. Figure 18 of Prieto-Márquez, Wagner & Lehman (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 5 of Kobayashi et al. (2019), Figure 11 of Prieto-Márquez et al. (2019), Figure 9 of Zhang et al. (2019), Figure 5 of Zhang et al. (2020), Figure 7 of Kobayashi et al. (2021), and Figure 10 of Longrich et al. (2021).
Composition. Under the primary reference phylogeny, Brachylophosaurini comprises Acristavus gagslarsoni, Brachylophosaurus canadensis, Maiasaura peeblesorum, and Probrachylophosaurus bergei (erroneously named ‘Probrachylophosaurus canadensis’ in the primary reference phylogeny).
Synonyms. The name Maiasaurini Sereno, 2005 is an approximate synonym of Brachylophosaurini. To our knowledge, the name was used only in two recent papers (McFeeters et al., 2021; McFeeters, Evans & Maddin, 2021) that attributed the name to Horner (1992). However, this attribution was due to the adherence of the authors to the Principle of Coordination, as Horner (1992) used the name Maiasaurinae. Nevertheless, all recent phylogenetic studies consistently use Brachylophosaurini (e.g., Freedman Fowler & Horner, 2015; Cruzado-Caballero & Powell, 2017; Xing, Mallon & Currie, 2017; Kobayashi et al., 2019; Zhang et al., 2019; Prieto-Márquez, Wagner & Lehman, 2020; Zhang et al., 2020; Kobayashi et al., 2021; McDonald et al., 2021). No other taxon names are currently in use for the same or approximate clade.
Comments. The name Brachylophosaurini has been (informally) defined before (Gates et al., 2011; Freedman Fowler & Horner, 2015). These definitions were maximum-clade and used Brachylophosaurus, Maiasaura, and Acristavus (Gates et al., 2011) or Brachylophosaurus, Probrachylophosaurus, Maiasaura, and Acristavus (Freedman Fowler & Horner, 2015) as the internal specifiers and Gryposaurus and Saurolophus as the external specifiers. The composition of Brachylophosaurini and the relationships of the clade to other hadrosaurids have been stable across studies since the introduction of the name. Therefore, using more than one internal specifier is unnecessary. We use a definition that ensures Brachylophosaurini does not cover taxa ‘traditionally’ comprised within Edmontosaurini, Kritosaurini, and Saurolophini.
Camptosauridae Marsh, 1885 (converted clade name)
Registration number: 595
Definition. The largest clade containing Camptosaurus dispar (Marsh, 1879) but not Iguanodon bernissartensis Boulenger in Beneden, 1881. This is a maximum-clade definition. Abbreviated definition: max ∇ (Camptosaurus dispar (Marsh, 1879) ~ Iguanodon bernissartensis Boulenger in Beneden, 1881).
Reference phylogeny. Figure 13 of Madzia, Jagt & Mulder (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 20 of Verdú et al. (2018), Figure 11 of Santos-Cubedo et al. (2021), and Figure 9 of Verdú et al. (2020).
Composition. Under the primary reference phylogeny, Camptosauridae comprises Camptosaurus dispar and Cumnoria prestwichii. Under alternative hypotheses, however, Camptosauridae includes only a single unequivocal member, Camptosaurus dispar (e.g., Madzia, Jagt & Mulder, 2020: Fig. 12).
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Camptosauridae was first (informally) defined by Sereno (1998: 62) who used the maximum-clade definition and selected Camptosaurus as the internal specifier and Parasaurolophus as the external specifier. We prefer to use Iguanodon bernissartensis as the external specifier to maintain the ‘node-branch triplet’ (‘node-stem triplet’ of Sereno (1998: 52–54)) comprising Ankylopollexia, Camptosauridae, and Styracosterna (all formally defined in the present paper). The inclusion of a different external specifier does not change the extent of Camptosauridae under any of the published phylogeny inferences.
Centrosaurinae Lambe, 1915 (converted clade name)
Registration number: 596
Definition. The largest clade containing Centrosaurus apertus Lambe, 1905 but not Chasmosaurus belli (Lambe, 1902) and Triceratops horridus Marsh, 1889. This is a maximum-clade definition. Abbreviated definition: max ∇ (Centrosaurus apertus Lambe, 1905 ~ Chasmosaurus belli (Lambe, 1902) & Triceratops horridus Marsh, 1889).
Reference phylogeny. Figure 9 of Chiba et al. (2018) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 10 of Ryan et al. (2017), Figure 13 of Dalman et al. (2018), Figure 10 of Wilson, Ryan & Evans (2020), Figure 4 of Yu et al. (2020), and Figure 23 of Dalman et al. (2021).
Composition. Under the primary reference phylogeny, Centrosaurinae comprises Albertaceratops nesmoi, Diabloceratops eatoni, Machairoceratops cronusi, Medusaceratops lokii, Sinoceratops zhuchengensis, Wendiceratops pinhornensis, Xenoceratops foremostensis, and members of the clades Eucentrosaura and Nasutoceratopsini.
Synonyms. No other taxon names are currently in use for the same or approximate clade. Although Ceratops montanus may fall within the largest clade containing Centrosaurus apertus but not Chasmosaurus belli and Triceratops horridus as well, the name Ceratopsinae Abel, 1919 has not been associated with the same contents as Centrosaurinae in the past. Therefore, Ceratopsinae is not considered to be an approximate synonym of Centrosaurinae. In any case, C. montanus does not seem to be diagnostic beyond Ceratopsidae at present (Dodson, Forster & Sampson, 2004; Mallon et al., 2016). Therefore, its position within the clade is uncertain. Lucas et al. (2016: 202) have argued that Pachyrhinosaurinae von Huene, 1950 has priority over Centrosaurinae under the Article 61 of the ICZN (International Commission on Zoological Nomenclature, 1999). However, the name Pachyrhinosaurinae has not been used in the literature recently and even Lucas et al. (2016) used Centrosaurinae for the clade in question.
Comments. The name Centrosaurinae has been (informally) defined before (Sereno, 1998; Dodson, Forster & Sampson, 2004; Sereno, 2005). These definitions were maximum-clade and used Pachyrhinosaurus (Sereno, 1998), Centrosaurus (Dodson, Forster & Sampson, 2004), or Centrosaurus apertus (Sereno, 2005) as the internal specifier and Triceratops (Sereno, 1998; Dodson, Forster & Sampson, 2004) or Triceratops horridus (Sereno, 2005) as the external specifier. We apply the name Centrosaurinae for the same known contents; adopting the mandatory Centrosaurus apertus as the internal specifier and Chasmosaurus belli and Triceratops horridus as the external specifiers.
Centrosaurini Ryan et al., 2017 (converted clade name)
Registration number: 687
Definition. The largest clade containing Centrosaurus apertus Lambe, 1905 but not Pachyrhinosaurus canadensis Sternberg, 1950. This is a maximum-clade definition. Abbreviated definition: max ∇ (Centrosaurus apertus Lambe, 1905 ~ Pachyrhinosaurus canadensis Sternberg, 1950).
Reference phylogeny. Figure 9 of Chiba et al. (2018) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 7 of Fiorillo & Tykoski (2012), Figure 10 of Ryan et al. (2017), Figure 13 of Dalman et al. (2018), and Figure 23 of Dalman et al. (2021).
Composition. Under the primary reference phylogeny, Centrosaurini comprises Centrosaurus apertus, Coronosaurus brinkmani, Rubeosaurus ovatus (?= Styracosaurus albertensis; see Holmes et al., 2020), Spinops sternbergorum, and Styracosaurus albertensis. Under an alternative hypothesis, Centrosaurini includes only a single unequivocal member, Centrosaurus apertus (Wilson, Ryan & Evans, 2020: Fig. 10). However, a Bayesian analysis of the same matrix and published in the same study reconstructed Centrosaurini to comprise Centrosaurus apertus, Coronosaurus brinkmani, and Spinops sternbergorum (Wilson, Ryan & Evans, 2020: Fig. 9).
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name was first (informally) defined by Ryan et al. (2017) who applied the maximum-clade definition and used Centrosaurus apertus as the internal specifier and Pachyrhinosaurus canadensis as the external specifier. We formalize this definition.
Cerapoda Sereno, 1986 (converted clade name)
Registration number: 597
Definition. The smallest clade containing Iguanodon bernissartensis Boulenger in Beneden, 1881, Pachycephalosaurus wyomingensis (Gilmore, 1931), and Triceratops horridus Marsh, 1889. This is a minimum-clade definition. Abbreviated definition: min ∇ (Iguanodon bernissartensis Boulenger in Beneden, 1881 & Pachycephalosaurus wyomingensis (Gilmore, 1931) & Triceratops horridus Marsh, 1889).
Reference phylogeny. Figure 4 of Madzia, Boyd & Mazuch (2018) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 16 of Han et al. (2018), Figure 25 of Herne et al. (2019), Figure 1 of Dieudonné et al. (2020), and Figure 57 of Barta & Norell (2021).
Composition. Under the primary reference phylogeny, Cerapoda comprises members of the clades Ornithopoda and Marginocephalia.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Cerapoda has been (informally) defined before (Weishampel, 2004; Butler, Upchurch & Norman, 2008). Both types of definitions, minimum-clade as well as maximum-clade, have been proposed for the name. Weishampel (2004) preferred a maximum-clade definition and used Triceratops as the internal specifier and Ankylosaurus as the external specifier, while Butler, Upchurch & Norman (2008) applied a minimum-clade definition, using Triceratops horridus and Parasaurolophus walkeri as the internal specifiers. Subsequent authors followed the latter definition (Boyd, 2015; Madzia, Boyd & Mazuch, 2018; Herne et al., 2019; Yang et al., 2020). We apply a minimum-clade definition as well and use Iguanodon bernissartensis, Pachycephalosaurus wyomingensis, and Triceratops horridus as the internal specifiers. Note that the internal specifiers Pachycephalosaurus wyomingensis and Triceratops horridus are not included in the primary reference phylogeny. The former belongs to Pachycephalosauria (see, e.g., Dieudonné et al., 2020), while the latter is part of Ceratopsia (e.g., Morschhauser et al., 2019), both within Marginocephalia that is indicated on Figure 4 of Madzia, Boyd & Mazuch (2018).
Ceratopsia Marsh, 1890 (converted clade name)
Registration number: 598
Definition. The largest clade containing Ceratops montanus Marsh, 1888 and Triceratops horridus Marsh, 1889 but not Pachycephalosaurus wyomingensis (Gilmore, 1931). This is a maximum-clade definition. Abbreviated definition: max ∇ (Ceratops montanus Marsh, 1888 & Triceratops horridus Marsh, 1889 ~ Pachycephalosaurus wyomingensis (Gilmore, 1931)).
Reference phylogeny. Figure 10 of Morschhauser et al. (2019) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 16 of Han et al. (2018), Figure S1 of Knapp et al. (2018), Figure 1 of Dieudonné et al. (2020), Figure 3 of Yu et al. (2020), and Figure 4 of Yu et al. (2020).
Composition. Under the primary reference phylogeny, Ceratopsia comprises Psittacosaurus spp. and members of the clades Chaoyangsauridae and Neoceratopsia.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Ceratopsia has been (informally) defined before (Dodson, 1997; Sereno, 1998; Sereno, 2005). These definitions were maximum-clade and used Ceratopsidae (Dodson, 1997), Triceratops (Sereno, 1998), or Triceratops horridus (Sereno, 2005) as the internal specifiers and Pachycephalosauridae (Dodson, 1997), Pachycephalosaurus (Sereno, 1998), or Pachycephalosaurus wyomingensis, Heterodontosaurus tucki, Hypsilophodon foxii, and Ankylosaurus magniventris (Sereno, 2005) as the external specifiers. Even though the position of Hypsilophodon foxii and Heterodontosaurus tucki is indeed somewhat unstable across studies (see, e.g., Han et al., 2018; Madzia, Boyd & Mazuch, 2018; Herne et al., 2019; Dieudonné et al., 2020; Yang et al., 2020), inclusion of these taxa among the external specifiers is not necessary. We use a definition similar to that of Sereno (1998) but include the mandatory Ceratops montanus as a second internal specifier. Note that the internal specifier Ceratops montanus and the external specifier Pachycephalosaurus wyomingensis are not included in the primary reference phylogeny. The former belongs to Ceratopsidae (e.g., Mallon et al., 2016), while the latter is part of Pachycephalosauria (see, e.g., Dieudonné et al., 2020).
Ceratopsidae Marsh, 1888 (converted clade name)
Registration number: 599
Definition. The smallest clade containing Centrosaurus apertus Lambe, 1905, Ceratops montanus Marsh, 1888, Chasmosaurus belli (Lambe, 1902), and Triceratops horridus Marsh, 1889. This is a minimum-clade definition. Abbreviated definition: min ∇ (Centrosaurus apertus Lambe, 1905 & Ceratops montanus Marsh, 1888 & Chasmosaurus belli (Lambe, 1902) & Triceratops horridus Marsh, 1889).
Reference phylogeny. Figure 4 of Yu et al. (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 14 of Mallon et al. (2016), Figure S1 of Knapp et al. (2018), Figure 9a of Fowler & Freedman Fowler (2020), Figure 10 of Wilson, Ryan & Evans (2020), and Figure 3 of Yu et al. (2020).
Composition. Under the primary reference phylogeny, Ceratopsidae comprises members of the clades Centrosaurinae and Chasmosaurinae.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Ceratopsidae has been (informally) defined before (Sereno, 1998, Dodson, Forster & Sampson, 2004; Sereno, 2005). These definitions were minimum-clade and used Triceratops and Pachyrhinosaurus (Sereno, 1998), Triceratops and Centrosaurus (Dodson, Forster & Sampson, 2004), and Triceratops horridus and Pachyrhinosaurus canadensis (Sereno, 2005) as the internal specifiers. Considering that Ceratopsidae ‘traditionally’ contains two subclades, Centrosaurinae and Chasmosaurinae, we include the nomenclatural types of these clades, Centrosaurus apertus and Chasmosaurus belli, respectively, as the internal specifiers, and additionally add Triceratops horridus, a common specifier in the nomenclature of ceratopsian clades and the only taxon that has always been used as an internal specifier in the definition of Ceratopsidae. Finally, we also include a fourth internal specifier, the mandatory Ceratops montanus. Even though the taxon is considered a nomen dubium (e.g., Dodson, Forster & Sampson, 2004; Mallon et al., 2016), its placement within the smallest clade comprising centrosaurines and chasmosaurines does not appear to be questionable (see, e.g., Mallon et al., 2016).
Ceratopsoidea Hay, 1902 (converted clade name)
Registration number: 601
Definition. The largest clade containing Ceratops montanus Marsh, 1888 and Triceratops horridus Marsh, 1889 but not Protoceratops andrewsi Granger & Gregory, 1923. This is a maximum-clade definition. Abbreviated definition: max ∇ (Ceratops montanus Marsh, 1888 & Triceratops horridus Marsh, 1889 ~ Protoceratops andrewsi Granger & Gregory, 1923).
Reference phylogeny. Figure 4 of Yu et al. (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure S1 of Knapp et al. (2018), Figure 10 of Morschhauser et al. (2019), and Figure 3 of Yu et al. (2020).
Composition. Under the primary reference phylogeny, Ceratopsoidea comprises Turanoceratops tardabilis, Zuniceratops christopheri, and members of the clade Ceratopsidae.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Ceratopsoidea has been (informally) defined before by Sereno (1998, 2005) who applied a maximum-clade definition and used Triceratops horridus as the internal specifier and Protoceratops andrewsi as the external specifier. We include an additional internal specifier, the mandatory Ceratops montanus.
Chaoyangsauridae Zhao, Cheng & Xu, 1999 (converted clade name)
Registration number: 602
Definition. The largest clade containing Chaoyangsaurus youngi Zhao, Cheng & Xu, 1999 but not Psittacosaurus mongoliensis Osborn, 1923 and Triceratops horridus Marsh, 1889. This is a maximum-clade definition. Abbreviated definition: max ∇ (Chaoyangsaurus youngi Zhao, Cheng & Xu, 1999 ~ Psittacosaurus mongoliensis Osborn, 1923 & Triceratops horridus Marsh, 1889).
Reference phylogeny. Figure 10 of Morschhauser et al. (2019) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 10 of Han et al. (2015), Figure 15 of Han et al. (2018), and Figure 3 of Yu et al. (2020).
Composition. Under the primary reference phylogeny, Chaoyangsauridae comprises Chaoyangsaurus youngi, Hualianceratops wucaiwanensis, Xuanhuaceratops niei, and Yinlong downsi.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Chaoyangsauridae has been (informally) defined before by Han et al. (2015) who applied a maximum-clade definition and used Chaoyangsaurus youngi as the internal specifier and Triceratops horridus and Psittacosaurus mongoliensis as the external specifiers. We formalize this definition.
Chasmosaurinae Lambe, 1915 (converted clade name)
Registration number: 603
Definition. The largest clade containing Chasmosaurus belli (Lambe, 1902) and Triceratops horridus Marsh, 1889 but not Centrosaurus apertus Lambe, 1905. This is a maximum-clade definition. Abbreviated definition: max ∇ (Chasmosaurus belli (Lambe, 1902) & Triceratops horridus Marsh, 1889 ~ Centrosaurus apertus Lambe, 1905).
Reference phylogeny. Figure 9a of Fowler & Freedman Fowler (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 3 of Brown & Henderson (2015), Figure 14 of Mallon et al. (2016), Figure S1 of Knapp et al. (2018), Figure 3 of Campbell et al. (2019), and Figure 4 of Yu et al. (2020).
Composition. Under the primary reference phylogeny, Chasmosaurinae comprises Agujaceratops mariscalensis, Anchiceratops ornatus, Arrhinoceratops brachyops, Bravoceratops polyphemus, Chasmosaurus spp., Coahuilaceratops magnacuerna, Kosmoceratops richardsoni, Navajoceratops sullivani, Pentaceratops sternbergii, Terminocavus sealyi, Utahceratops gettyi, Vagaceratops irvinensis, and members of the clade Triceratopsini.
Synonyms. The taxon Ceratops montanus may also fall within the largest clade containing Chasmosaurus belli and Triceratops horridus but not Centrosaurus apertus (see, e.g., Mallon et al., 2016). In such case, Ceratopsinae Abel, 1919 would be an approximate synonym. Though the name has been advocated to be the proper name for the clade (it has been (informally) defined by Sereno, 1998 and Sereno, 2005), it was actually introduced 4 years later than Chasmosaurinae. Note that the Principle of Coordination, which would make Ceratopsinae attributable to Marsh (1888), rather than to Abel (1919), does not apply under the ICPN (see Note 9.15A.3). Therefore, Ceratopsinae would not have priority over Chasmosaurinae under the ICPN. Anyway, C. montanus does not seem to be diagnostic beyond Ceratopsidae at present (Mallon et al., 2016), and its position within the clade is thus uncertain.
Comments. The name Chasmosaurinae has been (informally) defined before by Dodson, Forster & Sampson (2004) who applied a maximum-clade definition and used Triceratops as the internal specifier and Centrosaurus as the external specifier. We apply the name Chasmosaurinae for the same known contents; adopting Triceratops horridus and the mandatory Chasmosaurus belli as the internal specifiers and Centrosaurus apertus as the external specifier.
Clypeodonta Norman, 2014 (converted clade name)
Registration number: 604
Definition. The smallest clade within Ornithopoda containing Edmontosaurus regalis Lambe, 1917 and Hypsilophodon foxii Huxley, 1869. This is a minimum-clade definition. Abbreviated definition: min ∇ ∈ Ornithopoda (Edmontosaurus regalis Lambe, 1917 & Hypsilophodon foxii Huxley, 1869).
Reference phylogeny. Figure 50 of Norman (2015) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 25 of Herne et al. (2019) and Figure 2 of Dieudonné et al. (2020).
Composition. Under the primary reference phylogeny, Clypeodonta comprises a clade formed by Hypsilophodon foxii, Rhabdodontidae, and Tenontosaurus spp., and a clade uniting Dryosauridae and Ankylopollexia (termed Iguanodontia in Norman, 2015). However, see ‘Comments’ below for discussion of potential alternative composition of Clypeodonta.
Synonyms. No other taxon names are currently in use for the same or approximate clade. Iguanodontia, as reconstructed, for example, by Madzia, Jagt & Mulder (2020) covers a similar taxic composition; though the topology of Madzia, Jagt & Mulder (2020) differs from that of the primary reference phylogeny of Clypeodonta significantly.
Comments. The name Clypeodonta was claimed as being new in two different studies (Norman, 2014: 29; Norman, 2015: 102), although Norman (2015: 170) also cites Norman (2014) as the establishing reference. The use of the name Clypeodonta differed across studies. Originally, Norman (2014, 2015) intended to use it for a subclade of Ornithopoda that (approximately) comprises Hypsilophodon foxii and its relatives, and ornithopods later-diverging than H. foxii, and (informally) defined the name as pertaining to either, the branch of “Parasaurolophus walkeri and all taxa more closely related to P. walkeri than to Thescelosaurus neglectus” (Norman, 2014: 29) or the node of “Hypsilophodon foxii, Edmontosaurus regalis, their most recent common ancestor, and all of its descendants” (Norman, 2015: 170). In both these studies, Clypeodonta is said (Norman, 2014: 29) or figured (Norman, 2015: Fig. 50) to cover the same known contents although neither of the studies included taxa in their analyses that would fall outside the clade (except for Lesothosaurus diagnosticus). Madzia, Boyd & Mazuch (2018) followed the definition of Norman (2015). In their phylogenetic analysis, however, the name covers a much broader contents as one of the internal specifiers of Clypeodonta, Hypsilophodon foxii, is reconstructed outside Cerapoda in that study (Madzia, Boyd & Mazuch, 2018: Fig. 4). Still, Madzia, Boyd & Mazuch (2018: Appendix 1) stated that as Clypeodonta was a relatively new name with no ‘traditional’ meaning, they saw no reason for its redefinition. They also noted, though, that “given the unstable position of H. foxii among neornithischians, the name might have only limited utility” (Madzia, Boyd & Mazuch, 2018: Appendix 1).
Here we define the name Clypeodonta using the minimum-clade definition of Norman (2015). However, by including the part “within Ornithopoda” in the definition, we restrict the use of Clypeodonta only when H. foxii represents an ornithopod (see Article 11.14 of the ICPN), following the original intent of Norman (2014, 2015).
Coronosauria Sereno, 1986 (converted clade name)
Registration number: 605
Definition. The smallest clade containing Protoceratops andrewsi Granger & Gregory, 1923 and Triceratops horridus Marsh, 1889. This is a minimum-clade definition. Abbreviated definition: min ∇ (Protoceratops andrewsi Granger & Gregory, 1923 & Triceratops horridus Marsh, 1889).
Reference phylogeny. Figure 10 of Morschhauser et al. (2019) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure S1 of Knapp et al. (2018), Figure 8A of Arbour & Evans (2019), Figure 3 of Yu et al. (2020), and Figure 4 of Yu et al. (2020).
Composition. Under the primary reference phylogeny, Coronosauria comprises members of the clades Protoceratopsidae and Ceratopsoidea.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Coronosauria has been (informally) defined before by Sereno (1998, 2005) who applied the minimum-clade definition and used Triceratops horridus and Protoceratops andrewsi as the internal specifiers. We formalize this definition.
Corythosauria (new clade name)
Registration number: 746
Definition. The smallest clade containing Corythosaurus casuarius Brown, 1914a, Lambeosaurus lambei Parks, 1923, and Parasaurolophus walkeri Parks, 1922. This is a minimum-clade definition. Abbreviated definition: min ∇ (Corythosaurus casuarius Brown, 1914a & Lambeosaurus lambei Parks, 1923 & Parasaurolophus walkeri Parks, 1922).
Etymology. Derived from the stem of Corythosaurus Brown, 1914a, the name of an included taxon, which combines the Greek words korythos (helmet) and sauros (lizard, reptile).
Reference phylogeny. Figure 18 of Prieto-Márquez, Wagner & Lehman (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 5 of Kobayashi et al. (2019), Figure 11 of Prieto-Márquez et al. (2019), Figure 9 of Zhang et al. (2019), Figure 5 of Zhang et al. (2020), Figure 7 of Kobayashi et al. (2021), and Figure 10 of Longrich et al. (2021).
Composition. Under the primary reference phylogeny, Corythosauria comprises members of the clades Lambeosaurini and Parasaurolophini.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Corythosauria is established for the well-supported node uniting Lambeosaurini and Parasaurolophini, two lambeosaurine clades characterized by their distinctive, ‘crested’ crania.
Dryomorpha Sereno, 1986 (converted clade name)
Registration number: 606
Definition. The smallest clade containing Dryosaurus altus (Marsh, 1878) and Iguanodon bernissartensis Boulenger in Beneden, 1881. This is a minimum-clade definition. Abbreviated definition: min ∇ (Dryosaurus altus (Marsh, 1878) & Iguanodon bernissartensis Boulenger in Beneden, 1881).
Reference phylogeny. Figure 12 of Madzia, Jagt & Mulder (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 20 of Verdú et al. (2018), Figure 2 of Dieudonné et al. (2020), Figure 9 of Verdú et al. (2020), and Figure 11 of Santos-Cubedo et al. (2021).
Composition. Under the primary reference phylogeny, Dryomorpha comprises members of the clades Dryosauridae and Ankylopollexia.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Dryomorpha was first (informally) defined by Sereno (2005) who attributed the name to “(t)he most inclusive clade containing Dryosaurus altus (Marsh, 1878) and Parasaurolophus walkeri Parks, 1922”. However, due to the use of ‘most’, rather than ‘least’, such definition makes the name inapplicable within Ornithischia. Boyd (2015) later corrected the wording and proposed a minimum-clade definition using the same taxa as the internal specifiers. Here we use the same type of definition but replace P. walkeri with I. bernissartensis. This taxon has always been considered a part of Dryomorpha.
Dryosauridae Milner & Norman, 1984 (converted clade name)
Registration number: 607
Definition. The largest clade containing Dryosaurus altus (Marsh, 1878) but not Iguanodon bernissartensis Boulenger in Beneden, 1881. This is a maximum-clade definition. Abbreviated definition: max ∇ (Dryosaurus altus (Marsh, 1878) ~ Iguanodon bernissartensis Boulenger in Beneden, 1881).
Reference phylogeny. Figure 12 of Madzia, Jagt & Mulder (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 20 of Verdú et al. (2018), Figure 9 of Verdú et al. (2020), Figure 57 of Barta & Norell (2021), and Figure 11 of Santos-Cubedo et al. (2021).
Composition. Under the primary reference phylogeny, Dryosauridae comprises Callovosaurus leedsi, ‘Camptosaurus’ valdensis, Dryosaurus altus, Dysalotosaurus lettowvorbecki, Elrhazosaurus nigeriensis, Eousdryosaurus nanohallucis, and Valdosaurus canaliculatus.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. Dryosauridae was first (informally) defined by Sereno (1998: 61) who used the maximum-clade definition and Dryosaurus altus as the internal specifier and Parasaurolophus walkeri as the external specifier. Here we use the same type of definition but replace P. walkeri with I. bernissartensis. This taxon has always been considered outside Dryosauridae.
Edmontosaurini Glut, 1997 (converted clade name)
Registration number: 608
Definition. The largest clade containing Edmontosaurus regalis Lambe, 1917 but not Brachylophosaurus canadensis Sternberg, 1953, Hadrosaurus foulkii Leidy, 1858, Kritosaurus navajovius Brown, 1910, and Saurolophus osborni Brown, 1912. This is a maximum-clade definition. Abbreviated definition: max ∇ (Edmontosaurus regalis Lambe, 1917 ~ Brachylophosaurus canadensis Sternberg, 1953 & Hadrosaurus foulkii Leidy, 1858 & Kritosaurus navajovius Brown, 1910 & Saurolophus osborni Brown, 1912).
Reference phylogeny. Figure 18 of Prieto-Márquez, Wagner & Lehman (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 5 of Kobayashi et al. (2019), Figure 11 of Prieto-Márquez et al. (2019), Figure 9 of Zhang et al. (2019), Figure 5 of Zhang et al. (2020), Figure 7 of Kobayashi et al. (2021), and Figure 10 of Longrich et al. (2021).
Composition. Under the primary reference phylogeny, Edmontosaurini comprises Edmontosaurus spp., Kerberosaurus manakini, Kundurosaurus nagornyi, and Shantungosaurus giganteus.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Edmontosaurini has been (informally) defined before (Sereno, 2005; Xing et al., 2014). Sereno (2005) applied the maximum-clade definition and used Edmontosaurus regalis as the internal specifier and Maiasaura peeblesorum and Saurolophus osborni as the external specifiers. In turn, Xing et al. (2014) applied a minimum-clade definition, with Edmontosaurus and Kerberosaurus as the internal specifiers. We formalize a maximum-clade definition similar to that of Sereno (2005) but replace M. peeblesorum with Brachylophosaurus canadensis, as the representative of Brachylophosaurini, and further add Kritosaurus navajovius and Hadrosaurus foulkii.
Elasmaria Calvo, Porfiri & Novas, 2007 (converted clade name)
Registration number: 609
Definition. The smallest clade containing Macrogryphosaurus gondwanicus Calvo, Porfiri & Novas, 2007 and Talenkauen santacrucensis Novas, Cambiaso & Ambrosio, 2004, provided that it does not include Hypsilophodon foxii Huxley, 1869, Iguanodon bernissartensis Boulenger in Beneden, 1881, or Thescelosaurus neglectus Gilmore, 1913. This is a minimum-clade definition. Abbreviated definition: min ∇ (Macrogryphosaurus gondwanicus Calvo, Porfiri & Novas, 2007 & Talenkauen santacrucensis Novas, Cambiaso & Ambrosio, 2004 | ~ Hypsilophodon foxii Huxley, 1869 ∨ Iguanodon bernissartensis Boulenger in Beneden, 1881 ∨ Thescelosaurus neglectus Gilmore, 1913).
Reference phylogeny. Figure 31 of Rozadilla, Agnolín & Novas, 2019 is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 4 of Madzia, Boyd & Mazuch (2018), Figure 26 of Herne et al. (2019), Figure 2 of Dieudonné et al. (2020), and Figure 57 of Barta & Norell (2021).
Composition. Under the primary reference phylogeny, Elasmaria comprises Anabisetia saldiviai, Atlascopcosaurus loadsi, Fulgurotherium austral, Gasparinisaura cincosaltensis, Kangnasaurus coetzeei, Macrogryphosaurus gondwanicus, Morrosaurus antarcticus, Notohypsilophodon comodorensis, Quantassaurus intrepidus, and Trinisaura santamartaensis.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Elasmaria has been (informally) defined before (Calvo, Porfiri & Novas, 2007; Herne et al., 2019). The definition proposed by Calvo, Porfiri & Novas (2007) was minimum-clade, while the definition of Herne et al. (2019) was maximum-clade. However, both studies used Talenkauen santacrucensis and Macrogryphosaurus gondwanicus as the internal specifiers. Herne et al. (2019) proposed to add Iguanodon bernissartensis and Hypsilophodon foxii as the external specifiers to maintain the use of the name Elasmaria to the ‘traditional’ contents under a hypothesis in which one of the internal specifiers was reconstructed, for example, closer to iguanodontians. We keep the use of a minimum-clade definition (as first proposed for the name). However, even though all phylogenetic analyses consistently reconstruct close relationships between T. santacrucensis and M. gondwanicus, we follow Herne et al. (2019) in that the unsettled placement of elasmarians on the neornithischian phylogenetic tree warrants addition of external specifiers. We include Iguanodon bernissartensis and Hypsilophodon foxii as the external specifiers (following Herne et al., 2019) and further add a third external specifier, Thescelosaurus neglectus, to reflect that elasmarians were already inferred as a clade within Thescelosaurinae, as the sister taxon to Thescelosaurus spp. (Boyd, 2015).
Eucentrosaura Chiba et al., 2018 (converted clade name)
Registration number: 688
Definition. The smallest clade containing Centrosaurus apertus Lambe, 1905 and Pachyrhinosaurus canadensis Sternberg, 1950. This is a minimum-clade definition. Abbreviated definition: min ∇ (Centrosaurus apertus Lambe, 1905 & Pachyrhinosaurus canadensis Sternberg, 1950).
Reference phylogeny. Figure 9 of Chiba et al. (2018) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 7 of Fiorillo & Tykoski (2012), Figure 10 of Ryan et al. (2017), Figure 13 of Dalman et al. (2018), and Figure 23 of Dalman et al. (2021).
Composition. Under the primary reference phylogeny, Eucentrosaura comprises members of the clades Centrosaurini and Pachyrhinosaurini.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name was first (informally) defined by Chiba et al. (2018) who applied the minimum-clade definition and used Centrosaurus apertus and Pachyrhinosaurus canadensis as the internal specifiers. We formalize this definition.
Euceratopsia (new clade name)
Registration number: 610
Definition. The smallest clade containing Leptoceratops gracilis Brown, 1914b, Protoceratops andrewsi Granger & Gregory, 1923, and Triceratops horridus Marsh, 1889. This is a minimum-clade definition. Abbreviated definition: min ∇ (Leptoceratops gracilis Brown, 1914b & Protoceratops andrewsi Granger & Gregory, 1923 & Triceratops horridus Marsh, 1889).
Etymology. Derived from the Greek eu- (true) and formed to show its association to members of Ceratopsia. Note that Euceratopsia does not derive from the name Ceratops Marsh, 1888, and, as such, the taxon does not have to be the internal specifier in the used definition.
Reference phylogeny. Figure 4 of Yu et al. (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 16 of Han et al. (2018), Figure S1 of Knapp et al. (2018), Figure 10 of Morschhauser et al. (2019), and Figure 3 of Yu et al. (2020).
Composition. Under the primary reference phylogeny, Euceratopsia comprises members of the clades Leptoceratopsidae and Coronosauria.
Synonyms. The name Coronosauria Sereno, 1986 covers the same contents under the topology of You & Dodson (2004). However, see ‘Comments’. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Euceratopsia is established for the well-supported node uniting the three latest-diverging clades of ceratopsians – Leptoceratopsidae, Protoceratopsidae, and Ceratopsoidea. The monophyly of the grouping is supported by all recently published phylogenies that infer Euceratopsia to branch into two clades – leptoceratopsids and coronosaurs (protoceratopsids + ceratopsoids). Both these clades comprise representatives that are very close or survived to the Cretaceous/Paleogene mass extinction event (Fowler, 2017: Table S1). It is worth noting that You & Dodson (2004) reconstructed leptoceratopsids to be the sister taxon to Ceratopsoidea, and Protoceratopsidae to be the sister taxon to Leptoceratopsidae + Ceratopsoidea. Under such topology, Euceratopsia becomes a heterodefinitional synonym of Coronosauria, with the latter having priority.
Euhadrosauria Weishampel, Norman & Grigorescu, 1993 (converted clade name)
Registration number: 611
Definition. The smallest clade containing Lambeosaurus lambei Parks, 1923 and Saurolophus osborni Brown, 1912, provided that it does not include Hadrosaurus foulkii Leidy, 1858. This is a minimum-clade definition. Abbreviated definition: min ∇ (Lambeosaurus lambei Parks, 1923 & Saurolophus osborni Brown, 1912 | ~ Hadrosaurus foulkii Leidy, 1858).
Reference phylogeny. Figure 18 of Prieto-Márquez, Wagner & Lehman (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 11 of Prieto-Márquez et al. (2019), Figure 9 of Zhang et al. (2019), Figure 7 of Kobayashi et al. (2021), Figure 10 of Longrich et al. (2021), and Figure 11 of McDonald et al. (2021).
Composition. Under the primary reference phylogeny, Euhadrosauria comprises members of the clades Saurolophinae and Lambeosaurinae.
Synonyms. The name Hadrosauridae Cope, 1869 is an approximate synonym of Euhadrosauria. If Hadrosaurus foulkii nests within the smallest clade containing Saurolophus osborni and Lambeosaurus lambei, and within the ‘Saurolophus branch’ of the clade (see the entry for the name Saurolophinae), the name Hadrosauridae is used for the node instead, and Euhadrosauria becomes inapplicable. Additionally, the name Saurolophidae has been used for the same contents as well (see ‘Comments’).
Comments. The history and application of Euhadrosauria is complicated and has been thoroughly described and discussed by Madzia, Jagt & Mulder (2020: 14–16). We therefore refer to that study for details.
Euiguanodontia Coria & Salgado, 1996 (converted clade name)
Registration number: 612
Definition. The smallest clade containing Camptosaurus dispar (Marsh, 1879), Dryosaurus altus (Marsh, 1878), and Gasparinisaura cincosaltensis Coria & Salgado, 1996, provided that it does not include Tenontosaurus tilletti Ostrom, 1970. This is a minimum-clade definition. Abbreviated definition: min ∇ (Camptosaurus dispar (Marsh, 1879) & Dryosaurus altus (Marsh, 1878) & Gasparinisaura cincosaltensis Coria & Salgado, 1996 | ~ Tenontosaurus tilletti Ostrom, 1970).
Reference phylogeny. Figure 13 of Coria & Salgado (1996) is treated here as the primary reference phylogeny.
Composition. Under the primary reference phylogeny, Euiguanodontia comprises Gasparinisaura and members of the clades Dryosauridae and Ankylopollexia.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Euiguanodontia is applicable only on the condition that G. cincosaltensis, D. altus, and C. dispar form a clade exclusive of T. tilletti, as originally used by Coria & Salgado (1996). We follow the definition advocated by Madzia, Boyd & Mazuch (2018: Appendix 1) and refer to that study for additional comments. Note also that Euiguanodontia must be a subclade of Iguanodontia under the proposed definition because T. tilletti is an internal specifier in the definition of the name. Finally, note that the internal specifiers Dryosaurus altus and Camptosaurus dispar are not included in the primary reference phylogeny. The former belongs to Dryosauridae (e.g., Madzia, Boyd & Mazuch, 2018), while the latter is part of Ankylopollexia (see, e.g., Madzia, Jagt & Mulder, 2020). Both these clades are indicated on Figure 13 of Coria & Salgado (1996).
Euornithopoda Sereno, 1986 (converted clade name)
Registration number: 613
Definition. The largest clade within Ornithopoda containing Iguanodon bernissartensis Boulenger in Beneden, 1881 but not Heterodontosaurus tucki Crompton & Charig, 1962. This is a maximum-clade definition. Abbreviated definition: max ∇ ∈ Ornithopoda (Iguanodon bernissartensis Boulenger in Beneden, 1881 ~ Heterodontosaurus tucki Crompton & Charig, 1962).
Reference phylogeny. Figure 1 of Sereno (1999) is treated here as the primary reference phylogeny.
Composition. Under the primary reference phylogeny, Euornithopoda comprises Tenontosaurus spp. and members of the clades Ankylopollexia, Dryosauridae, and Hypsilophodontidae.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Euornithopoda has been (informally) defined before (Sereno, 1998; Sereno, 2005). These definitions were maximum-clade and used Parasaurolophus as the internal specifier and Heterodontosaurus tucki, Pachycephalosaurus wyomingensis, Triceratops horridus, and Ankylosaurus magniventris (Sereno, 2005) as the external specifiers. Here we define the name Euornithopoda using a similar maximum-clade definition as that of Sereno (1998) but replace Parasaurolophus with Iguanodon bernissartensis. Also, by including the part “within Ornithopoda” in the definition, we restrict the use of Euornithopoda to the branch only when Heterodontosaurus tucki represents an ornithopod (see Article 11.14 of the ICPN), thus maintaining the ‘traditional’ use (Sereno, 1998; Sereno, 2005).
Eurypoda Sereno, 1986 (converted clade name)
Registration number: 614
Definition. The smallest clade containing Ankylosaurus magniventris Brown, 1908 and Stegosaurus stenops Marsh, 1887. This is a minimum-clade definition. Abbreviated definition: min ∇ (Ankylosaurus magniventris Brown, 1908 & Stegosaurus stenops Marsh, 1887).
Reference phylogeny. Figure 3 of Thompson et al. (2012) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 16 of Han et al. (2018) and Figure 1 of Dieudonné et al. (2020).
Composition. Under the primary reference phylogeny, Eurypoda comprises members of the clades Ankylosauria and Stegosauria.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Eurypoda has been (informally) defined before by Sereno (1998) who used Ankylosaurus and Stegosaurus as the internal specifiers. Since Eurypoda has never been proposed an alternative use, we formalize this definition. Note that the internal specifier Stegosaurus stenops is not included in the primary reference phylogeny. The taxon is most closely related to the clade comprising the operational taxonomic units (OTUs) Stegosaurus armatus (nomen dubium according to Galton, 2010; S. armatus has long been the type species of Stegosaurus but was replaced by S. stenops as the type through an ICZN ruling (International Commission on Zoological Nomenclature, 2013)) and Huayangosaurus taibaii (see, e.g., Maidment et al., 2020).
Genasauria Sereno, 1986 (converted clade name)
Registration number: 615
Definition. The smallest clade containing Ankylosaurus magniventris Brown, 1908, Iguanodon bernissartensis Boulenger in Beneden, 1881, Stegosaurus stenops Marsh, 1887, and Triceratops horridus Marsh, 1889. This is a minimum-clade definition. Abbreviated definition: min ∇ (Ankylosaurus magniventris Brown, 1908 & Iguanodon bernissartensis Boulenger in Beneden, 1881 & Stegosaurus stenops Marsh, 1887 & Triceratops horridus Marsh, 1889).
Reference phylogeny. Figure 16 of Han et al. (2018) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 4 of Madzia, Boyd & Mazuch (2018), Figure 25 of Herne et al. (2019), Figure 1 of Dieudonné et al. (2020), Figure 12 of Yang et al. (2020), and Figure 57 of Barta & Norell (2021).
Composition. Under the primary reference phylogeny, Genasauria comprises members of the clades Neornithischia and Thyreophora.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Genasauria has been (informally) defined before (Currie & Padian, 1997; Sereno, 1998; Sereno, 2005; Butler, Upchurch & Norman, 2008). These definitions were minimum-clade and used Thyreophora and Cerapoda (Currie & Padian, 1997), Ankylosaurus and Triceratops (Sereno, 1998), Ankylosaurus magniventris, Triceratops horridus, and Parasaurolophus walkeri (Sereno, 2005), and Ankylosaurus magniventris, Stegosaurus stenops, Triceratops horridus, Parasaurolophus walkeri, and Pachycephalosaurus wyomingensis (Butler, Upchurch & Norman, 2008) as the internal specifiers. In order to maintain the ‘traditional’ concept of Genasauria as a clade comprising Neornithischia and Thyreophora, the internal specifiers in the definition of Genasauria are used from among the taxa representing the four major subclades – Ornithopoda (Iguanodon bernissartensis), Marginocephalia (Triceratops horridus), Ankylosauria (Ankylosaurus magniventris), and Stegosauria (Stegosaurus stenops). Addition of P. wyomingensis as another internal specifier (to include representatives of both marginocephalian clades – Ceratopsia and Pachycephalosauria) is considered unnecessary because pachycephalosaurs have always been inferred to be part of Genasauria as defined herein. Note that the internal specifiers Ankylosaurus magniventris and Triceratops horridus are not included in the primary reference phylogeny. The former belongs to Ankylosauria within Thyreophora (see, e.g., Thompson et al., 2012), while the latter is part of Ceratopsia (e.g., Morschhauser et al., 2019).
Hadrosauridae Cope, 1869 (converted clade name)
Registration number: 616
Definition. The smallest clade containing Hadrosaurus foulkii Leidy, 1858, Lambeosaurus lambei Parks, 1923, and Saurolophus osborni Brown, 1912. This is a minimum-clade definition. Abbreviated definition: min ∇ (Hadrosaurus foulkii Leidy, 1858 & Lambeosaurus lambei Parks, 1923 & Saurolophus osborni Brown, 1912).
Reference phylogeny. Figure 18 of Prieto-Márquez, Wagner & Lehman (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 5 of Kobayashi et al. (2019), Figure 11 of Prieto-Márquez et al. (2019), Figure 9 of Zhang et al. (2019), Figure 5 of Zhang et al. (2020), Figure 7 of Kobayashi et al. (2021), and Figure 10 of Longrich et al. (2021).
Composition. Under the primary reference phylogeny, Hadrosauridae comprises Hadrosaurus foulkii, Eotrachodon orientalis, Latirhinus uitstlani, Aquilarhinus palimentus, and members of the clade Euhadrosauria.
Synonyms. Several taxon names have been historically or recently used as approximate synonyms of Hadrosauridae. Of these, only the names Saurolophidae and Euhadrosauria have recently been attributed to a clade of the same or a similar composition (e.g., Prieto-Márquez, 2010; Verdú et al., 2018; Zhang et al., 2019; Madzia, Jagt & Mulder, 2020; Prieto-Márquez, Wagner & Lehman, 2020; Verdú et al., 2020; Zhang et al., 2020; Kobayashi et al., 2021; Ramírez-Velasco et al., 2021). See ‘Comments’ below.
Comments. The use of Hadrosauridae and other names applied to the same or similar clades (Saurolophidae and Euhadrosauria) have been thoroughly described and discussed by Madzia, Jagt & Mulder (2020: 14–16) who recommended to use Hadrosauridae for the smallest clade containing H. foulkii, S. osborni, and L. lambei; Euhadrosauria for the smallest clade containing S. osborni and L. lambei; and to abandon Saurolophidae. Note that under some phylogenies, in which H. foulkii is reconstructed within the smallest clade containing S. osborni and L. lambei, the names Hadrosauridae and Euhadrosauria, as (informally) defined by Madzia, Jagt & Mulder (2020), become heterodefinitional synonyms. Although such option may still be viewed acceptable, we decided to apply a minimum-clade definition for Euhadrosauria that makes the name inapplicable under such hypothesis.
Hadrosauriformes Sereno, 1997 (converted clade name)
Registration number: 617
Definition. The smallest clade containing Hadrosaurus foulkii Leidy, 1858 and Iguanodon bernissartensis Boulenger in Beneden, 1881. This is a minimum-clade definition. Abbreviated definition: min ∇ (Hadrosaurus foulkii Leidy, 1858 & Iguanodon bernissartensis Boulenger in Beneden, 1881).
Reference phylogeny. Figure 12 of Madzia, Jagt & Mulder (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 20 of Verdú et al. (2018), Figure 3 of Párraga & Prieto-Márquez (2019), Figure 8 of Słowiak et al. (2020), Figure 9 of Verdú et al. (2020) and Figure 11 of McDonald et al. (2021).
Composition. Under the primary reference phylogeny, Hadrosauriformes comprises members of the clades Iguanodontidae and Hadrosauroidea.
Synonyms. If Hypselospinus fittoni nests within the smallest clade containing Hadrosaurus foulkii and Iguanodon bernissartensis, the name Hadrosauriformes is a potential heterodefinitional synonym of Neoiguanodontia (see the name entry). In such case, the name Hadrosauriformes should have priority. The name Iguanodontoidea Hay, 1902 has been also used as an approximate synonym (Sereno, 1986; Norman, 2002). Note that Norman (2002) used Iguanodontoidea for a clade “(s)erially more derived than Camptosaurus” (Norman, 2002: 138) and defined it as “Iguanodon and all iguanodontians more closely related to Edmontosaurus than to Camptosaurus”. Such definition would make Iguanodontoidea applicable for the same clade as Styracosterna (see the name entry). However, Figure 35 of Norman (2002) shows that the name does not cover Lurdusaurus, which should be included within the clade under such maximum-clade definition. Since Norman (2002) considers Iguanodontoidea to be a synonym of Hadrosauriformes of Sereno (1997, 1998, 1999), it is apparent that Norman (2002) concept of Iguanodontoidea would be more similar to that of Hadrosauriformes rather than Styracosterna.
Comments. The name Hadrosauriformes has been (informally) defined before (Sereno, 1998; Norman, 2015; Madzia, Jagt & Mulder, 2020). However, only Madzia, Jagt & Mulder (2020: Table 1) included the mandatory H. foulkii as the internal specifier. We formalize the definition of Madzia, Jagt & Mulder (2020).
Hadrosaurinae Lambe, 1918 (converted clade name)
Registration number: 618
Definition. The largest clade containing Hadrosaurus foulkii Leidy, 1858 but not Lambeosaurus lambei Parks, 1923. This is a maximum-clade definition. Abbreviated definition: max ∇ (Hadrosaurus foulkii Leidy, 1858 ~ Lambeosaurus lambei Parks, 1923).
Reference phylogeny. Figure 5 of Kobayashi et al. (2019) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 13 of Cruzado-Caballero & Powell (2017), Figure 20 of Xing, Mallon & Currie (2017), Figure 5 of Zhang et al. (2020), and Figure 10 of Longrich et al. (2021).
Composition. Under the primary reference phylogeny, Hadrosaurinae comprises Hadrosaurus foulkii and members of the clades Brachylophosaurini, Edmontosaurini, Kritosaurini, and Saurolophini.
Synonyms. The name Saurolophinae Brown, 1914a has been recently used for the same clade (under the hypothesis in which H. foulkii is nested outside the smallest clade containing Saurolophus osborni and Lambeosaurus lambei). See the entry for the name Saurolophinae.
Comments. The name Hadrosaurinae has been (informally) defined before by (Sereno, 1998; Sereno, 2005). Sereno (1998) applied the maximum-clade definition and used Saurolophus as the internal specifier and Parasaurolophus as the external specifier. In turn, Sereno (2005), apparently erroneously, defined Hadrosaurinae as pertaining to “(t)he most inclusive taxon containing Saurolophus osborni Brown, 1912 and Parasaurolophus walkeri Parks, 1922 and including Hadrosaurus foulkii Leidy, 1858”. Our formal maximum-clade definition was formed to make Hadrosaurinae applicable regardless of whether the taxon lies ouside or within the smallest clade containing Saurolophus osborni and Lambeosaurus lambei.
Hadrosauroidea von Huene, 1952 (converted clade name)
Registration number: 619
Definition. The largest clade containing Hadrosaurus foulkii Leidy, 1858 but not Iguanodon bernissartensis Boulenger in Beneden, 1881. This is a maximum-clade definition. Abbreviated definition: max ∇ (Hadrosaurus foulkii Leidy, 1858 ~ Iguanodon bernissartensis Boulenger in Beneden, 1881).
Reference phylogeny. Figure 12 of Madzia, Jagt & Mulder (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 20 of Verdú et al. (2018), Figure 8 of Słowiak et al. (2020), Figure 9 of Verdú et al. (2020), Figure 11 of McDonald et al. (2021), and Figure 11 of Santos-Cubedo et al. (2021).
Composition. Under the primary reference phylogeny, Hadrosauroidea comprises Altirhinus kurzanovi, Batyrosaurus rozhdestvenskyi, Bolong yixianensis, Equijubus normani, Gongpoquansaurus mazongshanensis, Jinzhousaurus yangi, Koshisaurus katsuyama, Mantellisaurus atherfieldensis, Morelladon beltrani, Ouranosaurus nigeriensis, Penelopognathus weishampeli, Proa valdearinnoensis, Probactrosaurus gobiensis, Ratchasimasaurus suranareae, Sirindhorna khoratensis, Xuwulong yueluni, Zuoyunlong huangi, and members of the clade Hadrosauromorpha.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Hadrosauroidea was first (informally) defined by Sereno (1998: 62) who used the maximum-clade definition and Parasaurolophus walkeri as the internal specifier and Iguanodon bernissartensis as the external specifier. We formalize the definition of Madzia, Jagt & Mulder (2020: Table 1) who replaced P. walkeri with H. foulkii.
Hadrosauromorpha Norman, 2014 (converted clade name)
Registration number: 620
Definition. The largest clade containing Hadrosaurus foulkii Leidy, 1858 but not Probactrosaurus gobiensis Rozhdestvensky, 1966. This is a maximum-clade definition. Abbreviated definition: max ∇ (Hadrosaurus foulkii Leidy, 1858 ~ Probactrosaurus gobiensis Rozhdestvensky, 1966).
Reference phylogeny. Figure 12 of Madzia, Jagt & Mulder (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 20 of Verdú et al. (2018), Figure 9 of Verdú et al. (2020), Figure 7 of Kobayashi et al. (2021), and Figure 11 of Santos-Cubedo et al. (2021).
Composition. Under the primary reference phylogeny, Hadrosauromorpha comprises Bactrosaurus johnsoni, Datonglong tianzhenensis, Eolambia caroljonesa, Gilmoreosaurus mongoliensis, Jeyawati rugoculus, Jintasaurus meniscus, Levnesovia transoxiana, Nanyangosaurus zhugeii, ‘Orthomerus dolloi’, Plesiohadros djadokhtaensis, Protohadros byrdi, Tanius sinensis, Tethyshadros insularis, Shuangmiaosaurus gilmorei, Zhanghenglong yangchengensis, and members of the clade Hadrosauridae.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. Hadrosauromorpha was first (informally) defined by Norman (2014: 32) who used the maximum-clade definition and Parasaurolophus walkeri as the internal specifier and Probactrosaurus gobiensis as the external specifier. We formalize the definition of Madzia, Jagt & Mulder (2020: Table 1) who replaced P. walkeri with H. foulkii.
Heterodontosauridae Kuhn, 1966 (converted clade name)
Registration number: 622
Definition. The largest clade containing Heterodontosaurus tucki Crompton & Charig, 1962 but not Iguanodon bernissartensis Boulenger in Beneden, 1881, Pachycephalosaurus wyomingensis (Gilmore, 1931), Stegosaurus stenops Marsh, 1887, and Triceratops horridus Marsh, 1889. This is a maximum-clade definition. Abbreviated definition: max ∇ (Heterodontosaurus tucki Crompton & Charig, 1962 ~ Iguanodon bernissartensis Boulenger in Beneden, 1881 & Pachycephalosaurus wyomingensis (Gilmore, 1931) & Stegosaurus stenops Marsh, 1887 & Triceratops horridus Marsh, 1889).
Reference phylogeny. Figure 4 of Madzia, Boyd & Mazuch (2018) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 25 of Herne et al. (2019), Figure 12 of Yang et al. (2020), and Figure 57 of Barta & Norell (2021).
Composition. Under the primary reference phylogeny, Heterodontosauridae comprises Abrictosaurus consors, Echinodon becklesii, Eocursor parvus, Fruitadens haagarorum, Heterodontosaurus tucki, Lycorhinus angustidens, Manidens condorensis, Pegomastax africana, and Tianyulong confuciusi.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. We follow Sereno (2012) in recognizing Kuhn (1966), rather than Romer (1966), as the author establishing Heterodontosauridae. The name Heterodontosauridae has been (informally) defined before (Sereno, 1998; Sereno, 2005). These definitions were maximum-clade and used Heterodontosaurus as the internal specifier and Parasaurolophus (Sereno, 1998) or Parasaurolophus walkeri, Pachycephalosaurus wyomingensis, Triceratops horridus, and Ankylosaurus magniventris (Sereno, 2005) as the external specifiers. We apply the name Heterodontosauridae for the same known contents; adopting the mandatory Heterodontosaurus tucki as the internal specifier and representatives of all major ornithischian lineages, Ceratopsia (Triceratops horridus), Ornithopoda (Iguanodon bernissartensis), Pachycephalosauria (Pachycephalosaurus wyomingensis), and Thyreophora (Stegosaurus stenops), as the external specifiers. Note that the external specifiers Pachycephalosaurus wyomingensis, Stegosaurus stenops, and Triceratops horridus are not included in the primary reference phylogeny. P. wyomingensis and T. horridus belong to Marginocephalia that is indicated on Figure 4 of Madzia, Boyd & Mazuch (2018), while S. stenops is nested within Thyreophora (e.g., Maidment et al., 2020).
Huayangosauridae Dong, Tang & Zhou, 1982 (converted clade name)
Registration number: 623
Definition. The largest clade containing Huayangosaurus taibaii Dong, Tang & Zhou, 1982 but not Stegosaurus stenops Marsh, 1887. This is a maximum-clade definition. Abbreviated definition: max ∇ (Huayangosaurus taibaii Dong, Tang & Zhou, 1982 ~ Stegosaurus stenops Marsh, 1887).
Reference phylogeny. Figure 12 of Maidment et al. (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 11 of Maidment et al. (2008) and Figure 1 of Raven & Maidment (2017).
Composition. Under the primary reference phylogeny, Huayangosauridae comprises Chungkingosaurus jiangbeiensis and Huayangosaurus taibaii.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Huayangosauridae was first (informally) defined by Galton & Upchurch (2004: 358) who used the maximum-clade definition and selected Huayangosaurus as the internal specifier and Stegosaurus as the external specifier. We formalize this definition.
Hypsilophodontia Cooper, 1985 (converted clade name)
Registration number: 624
Definition. The smallest clade within Ornithopoda containing Hypsilophodon foxii Huxley, 1869 and Tenontosaurus tilletti Ostrom, 1970, provided that it does not include Iguanodon bernissartensis Boulenger in Beneden, 1881. This is a minimum-clade definition. Abbreviated definition: min ∇ ∈ Ornithopoda (Hypsilophodon foxii Huxley, 1869 & Tenontosaurus tilletti Ostrom, 1970 | ~ Iguanodon bernissartensis Boulenger in Beneden, 1881).
Reference phylogeny. Figure 50 of Norman (2015) is treated here as the primary reference phylogeny.
Composition. Under the primary reference phylogeny, Hypsilophodontia comprises Hypsilophodon foxii, Tenontosaurus spp., and members of the clade Rhabdodontidae. However, see ‘Comments’ below for discussion of potential alternative composition of Clypeodonta.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Hypsilophodontia was (informally) defined as pertaining to “Hypsilophodon foxii, Tenontosaurus tilletti, their most recent common ancestor, and all of its descendants” (Norman, 2015: 171). However, such definition does not reflect alternative topologies that do not support Hypsilophodontia as reconstructed by Norman (2015), making it applicable for markedly different contents (see, e.g., Madzia, Boyd & Mazuch, 2018: Fig. 4).
Here we define the name Hypsilophodontia using a similar minimum-clade definition as that of Norman (2015) but by including the part “within Ornithopoda” in the definition, and adding an external specifier, we restrict the use of Hypsilophodontia to the node only when H. foxii represents an ornithopod (see Article 11.14 of the ICPN) and when Hypsilophodon foxii and Tenontosaurus tilletti are more closely related to each other than either is to I. bernissartensis, following the original intent of Norman (2015). Note that the internal specifier Tenontosaurus tilletti is not indicated in the primary reference phylogeny. The taxon is the type species of Tenontosaurus Ostrom, 1970 and is comprised there within the ‘tenontosaurs’.
Hypsilophodontidae Dollo, 1882 (converted clade name)
Registration number: 625
Definition. The largest clade containing Hypsilophodon foxii Huxley, 1869 but not Iguanodon bernissartensis Boulenger in Beneden, 1881 and Rhabdodon priscus Matheron, 1869. This is a maximum-clade definition. Abbreviated definition: max ∇ (Hypsilophodon foxii Huxley, 1869 ~ Iguanodon bernissartensis Boulenger in Beneden, 1881 & Rhabdodon priscus Matheron, 1869).
Reference phylogeny. Figure 2 of Dieudonné et al. (2020) is treated here as the primary reference phylogeny.
Composition. Under the primary reference phylogeny, Hypsilophodontidae comprises Hypsilophodon foxii, Gasparinisaura cincosaltensis, and Parksosaurus warreni.
Synonyms. The name Parksosaurinae has been recently for the same contents (Yang et al., 2020), and attributed (apparently following the Principle of Coordination) to Buchholz (2002). No other taxon names are currently in use for the same or approximate clade.
Comments. Hypsilophodontidae was first (informally) defined by Sereno (1998: 61) who used the maximum-clade definition and Hypsilophodon foxii as the internal specifier and Parasaurolophus walkeri as the external specifier. Here we use the same type of definition but replace P. walkeri with I. bernissartensis. This taxon has always been considered outside Hypsilophodontidae. Additionally, we include Rhabdodon priscus as a second external specifier to prevent the inclusion of Rhabdodontidae within Hypsilophodontidae under the topology of Norman (2015: Fig. 50).
Iguanodontia Baur, 1891 (converted clade name)
Registration number: 626
Definition. The smallest clade containing Dryosaurus altus (Marsh, 1878), Iguanodon bernissartensis Boulenger in Beneden, 1881, Rhabdodon priscus Matheron, 1869, and Tenontosaurus tilletti Ostrom, 1970, provided that it does not include Hypsilophodon foxii Huxley, 1869. This is a minimum-clade definition. Abbreviated definition: min ∇ (Dryosaurus altus (Marsh, 1878) & Iguanodon bernissartensis Boulenger in Beneden, 1881 & Rhabdodon priscus Matheron, 1869 & Tenontosaurus tilletti Ostrom, 1970 | ~ Hypsilophodon foxii Huxley, 1869).
Reference phylogeny. Figure 12 of Madzia, Jagt & Mulder (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 16 of Han et al. (2018), Figure 20 of Verdú et al. (2018), Figure 25 of Herne et al. (2019), and Figure 9 of Verdú et al. (2020).
Composition. Under the primary reference phylogeny, Iguanodontia comprises members of the clade Rhabdodontomorpha, Tenontosaurus spp., and Dryomorpha.
Synonyms. No other taxon names are currently in use for the same or approximate clade. Clypeodonta, as reconstructed by Norman (2015) covers a similar taxic composition; though the topology of Norman (2015) differs from that of the primary phylogeny of Iguanodontia significantly.
Comments. The application of Iguanodontia has been described and discussed by Madzia, Boyd & Mazuch (2018: Appendix 1) and Madzia, Jagt & Mulder (2020: Table 1). We therefore refer to these studies for details. Our definition differs from that advocated by Madzia, Boyd & Mazuch (2018) and Madzia, Jagt & Mulder (2020) in that the name is newly applicable only if it is used for a clade that does not include Hypsilophodon foxii (e.g., it becomes inapplicable under the topology of Norman, 2015: Fig. 50).
Iguanodontidae Bonaparte, 1850 (converted clade name)
Registration number: 627
Definition. The largest clade containing Iguanodon bernissartensis Boulenger in Beneden, 1881 but not Hadrosaurus foulkii Leidy, 1858. This is a maximum-clade definition. Abbreviated definition: max ∇ (Iguanodon bernissartensis Boulenger in Beneden, 1881 ~ Hadrosaurus foulkii Leidy, 1858).
Reference phylogeny. Figure 13 of Madzia, Jagt & Mulder (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 3 of Madzia, Boyd & Mazuch (2018), Figure 20 of Verdú et al. (2018), Figure 32 of Tsogtbaatar et al. (2019), Figure 7 of Kobayashi et al. (2021), and Figure 11 of Santos-Cubedo et al. (2021).
Composition. Under the primary reference phylogeny, Iguanodontidae comprises Barilium dawsoni, Iguanodon bernissartensis, Iguanodon galvensis, and Lurdusaurus arenatus.
Synonyms. The name Iguanodontoidea Hay, 1902 is an approximate synonym of Iguanodontidae (see, e.g., Figure 20 of Verdú et al., 2018). Both these names have been used for various sets of taxa thought or reconstructed to be more closely related to Iguanodon bernissartensis than to hadrosaurids. Considering that significant differences exist between phylogeny reconstructions of Iguanodon-grade ornithopods (e.g., Madzia, Boyd & Mazuch, 2018; Verdú et al., 2018; Madzia, Jagt & Mulder, 2020; McDonald et al., 2021), it is difficult to link either of the names to a certain, stable composition. Here, we prefer to apply the name Iguanodontidae because it is more frequent in the literature and because it was coined 52 years before Iguanodontoidea. It is worth noting that the name Iguanodontoidea has been also used as an approximate synonym of Hadrosauriformes (see the name entry).
Comments. The name Iguanodontidae was first (informally) defined before (Sereno, 1998; Sereno, 2005; Santos-Cubedo et al., 2021). These definitions were maximum-clade and used Iguanodon bernissartensis as the internal specifier and Parasaurolophus walkeri (Sereno, 1998; Sereno, 2005) or Corythosaurus casuarius (Santos-Cubedo et al., 2021) as the external specifier. We apply a similar definition but replace P. walkeri/Corythosaurus casuarius with H. foulkii. Note that even though the study of Santos-Cubedo et al. (2021) appeared after the publication of Phylonyms (de Queiroz, Cantino & Gauthier, 2020), the work does not meet the general requirements for establishing Iguanodontidae as a phylogenetically defined clade name (see Articles 7 of the ICPN), nor it provides anything that would indicate such intention. Specifically, the name Iguanodontidae is not explicitly designated as a converted clade name, no bibliographic citations demonstrating prior application of the name to a taxon approximating the clade for which it is being established have been provided (including the authorship of the preexisting name), and no evidence is provided that the required information has been submitted to the registration database for phylogenetically defined names, the RegNum (registration number is missing). The study specifies the phylogenetic information, such as the placement of the clade on the ornithopod tree and the distribution of apomorphies supporting the existence of the clade, and presents the hypothesized composition of the clade. This information alone, however, would not be sufficient for the name Iguanodontidae to be established as a converted clade name, as required by the ICPN.
Jeholosauridae Han et al., 2012 (converted clade name)
Registration number: 628
Definition. The largest clade outside Hypsilophodontidae or Thescelosauridae containing Jeholosaurus shangyuanensis Xu, Wang & You, 2000 but not Hypsilophodon foxii Huxley, 1869, Iguanodon bernissartensis Boulenger in Beneden, 1881, Pachycephalosaurus wyomingensis (Gilmore, 1931), Thescelosaurus neglectus Gilmore, 1913, and Triceratops horridus Marsh, 1889. This is a maximum-clade definition. Abbreviated definition: max ∇ ∉ Hypsilophodontidae ∨ Thescelosauridae (Jeholosaurus shangyuanensis Xu, Wang & You 2000 ~ Hypsilophodon foxii Huxley, 1869 & Iguanodon bernissartensis Boulenger in Beneden, 1881 & Pachycephalosaurus wyomingensis (Gilmore, 1931) & Thescelosaurus neglectus Gilmore, 1913 & Triceratops horridus Marsh, 1889).
Reference phylogeny. Figure 25 of Herne et al. (2019) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 16 of Han et al. (2018), Figure 4 of Madzia, Boyd & Mazuch (2018), and Figure 57 of Barta & Norell (2021).
Composition. Under the primary reference phylogeny, Jeholosauridae comprises Changchunsaurus parvus, Haya griva, and Jeholosaurus shangyuanensis. Under alternative hypotheses, however, Jeholosauridae includes Jeholosaurus shangyuanensis and Yueosaurus tiantaiensis (e.g., Madzia, Boyd & Mazuch, 2018: Fig. 4; Barta & Norell, 2021: Fig. 57).
Synonyms. The name Jeholosaurinae has been used recently for the same contents (Yang et al., 2020), and attributed (apparently following the Principle of Coordination) to Han et al. (2012). No other taxon names are currently in use for the same or approximate clade.
Comments. We use a maximum-clade definition similar to that of Han et al. (2012), which is the only definition (informally) used for Jeholosauridae. Our definition differs in that we replaced the original representative of Ceratopsia (Protoceratops andrewsi) with a taxon that is widely used in phylogenetic definitions of ornithischian clade names (Triceratops horridus). Additionally, our definition prevents the use of Jeholosauridae under the potential hypotheses in which Jeholosaurus is inferred as part of Hypsilophodontidae or Thescelosauridae. Note that the internal specifiers Pachycephalosaurus wyomingensis and Triceratops horridus are not included in the primary reference phylogeny. The former belongs to Pachycephalosauria (see, e.g., Dieudonné et al., 2020), while the latter is part of Ceratopsia (e.g., Morschhauser et al., 2019), both within Marginocephalia that is indicated on Figure 25 of Herne et al. (2019).
Kritosaurini Glut, 1997 (converted clade name)
Registration number: 629
Definition. The largest clade containing Kritosaurus navajovius Brown, 1910 but not Brachylophosaurus canadensis Sternberg, 1953, Edmontosaurus regalis Lambe, 1917, Hadrosaurus foulkii Leidy, 1858, and Saurolophus osborni Brown, 1912. This is a maximum-clade definition. Abbreviated definition: max ∇ (Kritosaurus navajovius Brown, 1910 ~ Brachylophosaurus canadensis Sternberg, 1953 & Edmontosaurus regalis Lambe, 1917 & Hadrosaurus foulkii Leidy, 1858 & Saurolophus osborni Brown, 1912).
Reference phylogeny. Figure 18 of Prieto-Márquez, Wagner & Lehman (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 5 of Kobayashi et al. (2019), Figure 11 of Prieto-Márquez et al. (2019), Figure 9 of Zhang et al. (2019), Figure 5 of Zhang et al. (2020), Figure 7 of Kobayashi et al. (2021), and Figure 10 of Longrich et al. (2021).
Composition. Under the primary reference phylogeny Kritosaurini comprises Gryposaurus spp., Kritosaurus spp., Rhinorex condrupus, Secernosaurus koerneri, and the specimen ‘Big Bend UTEP 37.7’.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The study of Lapparent & Lavocat (1955) has been cited to be the reference establishing the name Kritosaurini (e.g., Prieto-Márquez, 2014). However, Lapparent & Lavocat (1955) used ‘Kritosaurinés’ rather than ‘Kritosaurini’. The name Kritosaurini was then used by Brett-Surman (1989) and by Glut (1997). Since Brett-Surman (1989) is an unpublished doctoral dissertation, we consider Glut (1997) to be the earliest publication to spell the name Kritosaurini. The name was first (informally) defined by Prieto-Márquez (2014) who applied the minimum-clade definition and used Kritosaurus navajovius, Gryposaurus notabilis, and Naashoibitosaurus ostromi as the internal specifiers. We preserve the original intent of Prieto-Márquez (2014) but prefer to apply the maximum-clade definition. Kritosaurus navajovius is used as the internal specifier and Hadrosaurus foulkii, and representatives of Brachylophosaurini (Brachylophosaurus canadensis), Edmontosaurini (Edmontosaurus regalis), and Saurolophini (Saurolophus osborni), as the external specifiers.
Lambeosaurinae Parks, 1923 (converted clade name)
Registration number: 630
Definition. The largest clade containing Lambeosaurus lambei Parks, 1923 but not Hadrosaurus foulkii Leidy, 1858 and Saurolophus osborni Brown, 1912. This is a maximum-clade definition. Abbreviated definition: max ∇ (Lambeosaurus lambei Parks, 1923 ~ Hadrosaurus foulkii Leidy, 1858 & Saurolophus osborni Brown, 1912).
Reference phylogeny. Figure 18 of Prieto-Márquez, Wagner & Lehman (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 5 of Kobayashi et al. (2019), Figure 11 of Prieto-Márquez et al. (2019), Figure 9 of Zhang et al. (2019), Figure 5 of Zhang et al. (2020), Figure 7 of Kobayashi et al. (2021), and Figure 10 of Longrich et al. (2021).
Composition. Under the primary reference phylogeny, Lambeosaurinae comprises Aralosaurus tuberiferus, Canardia garonnensis, Jaxartosaurus aralensis, and members of the clades Corythosauria and Tsintaosaurini.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Lambeosaurinae has been (informally) defined before (Sereno, 1998; Sereno, 2005; Prieto-Márquez, 2010). These definitions were maximum-clade and used Parasaurolophus (Sereno, 1998) or Lambeosaurus lambei (Prieto-Márquez, 2010) as the internal specifiers and Saurolophus (Sereno, 1998) or Hadrosaurus foulkii, Saurolophus osborni, and Edmontosaurus regalis (Prieto-Márquez, 2010) as the external specifiers. Sereno (2005), apparently erroneously, defined Lambeosaurinae as pertaining to “(t)he most inclusive taxon containing Saurolophus osborni Brown, 1912 but not Parasaurolophus walkeri Parks, 1922 and including Lambeosaurus lambei Parks, 1923”. Our formal maximum-clade definition is similar to that of Prieto-Márquez (2010) though we have removed E. regalis from the external specifiers because the taxon is consistently inferred outside Lambeosaurinae (Kobayashi et al., 2019; Prieto-Márquez et al., 2019; Prieto-Márquez, Wagner & Lehman, 2020; Zhang et al., 2019; Zhang et al., 2020; Gates, Evans & Sertich, 2021; Kobayashi et al., 2021; Longrich et al., 2021; Ramírez-Velasco et al., 2021).
Lambeosaurini Sullivan et al., 2011 (converted clade name)
Registration number: 631
Definition. The largest clade containing Lambeosaurus lambei Parks, 1923 but not Aralosaurus tuberiferus Rozhdestvensky, 1968, Parasaurolophus walkeri Parks, 1922, and Tsintaosaurus spinorhinus Young, 1958. This is a maximum-clade definition. Abbreviated definition: max ∇ (Lambeosaurus lambei Parks, 1923 ~ Aralosaurus tuberiferus Rozhdestvensky, 1968 & Parasaurolophus walkeri Parks, 1922 & Tsintaosaurus spinorhinus Young, 1958).
Reference phylogeny. Figure 18 of Prieto-Márquez, Wagner & Lehman (2020) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 5 of Kobayashi et al. (2019), Figure 11 of Prieto-Márquez et al. (2019), Figure 9 of Zhang et al. (2019), Figure 5 of Zhang et al. (2020), Figure 7 of Kobayashi et al. (2021), and Figure 10 of Longrich et al. (2021).
Composition. Under the primary reference phylogeny, Lambeosaurini comprises Amurosaurus riabinini, Arenysaurus ardevoli, Blasisaurus canudoi, Corythosaurus spp., Hypacrosaurus stebingeri, Hypacrosaurus altispinus, Lambeosaurus spp., Magnapaulia laticaudus, Olorotitan arharensis (misspelled as ‘ararhensis’ in the primary reference phylogeny), Sahaliyania elunchunorum, and Velafrons coahuilensis.
Synonyms. The name Corythosaurini Glut, 1997 is an approximate synonym of Lambeosaurini (e.g., Evans & Reisz, 2007; Gates et al., 2007; Pereda-Suberbiola et al., 2009). However, its use has been discouraged (Prieto-Márquez et al., 2013) and all recent phylogenetic studies preferred to use Lambeosaurini instead (e.g., Xing, Mallon & Currie, 2017; Kobayashi et al., 2019; Prieto-Márquez et al., 2019; Zhang et al., 2020; Kobayashi et al., 2021; Longrich et al., 2021; Ramírez-Velasco et al., 2021). No other taxon names are currently in use for the same or approximate clade.
Comments. Even though Sullivan et al. (2011) did not explicitly formulate the definition of their newly proposed name Lambeosaurini, they noted that their “definition of the Lambeosaurini would be equivalent to node 38 of Prieto-Márquez (2010: fig. 9)” (Sullivan et al., 2011: 417). The name Lambeosaurini was first (informally) defined by Prieto-Márquez et al. (2013) who applied the maximum-clade definition and used Lambeosaurus lambei as the internal specifier and Parasaurolophus walkeri, Tsintaosaurus spinorhinus, and Aralosaurus tuberiferus as the external specifier. Such defined, the use of Lambeosaurini adheres to the original intent of Sullivan et al. (2011). We formalize this definition.
Leptoceratopsidae Nopcsa, 1923 (converted clade name)
Registration number: 632
Definition. The largest clade containing Leptoceratops gracilis Brown, 1914b but not Protoceratops andrewsi Granger & Gregory, 1923 and Triceratops horridus Marsh, 1889. This is a maximum-clade definition. Abbreviated definition: max ∇ (Leptoceratops gracilis Brown, 1914b ~ Protoceratops andrewsi Granger & Gregory, 1923 & Triceratops horridus Marsh, 1889).
Reference phylogeny. Figure 10 of Morschhauser et al. (2019) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure S1 of Knapp et al. (2018), Figure 8A of Arbour & Evans (2019), Figure 3 of Yu et al. (2020), and Figure 4 of Yu et al. (2020).
Composition. Under the primary reference phylogeny, Leptoceratopsidae comprises Cerasinops hodgskissi, Gryphoceratops morrisoni, Helioceratops brachygnathus, Ischioceratops zhuchengensis, Koreaceratops hwaseongensis, Leptoceratops gracilis, Montanoceratops cerorhynchus, Prenoceratops pieganensis, Udanoceratops tchizhovi, Unescoceratops koppelhusae, and Zhuchengceratops inexpectus.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Leptoceratopsidae has been (informally) defined before by Makovicky (2001) who used Leptoceratops gracilis as the internal specifier and Triceratops horridus as the external specifier. Since Leptoceratopsidae has never been proposed an alternative use, we formalize a similar definition that differs only in adding Protoceratops andrewsi as a second external specifier.
Marginocephalia Sereno, 1986 (converted clade name)
Registration number: 633
Definition. The smallest clade containing Ceratops montanus Marsh, 1888, Pachycephalosaurus wyomingensis (Gilmore, 1931), and Triceratops horridus Marsh, 1889. This is a minimum-clade definition. Abbreviated definition: min ∇ (Ceratops montanus Marsh, 1888 & Pachycephalosaurus wyomingensis (Gilmore, 1931) & Triceratops horridus Marsh, 1889).
Reference phylogeny. Figure 16 of Han et al. (2018) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 4 of Madzia, Boyd & Mazuch (2018), Figure 25 of Herne et al. (2019), Figure 1 of Dieudonné et al. (2020), Figure 12 of Yang et al. (2020), and Figure 57 of Barta & Norell (2021).
Composition. Under the primary reference phylogeny, Marginocephalia comprises members of the clades Ceratopsia and Pachycephalosauria.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Marginocephalia has been (informally) defined before (Currie & Padian, 1997; Sereno, 1998; Sereno, 2005; Madzia, Boyd & Mazuch, 2018; Herne et al., 2019). These definitions, except for that of Herne et al. (2019), were minimum-clade and used Ceratopsia and Pachycephalosauria (Currie & Padian, 1997) or Triceratops horridus and Pachycephalosaurus wyomingensis (Sereno, 1998; Sereno, 2005; Madzia, Boyd & Mazuch, 2018) as the internal specifiers. Madzia, Boyd & Mazuch (2018) further included Ceratops montanus as a third internal specifier, stating that “(t)he first definition of Marginocephalia was node-based and used ‘Ceratopsia’ and ‘Pachycephalosauria’ as the internal specifiers […]. To follow the definition, and adhere to the ICPN (Art. 11), we have to use name-bearing species or their type specimens as specifiers which makes the name to be anchored on the types of Ceratops montanus and Pachycephalosaurus wyomingensis. Even if C. montanus may be a nomen dubium, its type specimen is unequivocally nested deeply within Ceratopsia and thus its use does not change the extent of the name” (Madzia, Boyd & Mazuch, 2018: Appendix 1). In turn, Herne et al. (2019) preferred a maximum-clade definition with T. horridus and P. wyomingensis as the internal specifiers and Parasaurolophus walkeri as the external specifier, arguing that “(previous) definitions (were) not complementary with present definitions of Cerapoda and Ornithopoda within a node-stem triplet arrangement of clades” and that “re-definition of Marginocephalia as a stem now mirrors its sister stem clade, Ornithopoda, within a node-based Cerapoda. As a result, this stabilization of definition allows for the definitive assignment of all cerapodan OTUs either as ornithopods or marginocephalians” (Herne et al., 2019: Supplemental Text S1: 4). However, Marginocephalia has never formed such ‘triplet’. When its use in a ‘node-branch triplet’ is considered, it is more closely tied with Ceratopsia and Pachycephalosauria rather than with Cerapoda and Ornithopoda. Here, the internal specifiers in the definition of Marginocephalia are used from among the taxa representing the two major subclades – Ceratopsia (Ceratops montanus and Triceratops horridus) and Pachycephalosauria (Pachycephalosaurus wyomingensis). Note that none of the internal specifiers is included in the primary reference phylogeny. Ceratops montanus and Pachycephalosaurus wyomingensis are name-bearers of Ceratopsia and Pachycephalosauria, respectively, and are deeply nested within these clades (e.g., Mallon et al., 2016; Dieudonné et al., 2020). Triceratops horridus is a late-diverging member of Chasmosaurinae within Ceratopsia (e.g., Morschhauser et al., 2019; Fowler & Freedman Fowler, 2020).
Nasutoceratopsini Ryan et al., 2017 (converted clade name)
Registration number: 689
Definition. The largest clade containing Nasutoceratops titusi Sampson et al., 2013 but not Centrosaurus apertus Lambe, 1905. This is a maximum-clade definition. Abbreviated definition: max ∇ (Nasutoceratops titusi Sampson et al., 2013 ~ Centrosaurus apertus Lambe, 1905).
Reference phylogeny. Figure 9 of Chiba et al. (2018) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 7 of Fiorillo & Tykoski (2012), Figure 10 of Ryan et al. (2017), and Figure 13 of Dalman et al. (2018).
Composition. Under the primary reference phylogeny, Nasutoceratopsini comprises Avaceratops lammersi, Nasutoceratops titusi, and the specimens CMN 8804, MOR 692, and the ‘Malta New Taxon’ (GPDM 63). Under an alternative hypothesis, however, Nasutoceratopsini includes only a single unequivocal member, Nasutoceratops titusi (Dalman et al., 2021: Fig. 23).
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name was first (informally) defined by Ryan et al. (2017) who applied the maximum-clade definition and used Nasutoceratops titusi as the internal specifier and Centrosaurus apertus as the external specifier. We formalize this definition.
Neoceratopsia Sereno, 1986 (converted clade name)
Registration number: 634
Definition. The largest clade containing Triceratops horridus Marsh, 1889 but not Chaoyangsaurus youngi Zhao, Cheng & Xu, 1999 and Psittacosaurus mongoliensis Osborn, 1923. This is a maximum-clade definition. Abbreviated definition: max ∇ (Triceratops horridus Marsh, 1889 ~ Chaoyangsaurus youngi Zhao, Cheng & Xu, 1999 & Psittacosaurus mongoliensis Osborn, 1923).
Reference phylogeny. Figure 10 of Morschhauser et al. (2019) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 16 of Han et al. (2018), Figure S1 of Knapp et al. (2018), and Figure 4 of Yu et al. (2020).
Composition. Under the primary reference phylogeny, Neoceratopsia comprises Aquilops americanus, Archaeoceratops oshimai, Asiaceratops salsopaludalis, Auroraceratops rugosus, ZPAL MgD-I/156 (= Graciliceratops mongoliensis), Liaoceratops yanzigouensis, Mosaiceratops azumai, Stenopelix valdensis, Yamaceratops dorngobiensis, and members of the clade Euceratopsia.
Synonyms. No other taxon names are currently in use for the same or approximate clade.
Comments. The name Neoceratopsia has been (informally) defined before by Sereno (1998, 2005) who applied a maximum-clade definition and used Triceratops horridus as the internal specifier and Psittacosaurus mongoliensis as the external specifier. We further include a second external specifier, Chaoyangsaurus youngi, to ensure that Chaoyangsauridae, a clade usually reconstructed as some of the earliest-diverging ceratopsians (e.g., Han et al., 2018; Knapp et al., 2018; Yu et al., 2020), are maintained outside Neoceratopsia.
Neoiguanodontia Norman, 2014 (converted clade name)
Registration number: 635
Definition. The smallest clade containing Hypselospinus fittoni (Lydekker, 1889), Iguanodon bernissartensis Boulenger in Beneden, 1881, and Parasaurolophus walkeri Parks, 1922. This is a minimum-clade definition. Abbreviated definition: min ∇ (Hypselospinus fittoni (Lydekker, 1889) & Iguanodon bernissartensis Boulenger in Beneden, 1881 & Parasaurolophus walkeri Parks, 1922).
Reference phylogeny. Figure 2.26 of Norman (2014) is treated here as the primary reference phylogeny. Additional reference phylogenies include Figure 50 of Norman (2015), Figure 3 of Párraga & Prieto-Márquez (2019), and Figure 11 of McDonald et al. (2021).
Composition. Under the primary reference phylogeny, Neoiguanodontia comprises Hypselospinus fittoni and members of the clade Hadrosauriformes.
Synonyms. Neoiguanodontia is a potential heterodefinitional synonym of Hadrosauriformes. If Hypselospinus fittoni nests within the smallest clade containing Hadrosaurus foulkii and Iguanodon bernissartensis (e.g., Verdú et al., 2018; Santos-Cubedo et al., 2021: Fig. 11), the name Hadrosauriformes should have priority.
Comments. The application of Neoiguanodontia has been described
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Armored (Genus: 143, Species 174)
MAGNORDER: ORNITHISCHIA SEELEY, 1888
= SUPERORDER: ORNITHISCHIA SEELEY, 1888
= COHORT: ORNITHISCHIFORMES COOPER, 1985
= SUPERORDER: PACHYPODOSAURIA COOPER, 1985
CLADE NAME: PARAPREDENTATA NORMAN, BARON, GARCIA & MȔLLER, 2022
CLADE NAME: PRIONODONTIA OWEN, 1874 emend NORMAN, BARON, GARCIA & MȔLLER, 2022
CLADE NAME: GENASAURIA SERENO, 1986
THYREOPHORA NOPCSA, 1915
= SUBORER: THYREOPHORNIA NOPCSA, 1915
Incertae sedis
Genus: Andhrasaurus ULANSKY, 2014a, b (nomen nudum)
A. indicus ULANSKY, 2014a, b (Type)
Genus: Bienosaurus DONG, 2001 (nomen dubium)
B. lufengensis DONG, 2001 (nomen dubium) (Type)
Genus: Emausaurus HAUBOLD, 1990
E. ernsti HAUBOLD, 1990 (Type)
Genus: Jakapil RIGUETTI, APESTEGUIA & PEREDA-SUBERBIOLA, 2022
J. kaniukura RIGUETTI, APESTEGUIA & PEREDA-SUBERBIOLA, 2022 (Type)
Genus: Scelidosaurus non OWEN, 1859 (nomen conservandum)
S. arizonensis ULANSKY, 2014, a, c (nomen dubium)
Genus: Sinopeltosaurus ULANSKY, 2014b (nomen dubium)
= Sinopelta ULANSKY, 2014a
S. minimus ULANSKY, 2014b (Type)
= Genus: Nova? IRMIS, 2002
= Sinopelta minima ULANSKY, 2014a
= Ornithischia incertae sedis IRMIS & KNOLL, 2008
Genus: Tatisaurus SIMMONS, 1965
T. oehleri SIMMONS, 1965 (Type)
= Scelidosaurus oehleri (SIMMONS, 1965) LUCAS, 199
Genus: Nova? DELAIR & WIMBLEDON, 1993
Gen. sp indet.
Bibliography
Family: SCUTELLOSAURIDAE Lambert, 1990
Genus: Scutellosaurus COLBERT, 1981
S. lawleri COLBERT, 1981 (Type)
S. nova? GAY, 2012
CLADE NAME THYREOPHOROIDEA NOPCSA, 1928 (as per FONSECA, REID, VENNER, DUCNA, GARCIA & MȔLLER, 2024)
ORDER: SCELIDOSAURIA COOPER, 1985
= INFRAORDER: SCELIDOSAURIA COOPER, 1985
= GRADE: BRACHYPODA THULBORN, 1973 (partim)
= Superfamily: SCELIDOSAUROIDEA Cope, 1869 (sensu Zhao, 1983)
Family: SCELIDOSAURIDAE Cope, 1869
= Family: SCELIDOSAURIDIDAE Nopcsa, 1917 (sic)
= Subfamily: SCELIDOSAURINAE Cope, 1869 (sensu Nopcsa, 1923)
Genus: Lusitanosaurus de LAPPARENT & ZBYSZEWSKI, 1957
L. liasicus de LAPPARENT & ZBYSZEWSKI, 1957 (Type)
Genus: Scelidosaurus OWEN, 1859 (nomen conservandum)
= Scelodosaurus GILMORE, 1920 (sic)
= Scelidotherium JAEKEL, 1909 (sic)
S. harrisonii OWEN, 1861 (Type)
= Scelidosaurus harrisoni (OWEN, 1861)
S. sp
Genus: Yuxisaurus YAO, BARRETT, YANG, XU & BI, 2022
Y. kopchicki YAO, BARRETT, YANG, XU & BI, 2022 (Type)
Genus: Nova? WEISHAMPEL, 1990/1992/WANG et al., 1985
Bibliography
SUPERORDER: EURYOPDA SERENO, 1986
GRANDORDER: ANKYLOSAUROMORPHA CARPENTER, 2001
= SUBORDER: ARMATOSAURIA ZHAO, 1983
= PARAORDER: ANKYLOSAURIA OLSHEVSKY, 1998
Incertae sedis
Genus: Nova RIDGWELL & SERENO, 2010
ORDER: ANKYLOSAURIA OSBORN, 1923
SUBORDER: PARANKYLOSAURIA
emend SOTO-ACUNA, VARGAS, KALUZA, LEPPE, BOTELHO, PALMA-LIBERONA, SIMON-GUTSTEIN, FERNANDEZ, ORTIZ, MILLA, ARAVENA, MANRIQUEZ, ALARCON-MUNOZ, PINO, TREVISAN, MANSILLA, HINOJOSA, MUNOZ-WALTHER & RUBILAR-ROGERS, 2021
= CLADE NAME: PARANKYLOSAURIA
SOTO-ACUNA, VARGAS, KALUZA, LEPPE, BOTELHO, PALMA-LIBERONA, SIMON-GUTSTEIN, FERNANDEZ, ORTIZ, MILLA, ARAVENA, MANRIQUEZ, ALARCON-MUNOZ, PINO, TREVISAN, MANSILLA, HINOJOSA, MUNOZ-WALTHER & RUBILAR-ROGERS, 2021
Genus: Antarctopelta SALGADO & GASPARINI, 2006
A. oliveroi SALGADO & GASPARINI, 2006 (Type)
Genus: Kunbarrasaurus LEAHEY, MOLNAR, CARPENTER, WITMER, & SALISBURY, 2015
K. ieversi LEAHEY, MOLNAR, CARPENTER, WITMER, & SALISBURY, 2015 (Type)
= Minmi sp MOLNAR, 1996
K. sp
Genus: Stegouros SOTO-ACUNA, VARGAS, KALUZA, LEPPE, BOTELHO, PALMA-LIBERONA, SIMON-GUTSTEIN, FERNANDEZ, ORTIZ, MILLA, ARAVENA, MANRIQUEZ, ALARCON-MUNOZ, PINO, TREVISAN, MANSILLA, HINOJOSA, MUNOZ-WALTHER & RUBILAR-ROGERS, 2021
S. elengassne SOTO-ACUNA, VARGAS, KALUZA, LEPPE, BOTELHO, PALMA-LIBERONA, SIMON-GUTSTEIN, FERNANDEZ, ORTIZ, MILLA, ARAVENA, MANRIQUEZ, ALARCON-MUNOZ, PINO, TREVISAN, MANSILLA, HINOJOSA, MUNOZ-WALTHER & RUBILAR-ROGERS, 2021 (Type)
SUBORDER: EUANKYLOSAURIA emend SOTO-ACUNA, VARGAS, KALUZA, LEPPE, BOTELHO, PALMA-LIBERONA, SIMON-GUTSTEIN, FERNANDEZ, ORTIZ, MILLA, ARAVENA, MANRIQUEZ, ALARCON-MUNOZ, PINO, TREVISAN, MANSILLA, HINOJOSA, MUNOZ-WALTHER & RUBILAR-ROGERS, 2021
= CLADE NAME: SOTO-ACUNA, VARGAS, KALUZA, LEPPE, BOTELHO, PALMA-LIBERONA, SIMON-GUTSTEIN, FERNANDEZ, ORTIZ, MILLA, ARAVENA, MANRIQUEZ, ALARCON-MUNOZ, PINO, TREVISAN, MANSILLA, HINOJOSA, MUNOZ-WALTHER & RUBILAR-ROGERS, 2021
= INFRAORDER: ANKYLOSAURA OSBORN, 1923
= SUBORDER: ANKYLOSAURIA OSBORN, 1923
Incertae sedis
Genus: Crichtonsaurus DONG, 2002
C. bohlini DONG, 2002 (Type)
Genus: Minmi MOLNAR, 1980
M. paravertebra MOLNAR, 1980 (Type)
M. sp
Genus: Sinankylosaurus WANG, ZHANG, CHEN, CHEN & WANG, 2020
S. zhuchengensis WANG, ZHANG, CHEN, CHEN & WANG, 2020 (Type)
Genus: Spicomellus MAIDMENT, STRACHAN, OURHACHE, SCHEYER, BROWN, FERNANDEZ, JOHANSON, RAVEN & BARRET, 2021
S. afer MAIDMENT, STRACHAN, OURHACHE, SCHEYER, BROWN, FERNANDEZ, JOHANSON, RAVEN & BARRET, 2021 (Type)
Genus: Nova LEAHEY, MOLNAR & SALISBURY, 2019
Gen. sp indet.
Family: POLACANTHIDAE (Weiland, 1911) KIRKLAND, 1998
= Family: HYLAEOSAURIDAE Nopcsa, 1917
= Family: HYLAEOSAURIDIDAE Nopcsa, 1917 (sic)
= Family: POLACANTHINES Lavocat, 1955
= Family: APRAEDENTALIDAE Huene, 1956 (partim)
Genus: Dracopelta GALTON, 1980
D. zbyszewskii GALTON, 1980 (Type)
Genus: Gargoyleosaurus CARPENTER, MILES & CLOWARD, 1998
G. parkpinorum CARPENTER, MILES & CLOWARD, 1998 (Type)
= Gargoyleosaurus parkpini CARPENTER, MILES & CLOWARD, 1998
Genus: Gastonia KIRKLAND, 1998
= Gastonia KIKRLAND, 1997 (nomen nudum)
G. burgei KIRKLAND, 1998 (Type)
= Gastonia burgei KIRKLAND, 1997 (nomen nudum)
G. lorriemcwhinneyae KINNEER, CARPENTER, & SHAW, 2016
G. sp
Genus: Hoplitosaurus LUCAS, 1902
H. marshi (LUCAS, 1901) LUCAS, 1902 (Type)
= Stegosaurus marshi LUCAS, 1901
= Polacanthus marshi (LUCAS, 1901) PEREDA-SUBERBIOLA, 1991
Genus: Horshamosaurus BLOWS, 2015
H. rudgwickensis (BLOWS, 1996) BLOWS, 2015 (Type)
= Polacanthus rudgwickensis BLOWS, 1996
= Polacanthus Nova (Anonymous, 1995 as per BLOWS)
Genus: Hylaeosaurus MANTELL, 1833
= Hyaelosaurus HUENE, 1908 (sic)
= Hylaosaurus PLIENINGER, 1847 (sic)
= Hylacosaurus del CORRO, 1974 (sic)
= Hylaeosaurus SAUVAGE, 1883 (sic)
= Hylaesaurus SAUVAGE, 1883 (sic)
= Hyleosaurus MANTELL, 1837 (sic)
= Hyloesaurus GERVAIS, 1859 (sic)
= Hylosaurus FITZINGER, 1843 (sic)
H. armatus MANTELL, 1833 (Type)
= Hylaeosaurus oweni MANTELL, 1844
H. sp
Genus: Mymoorapelta KIRKLAND & CARPENTER, 1994
M. maysi KIRKLAND & CARPENTER, 1994 (Type)
M. sp
Genus: Polacanthoides NOPCSA, 1928
P. ponderosus NOPCSA, 1928 (Type)
Genus: Polacanthus FOX, 1865 (OWEN, vide HULKE, 1881)
= Euacanthus (A. TENNYSON) H. TENNYSON, 1897 (nomen nudum)
= Palacenthus SAUVAGE, 1883 (sic)
= Polacanthus OWEN vide [Anonymous] 1865 (nomen nudum)
= Polacanthus HUXLEY, 1867 (nomen dubium)
= Polanthus NAISH & MARTILL, 2001 (sic)
= Polecanthus McLOUGHLIN, 1979 (sic)
= Polycacanthus COPE, 1869/KUHL, 1831 (sic)
= Vectensia DELAIR, 1982 (nomen nudum)
= Xoplitosaurus MALEEV, 1954 (sic)
P. foxii HULKE, 1881 (Type)
= Hylaeosaurus foxii (HULKE, 1881) COOMBS, 1971
= Euacanthus vectianus (A. TENNYSON) H. TENNYSON, 1897 (nomen nudum)
Note: The first name of Polacanthus?
= Polcanathus becklesi HENNING, 1924 (nomen nudum)
= Polacanthus foxi SEELEY, 1891 non HULKE, 1881
= Vectensia (no species name) DELAIR, 1982
= Polanthus foxii NAISH & MARTILL, 2001 (sic)
P. sp
Genus: Taohelong YANG, YOU, LI, & KONG, 2013
T. jinchengensis YANG, YOU, LI, & KONG, 2013 (Type)
Genus: Nova? = Hoplitosaurus (?) sp BODILY, 1970 (= cf. Sauropelta sp CARPENTR, KIRKLAND, BURGE & BIRD, 1999)
Gen. sp indet.
CLADE NAME: PANOPLOSAURINI MADZIA, ARBOUR, BOYD, FARKE, CRUZADO-CABALLERO & EVANS, 2021
= CLADE NAME: STRUTHIOSAURINI MADZIA, ARBOUR, BOYD, FARKE, CRUZADO-CABALLERO & EVANS, 2021
Family: NODOSAURIDAE Marsh, 1890
= Family: NODOSAURIDIDAE Nopcsa, 1923 (sic)
= Family: NOTOSAURIDAE Koken, 1900 (sic)
= Subfamily: NODOSAURINAE Marsh, 1890 (sensu Abel, 1919)
= Family: APRAEDENTALIDAE Huene, 1956 (partim)
Genus: Acantholipan RIVERA-SYLVA, FREY, STINNESBECK, CARBOT-CHANONA, SANCHEZ-URIBE & GUZMAN-GUTIERREZ, 2018
A. gonzalezi RIVERA-SYLVA, FREY, STINNESBECK, CARBOT-CHANONA, SANCHEZ-URIBE & GUZMAN-GUTIERREZ, 2018 (Type)
Genus: Animantarx CARPENTER, KIRKLAND, BURGE & BIRD, 1999
A. ramaljonesi CARPENTER, KIRKLAND, BURGE & BIRD, 1999 (Type)
Genus: Borealopelta BROWN, HENDERSON, VINTHER, FLETCHER, SISTIAGA, HERRERA & SUMMONS, 2017
B. markmitchelli BROWN, HENDERSON, VINTHER, FLETCHER, SISTIAGA, HERRERA & SUMMONS, 2017 (Type)
Genus: Hierosaurus WIELAND, 1909 (nomen dubium)
= Heirosaurus COLBERT, 1961 (sic)
= Xierosaurus MALEEV, 1954 (sic)
H. sternbergi WIELAND, 1909 (Type)
= Nodosaurus sternbergi (WIELAND, 1909) COOMBS, 1978
= Hadrosaurus sternbergi (WIELAND, 1909) LANE, 1946 (sic)
Genus: Invictarx McDONALD & WOLFE, 2018
I. zephyri McDONALD & WOLFE, 2018 (Type)
Genus: Niobrarasaurus CARPENTER, DILKES & WEISHAMPEL, 1995
N. coleii (MEHL 1936) CARPENTER, DILKES & WEISHAMPEL, 1995 (Type)
= Hierosaurus coleii MEHL, 1936
= Nodosaurus coleii (MEHL, 1936) COOMBS, 1978
Genus: Patagopelta RIGUETTI, PEREDA-SUBERBIOLA, PONCE, SALGADO, APESTEGUIA, ROZADILLA & ARBOUR, 2022
P. cristata RIGUETTI, PEREDA-SUBERBIOLA, PONCE, SALGADO, APESTEGUIA, ROZADILLA & ARBOUR, 2022 (Type)
Genus: Pawpawsaurus LEE, 1996
P. campbelli LEE, 1996 (Type)
Genus: Peloroplites CARPENTER, BARTLETT, BIRD & BARRICK, 2008
P. cedrimontanus CARPENTER, BARTLETT, BIRD & BARRICK, 2008 (Type)
Genus: Texasetes COOMBS, 1995
T. pleurohalio COOMBS, 1995 (Type)
Genus: Vectipelta POND, STRACHAN, RAVEN, SIMPSON, MORGAN & MAIDMENT, 2023
V. barretti POND, STRACHAN, RAVEN, SIMPSON, MORGAN & MAIDMENT, 2023 (Type)
Genus: Zhejiangosaurus LU, JIN, SHENG, LI, WANG & AZUMA, 2007
Z. lishuiensis LU, JIN, SHENG, LI, WANG & AZUMA, 2007 (Type)
Genus: Zhongyuansaurus XU, LU, ZHANG, JIA, HU, ZHANG, WU & JI, 2007
Z. luoyangensis XU, LU, ZHANG, JIA, HU, ZHANG, WU & JI, 2007 (Type)
Genus: Nova BIRD, BURGE & CARPENTER, 2003
Genus: Nova DONG, 1992
Genus: Nova HUNT, HUPS, LOCKLEY, KIRKLAND, BRITT, 1994/HUNT, LOCKLEY, HUPS, & SCHULTZE, 1995
Incertae sedis
= Family: ACANTHOPHOLIDIDAE Nopcsa, 1902
= Family: ACANTHOPHOLIDAE Nopcsa, 1917 (sic)
= Subfamily: ACANTHOPHOLIDINAE Nopcsa, 1902 (sensu Huene, 1956)
= Subfamily: ACANTHOPHOLINAE Nopcsa, 1902 (sensu Nopcsa, 1923)
= Family: PALAEOSCINCIDAE Nopcsa, 1918
Genus: Acanthopholis HUXLEY, 1867 (nomen dubium)
= Acanthopholis KUHN, 1939 (sic)
A. horridus HUXLEY, 1867 (Type)
= Acanthopholis horrida LOMAX & TAMURA, 2014 (sic)
A. hughesii (SEELEY, 1871) (nomen nudum) as per PEREDA-SUBERBIOLA & BARRETT, 1998
A. keepingi SEELEY, 1869 (nomen nudum) as per PEREDA-SUBERBIOLA & BARRETT, 1998
A? platypus SEELEY, 1871 (nomen dubium)
= Acanthopholis platypus SEELEY, 1869 (nomen nudum)
= Acantopholis platypus KUHN, 1939 (sic)
A. macrocercus SEELEY, 1879 (nomen dubium)
A? stereocercusSEELEY, 1879 (nomen dubium)
A? eucercus SEELEY, 1879 (nomen dubium)
A. sp
Genus: Anoplosaurus SEELEY, 1879
= Anoplocephalus HENNING, 1924 (sic)
A. curtonotus SEELEY, 1879 (Type)
= Acanthopholis curtonotus (SEELEY, 1879) NOPCSA, 1902
A. major SEELEY, 1879 (nomen dubium)
= Acanthopholis major (SEELEY, 1879) NOPCSA, 1902 (nomen dubium)
Genus: Brachypodosaurus CHAKRAVARTI, 1934 (nomen dubium)
B. gravis CHAKRAVARTI, 1934 (Type)
Genus: Cryptosaurus SEELEY, 1869
= Cryptodraco LYDEKKER, 1889 (nomen dubium)
= Cryptodraco DELAIR, 1959 (sic)
Note: Lydekker’s (1889) incorrectly believed Crytposaurus was preoccupied byGeoffroy Saint-Hilaire (1833), which was a typographical error for Cystosaurus.
C. eumerus SEELEY, 1869 (Type)
= Cryptodraco eumerus (SEELEY, 1869) LYDEKKER, 1889 (nomen dubium)
Genus: Danubiosaurus BUNZEL, 1871 (nomen dubium)
= Danubriosaurus ROMER, 1966 (sic)
D. anceps BUNZEL, 1871 (Type)
Genus: Denversaurus BAKKER, 1988
D. schlessmani BAKKER, 1988 (Type)
= Denversarus schlessmani HUNT & LUCAS, 1992 (sic)
= Edmontonia sp CARPENTER & BREITHAUPT, 1986
Genus: Palaeoscincus LEIDY, 1856 (nomen dubium)
= Palaeosincus MALEEV, 1956 (sic)
P. costatus LEIDY, 1856 (Type)
P. latus MARSH, 1892 (nomen dubium)
P. sp
Genus: Priconodon MARSH, 1888 (nomen dubium)
= Princonodon LULL, 1911 (sic)
P. crassus MARSH, 1888 (Type)
= Stegosaurus crassus (MARSH, 1888) HENNING, 1915 (nomen dubium)
Genus: Priodontognathus SEELEY, 1875 (nomen dubium)
= Priodontosaurus ROMER, 1966 (sic)
P. phillipsii (SEELEY, 1869) SEELEY, 1875 (Type)
= Iguanodon phillipsii SEELEY, 1869 (nomen dubium)
Genus: Propanoplosaurus STANFORD, WEISHAMPEL & DELEON, 2011
P. marylandicus STANFORD, WEISHAMPEL & DELEON, 2011 (Type)
Note: I believe this to be a concretion and not a real fossil.
Genus: Sarcolestes LYDEKKER, 1893
S. leedsi LYDEKKER, 1893 (Type)
S. sp
Gen. sp indet.
Subfamily: EDMONTONIINAE L.S. Russell, 1940
= Family: EDMONTONIIDAE L. S. Russell, (sensu Bakker, 1988)
Genus: Chassternbergia (BAKKER, 1988) OLSHEVSKY, 1991
= Edmontonia Subgenus Chasternbergia BAKKER, 1988
C. rugosidens (GILMORE, 1930 ) OLSHEVSKY, 1991 (Type)
= Palaeoscincus rugosidens GILMORE, 1930
= Edmontonia rugosidens (GILMORE, 1930) RUSSELL, 1939
= Panoplosaurus rugosidens (GILMORE, 1930) COOMBS, 1979
= Edmontonia Subgenus Chassternbergia rugosidens (GILMORE, 1930) BAKKER, 1988
= Edmontia rugosidens HUNT & LUCAS, 1992 (sic)
Sp 1 or sbsp 1, BAKKER, 1988
Genus: Edmontonia C. M. STERNBERG, 1928
= Edmontonia Subgenus Edmontonia BAKKER, 1988
= Edmontia HUNT & LUCAS, 1992 (sic)
E. longiceps C. M. STERNBERG, 1928 (Type)
= Panoplosaurus longiceps (C. M. STERNBERG, 1928) COOMBS, 1979
= Edmontonia Subgenus Edmontonia longiceps (C. M. STERNBERG, 1928) BAKKER, 1988
= Edmontia longiceps HUNT & LUCAS, 1992 (sic)
E. australis FORD, 2000
E. sp
Subfamily: PANOPLOSAURINAE Nopcsa, 1919
= Subfamily: PANOPLOSAURINES Nopcsa, 1919 (sensu de Lapparent & Lavocat, 1955)
Genus: Panoplosaurus LAMBE, 1919
= Panaplosaurus GALTON, 1981 (sic)
P. mirus LAMBE, 1919 (Type)
P. sp
Subfamily: NODOSAURINAE Marsh, 1890 (sensu Abel, 1919)
Genus: Nodosaurus MARSH, 1889
= Modosaurus KEYES, 1894 (sic)
N. textilis MARSH, 1889 (Type)
= Modosaurus textilis KEYES, 1894 (sic)
Subfamily: SAUROPELTINAE Ford, 2000
Genus: Sauropelta OSTROM, 1970
= Peltosaurus BROWN vide CHURE & McINTOSH, 1989/COPE, 1873
=Peltosaurus GLUT, 1972/COPE, 1873 (nomen nudum)
S. edwardsorum OSTROM, 1970 (Type)
= Sauropelta edwardsi OSTROM, 1970
S. sp
Genus: Silvisaurus EATON, 1960
S. condrayi EATON, 1960 (Type)
Genus: Tatankacephalus PARSONS & PARSONS, 2009
T. cooneyorum PARSONS & PARSONS, 2009 (Type)
Genus: Nova KRUMENACKER, CARPENTER, MOORE & VARRICCHO, 2023
Subfamily: STRUTHIOSAURINAE Nopcsa, 1929
= Family: STRUTHIOSAURIDAE Nopcsa, 1929 (sensu Kuhn, 1966)
Genus: Europelta KIRKLAND, ALCALA, LOEWEN, ESPILEZ, MAMPEL & WIERSMA, 2013
E. carbonensis KIRKLAND, ALCALA, LOEWEN, ESPILEZ, MAMPEL & WIERSMA, 2013 (Type)
Genus: Hoplosaurus SEELEY, 1881 (nomen dubium)
H. ischyrus SEELEY, 1881 (Type)
= Nodosaurus ischyrus (SEELEY, 1881) NOPCSA, 1901 (nomen dubium)
= Nodosaurus (Hoplosaurus) ischyrus (SEELEY, 1881) NOPCSA, 1901 (nomen dubium)
= Hoplosaurus insignis SAUVAGE, 1882 (sic)
Genus: Hungarosaurus OSI, 2005
H. tormai OSI, 2005 (Type)
Genus: Leipsanosaurus NOPCSA, 1918 (nomen dubium)
= Lepanosaurus ROMER, 1966 (sic)
L. noricus NOPCSA, 1918 (Type)
= Struthiosaurus noricus (NOPCSA, 1918) NOPCSA, 1923 (nomen dubium)
Genus: Rhodanosaurus NOPCSA, 1929 (nomen dubium)
= Rhodanosaurus ludguensis NOPCSA, 1929 (Type)
= Struthiosaurus? ludgunensis (NOPCSA, 1929) (nomen dubium)
= Struthiosaurus lugdunensis de LAPPARENT & LAVOCAT, 1955 (sic)
Genus: Struthiosaurus BUNZEL, 1870
= Crataeomus SEELEY, 1881 (nomen dubium)
= Plerropletus TUMANOVA, 1987 (sic)
= Pleuropeltis COOMBS, 1971 (sic)
= Pleuropeltus SEELEY, 1881 (nomen dubium)
Note: Originally described as a skull element, 2 costals and scapula of a large turtle and referred to Struthiosaurus by PEREDA-SUBERBIOLA & GALTON, 2001)
= Pluropeltus ROZHDESTVENSKY & TATARINOV, 1964 (sic)
S. austriacus BUNZEL, 1871 (Type)
= Danubiosaurus anceps BUNZEL, 1871 (nomen dubium) (partim)
= Hylaeosaurus sp BUNZEL, 1871
= Scelidosaurus sp BUNZEL, 1871
S? transilvanicus NOPCSA, 1915
= Struthiosaurus transylvanicus NOPCSA, 1915 (sic)
= Struthiosaurus transsylvanicus NOPCSA, 1929 (sic)
= Struthiosaurus transilvaticus COOMBS, 1971 (sic)
= Crataeomus lepidophorus SEELEY, 1881 (nomen dubium)
= Struthiosaurus lepidophorus (SEELEY, 1881) NOPCSA, 1923 (nomen dubium)
= Crataeomus pawlowitschii SEELEY, 1881 (nomen dubium)
= Struthiosaurus pawolowitschii (SEELEY, 1881) NOPCSA, 1915 (nomen dubium)
= Pleuropletus suessii SEELEY, 1881(nomen diubium)
S. languedocensis GARCIA & PEREDA SUBERBIOLA, 2003
S. sp
Bibliography
Family: ANKYLOSAURIDAE Brown, 1908
= Superfamily: ANKYLOSAUROIDEA Brown, 1908 (sensu Huene, 1914)
Genus: Datai XING, NIU, MALLON & MIYASHITA, 2024
D. yingliangis XING, NIU, MALLON & MIYASHITA, 2024 (Type)
Incertae sedis
Genus: Amtosaurus KURZANOV & TUMANOV, 1978
A. magnus KURZANOV & TUMANOV, 1978 (Type)
Genus: Chuanqilong HAN, ZHENG, HU, XU & BARRETT, 2014
C. chaoyangensis HAN, ZHENG, HU, XU & BARRETT, 2014 (Type)
Genus: Dyoplosaurus non PARKS, 1924
D. giganteus MALEEV, 1956 (nomen dubium)
= Euoplocephalus giganteus (MALEEV, 1956) COOMBS, 1978
Genus: Heishansaurus BOHLIN, 1953 (nomen dubium)
= Heischansaurus ROZHDESTVENSKY, 1977 (sic)
= Heishanasaurus GLUT, 1972 (sic)
= Heishanosaurus SWINTON, 1979 (sic)
H. pachycephalus BOHLIN, 1953 (Type)
Genus: Liaoningosaurus XU, WANG & YOU, 2001
L. paradoxus XU, WANG & YOU, 2001 (Type)
Genus: Stegosaurides BOHLIN, 1953 (nomen dubium)
= Stegosauroides COLBERT, 1961 (sic)
S. excavatus BOHLIN, 1953 (Type)
Genus: Tianchiasaurus DONG, 1994
= Tianchiasaurus DONG, 1993 (sic)
= Jurassosaurus Anonymous, 1993 (nomen nudum)
= SangonghesaurusZHAO, 1983 (nomen nudum)
T. nedegoapeferimoroum emend DONG, 1993 (Type)
= Tianchiasaurus nedegoapeferima DONG, 1993
= Jurassosaurus nedegoapeferkimorum Anonymous, 1993 (nomen nudum)
Genus: Nova? CHATTERJEE, 1995
Gen. sp indet.
Subfamily: ANKYLOSAURINAE Brown, 1908 (sensu Nopcsa, 1923)
Genus: Ahshislepelta BURNS & SULLIVAN, 2011
A. minor BURNS & SULLIVAN, 2011 (Type)
Genus: Bissektipelta PARISH & BARRETT, 2004
B. archibaldi (AVERIANOV, 2002) PARISH & BARRETT, 2004
= Amtosaurus archibaldi AVERIANOV, 2002
Genus: Crichtonpelta ARBOUR & CURRIE, 2015
C. benxiensis (LÜ, JI, GAO & LI, 2007) ARBOUR & CURRIE, 2015 (Type)
= Crichtonsaurus benxiensis LÜ, JI, GAO & LI, 2007
Genus: Jinyunpelta ZHENG, JIN, AZUMA, WANG, MIYATA & XU, 2018
J. sinensis ZHENG, JIN, AZUMA, WANG, MIYATA & XU, 2018 (Type)
Genus: Oohkotokia PENKALKSI, 2014
O. horneri PENKALKSI, 2014 (Type)
= Genus: Nova PENKALSKI, 1998
= Dyoplosaurus sp GILMORE, 1930
Genus: Zuul ARBOUR & EVANS, 2017
Z. crurivastator ARBOUR & EVANS, 2017 (Type)
Tribe: ANKYLOSAURINI Brown, 1908 emend Arbour & Currie, 2016
Genus: Ankylosaurus BROWN, 1908
A. magniventris BROWN, 1908 (Type)
= Euoplocephalus magniventris (BROWN, 1908) STEEL, 1969
A. sp
Genus: Dongyangopelta CHEN, ZHENG, AZUMA, SHIBATA, LOU, JIN & JIN, 2013
D. yangyanensis CHEN, ZHENG, AZUMA, SHIBATA, LOU, JIN & JIN, 2013 (Type)
Genus: Sauroplites BOHLIN, 1953
S. scutiger BOHLIN, 1953 (Type)
= Sauroplites spiniger MARYANASKA, 1971 (sic)
Tribe: EUOPLOCEPHALINI Penkalski, 2018
Genus: Anodontosaurus C. M. STERNBERG, 1929
= Andontosaurus BODILY, 1969 (sic)
A. lambei C. M. STERNBERG, 1929 (Type)
A. inceptus PENKALSKI, 2018
A. sp
Genus: Dyoplosaurus PARKS, 1924
= Dioplosaurus HAY, 1930 (sic)
= Dyoplasaurus MARYANSKA, 1977 (sic)
D. acutosquameus PARKS, 1924 (Type)
Genus: Euoplocephalus LAMBE, 1910
= Paragenus: Euoplocephalus OLSHEVSKY, 1998
= Eoplocephalius HUNT & LUCAS, 1992 (sic)
= Erroplocephalus NOPCSA, 1928 (sic)
= Euopliocephalus [ANONYMOUS] 1979 (sic)
= Euoplasurus von HUENE, 1956 (sic)
= Euoplocophalus GLUT, 1972 (sic)
= Euoplogy HOU, 1977 (sic)
= Euoplosaurus MALEEV, 1956 (sic)
= Euplocephalus LAMBE, 1920 (sic)
= Europlocephalus C. H. STERNBERG, 1915 (sic)
= Europlosaurus von HUENE, 1929 (sic)
= Europocephalus NOPCSA, 1923 (sic)
= Sterecephalus MALEEV, 1956 (sic)
= Stereocephalus LAMBE, 1902 non ARRIBALZAGA, 1884 (Stereocephalus seriatipennis, Insecta, Staphylinidae)
= Sterocephalus MALEEV, 1956 (sic)
E. tutus (LAMBE, 1902) LAMBE, 1910 (Type)
= Stereocephalus tutus LAMBE, 1902
= Palaeoscincus tutus (LAMBE, 1902) HENNING, 1915
= Eoplocephalus tutus HUNT & LUCAS, 1992 (sic)
= Palaeoscincus asper LAMBE, 1902 (nomen dubium)
= Ankylosaurus tutus (LAMBE, 1902) KUHN, 1964
Platypelta PENKALSKI, 2018
P. coombsi PENKALSKI, 2018 (Type)
Genus: Scolosaurus NOPCSA, 1928
= Scalosaurus MEHL, 1936 (sic)
= Scholosaurus MINELLIL, 1987 (sic)
= Scolasaurus CHEVRAUX, 1980 (sic)
= Skolosaurus von HUENE, 1954 (sic)
Scolosaurus cutleri NOPCSA, 1928 (Type)
S. thronus PENKALSKI, 2018
Genus: Ziapelta ARBOUR, BURNS, SULLIVAN, LUCAS, CANTRELL, FRY & SUAZO, 2014
Z. sanjuanensis ARBOUR, BURNS, SULLIVAN, LUCAS, CANTRELL, FRY & SUAZO, 2014 (Type)
Subfamily: SYRMOSAURINAE Maleev, 1952
= Family: SYRMOSAURIDAE Maleev, 1952
Genus: Akainacephalus WIERSMA & IRMIS, 2018
A. johnsoni WIERSMA & IRMIS, 2018 (Type)
Genus: Maleevus TUMANOVA, 1987
M. disparoserratus (MALEEV, 1952) TUMANOVA, 1987 (Type)
= Syrmosaurus disparoserratus MALEEV, 1952
= Pinacosaurus disparoserratus (MALEEV, 1952) OLSHEVSKY, 1991
= Talarurus disparoserratus (MALEEV, 1952) MARYANASKA, 1977
= Symrosaurus disparoserrata KUHN, 1964 (sic)
= Talarurus disparsoserratus MARYANASKA, 1977 (sic)
Genus: Minotaurasaurus MILES & MILES, 2009
M. ramachandrani MILES & MILES, 2009 (Type)
Genus: Nodocephalosaurus SULLIVAN, 1999
= Nodocephalosaurus SULLIVAN, 1998 (nomen nudum)
N. kirtlandensis SULLIVAN, 1999 (Type)
= Nodocephalosaurus kirtlandensis SULLIVAN, 1998 (nomen nudum)
Genus: Pinacosaurus GILMORE, 1933
= Ninghsiasaurus YOUNG, 1965 (sic)
= Syrmosaurus MALEEV, 1952
= Viminicaudus von HUENE, 1958 (sic)
P. grangeri GILMORE, 1933 (Type)
= Pinacosaurus ninghsieneisYOUNG, 1935
= Syrmosaurus viminocaudus MALEEV, 1952
= Syrmosaurus viminicaudus MALEEV, 1956 (sic)
P. mephistocephalus GODEFROIT, PEREDA-SUBERBIOLA, LI & DONG, 1999
P. sp
Genus: Saichania MARYANSKA, 1977
= Saichana MARYANSKA, 1977 (sic)
S. chulsanensis MARYANSKA, 1977 (Type)
S. sp
Genus: Shanxia BARRETT, HAILU, UPCHURCH & BURTON, 1998
S. tianzhenensis BARRETT, HAILU, UPCHURCH & BURTON, 1998 (Type)
Genus: Talarurus MALEEV, 1952
= Talararus SWINTON, 1970 (sic)
= Talarusus GALTON, 1970 (sic)
T. plicatospineus MALEEV, 1952 (Type)
Genus: Tarchia MARYANSKA, 1977
= Tarcia [Anonymous 1992]
T. kielanae MARYANSKA, 1977 (Type)
T. teresae PENKALSKI & TUMANOVA, 2016
= Tarchia gigantea TUMANOVA, 1977
= Tarcia gigantea [ANONYMOUS, 1992] (sic)
T. tumanovae PARK, LEE, KOBAYASHI, JACOBS, BARSBOLD, LEE, KIM, SONG & POLCYN, 2021
T. sp
Genus: Tianzhenosaurus PANG & CHENG, 1998
T. youngi PANG & CHENG, 1998 (Type)
Genus: Zaraapelta ARBOUR, CURRIE, & BADAMGARAV, 2014
Z. nomadis ARBOUR, CURRIE, & BADAMGARAV, 2014 (Type)
Gen. sp indet.
Subfamily: STEGOPELTINAE Ford, 2000
Genus: Aletopelta FORD & KIRKLAND, 2001
A. coombsi FORD & KIRKLAND, 2001 (Type)
A. sp
Genus: Glyptodontopelta FORD, 2000
G. mimus FORD, 2000 (Type)
Genus: Stegopelta WILLISTON, 1905
S. landerensis WILLISTON, 1905 (Type)
= Nodosaurus landerensis (WILLISTON, 1905) COOMBS, 1978
Subfamily: SHAMOSAURINAE Tumanova, 1983
Genus: Cedarpelta CARPENTER, KIRKLAND, BURGE & BIRD, 2001
= Bilbeyhallorum BURGE, BIRD, McCLELLAND & CICCONETTI, 1999 (nomen nudum)
C. bilbeyhallorum CARPENTER, KIRKLAND, BURGE & BIRD, 2001 (Type)
Genus: Gobisaurus VICKARYOUS, RUSSELL, CURRIE & ZHAO, 2001
G. domoculus VICKARYOUS, RUSSELL, CURRIE & ZHAO, 2001 (Type)
Genus: Shamosaurus TUMANOVA, 1983
= Shamosaurus TUMANOVA, 1981 (nomen nudum)
S. scutatus TUMANOVA, 1983 (Type)
Genus: Tsagantegia TUMANOVA, 1993
T. longicranialis TUMANOVA, 1993 (Type)
Genus: Zhongyuansaurus XU, LU, ZHANG, JIA, HU, ZHANG, WU & JI, 2007
Z. luoyangensis XU, LU, ZHANG, JIA, HU, ZHANG, WU & JI, 2007 (Type)
Gen. sp indet.
Bibliography
Thyreophora in general
GRANDORDER: STEGOSAUROMORPHA COOPER, 1985 (emend)
= INFRAORDER: STEGOSAUROMORPHA COOPER, 1985
= ORDER: THYREOPHORA NOPCSA, 1915 (partim)
= SUBORER: THYREOPHORNIA NOPCSA, 1915 (partim)
= Superfamily: STEGOSAUROIDEA Marsh, 1877 (sensu Hay, 1902)
ORDER: STEGOSAURIA MARSH, 1877
Incertae sedis
= Superfamily: OLIGOSACRALOSAUROIDEA Zhao, 1983
= Superfamily: POLYSACRALOSAUROIDEA Zhao, 1983
Genus: Adratiklit MAIDMENT, RAVEN, OUARHACHE & BARRETT, 2019
A. boulafa MAIDMENT, RAVEN, OUARHACHE & BARRETT, 2019 (Type)
Genus: Amargastegos ULANSKY, 2014a, b (nomen dubium)
A. brevicollus ULANSKY, 2014 a, b (Type)
Genus: Baiyinosaurus LI, MAIDMENT, LI, YOU & GUANGZHAO, 2024
B. baojiensis LI, MAIDMENT, LI, YOU & GUANGZHAO, 2024 (Type)
Genus: Bashanosaurus DAI, LI, MAIDMENT, WEI, ZHOU, HU, MA, WANG, HU & PENG, 2022
B. primitivus DAI, LI, MAIDMENT, WEI, ZHOU, HU, MA, WANG, HU & PENG, 2022 (Type)
Genus: Changdusaurus ZHAO 1986 (nomen nudum)
= Changtusaurus ZHAO, 1983 (nomen nudum)
C. laminaplacodus ZHAO 1986 (Type)
Genus: Chialingosaurus YOUNG 1959
= Chialangosaurus COLBERT, 1961 (sic)
C. kuani YOUNG 1959 (Type)
C. guangyuanensis (listed in LI & CAI, 1998) (nomen nudum)
Genus: Craterosaurus SEELEY 1874 (nomen dubium)
C. pottonensis SEELEY 1874 (Type)
Genus: Diracodon MARSH, 1881 (nomen dubium)
= Diraconodon MARSH, 1887 (sic)
D. laticeps MARSH, 1881 (Type)
= Stegosaurus laticeps (MARSH, 1881) HENNING, 1915
Genus: Dravidosaurus YADAGIRI & AYYASAMI, 1979
D. blanfordi YADGIRI & AYYASAMI, 1979 (Type)
Genus: Eoplophysis ULANSKY, 2014a, b (nomen dubium)
E. vetustus (HUENE, 1910) ULANSKY, 2014a, b. (Type)
= Omosaurus vetustus HUENE, 1910 (nomen dubium)
= Dacentrurus vetustus (HUENE, 1910) HENNING, 1915 (nomen dubium)
= Omosaurus (Dacentrurus) vetustus (HUENE, 1910) HOFFSTETTER, 1957 (nomen dubium)
= Lexovisaurus? vetustus (HUENE, 1910) GALTON & POWELL, 1983 (nomen dubium)
Genus: Ferganastegos ULANSKY, 2014a, b (nomen dubium)
F. callovicus ULANSKY, 2014a, b (Type)
Genu: Isaberrysaura SALGADO, CANUDO, GARRIDO, MORENO-AZANZA, MARTINEZ, CORIA & GASCA, 2017
I. mollensis SALGADO, CANUDO, GARRIDO, MORENO-AZANZA, MARTINEZ, CORIA & GASCA, 2017 (Type)
Genus: Monkonosaurus ZHAO, 1983. (nomen nudum)
M. lawulacus ZHAO, 1983 (Type)
Genus: Saldamosaurus ULANSKY, 2014a, b (nomen dubium)
S. tuvensis ULANSKY, 2014a, b (Type)
Genus: Siamodracon ULANSKY, 2014a, b (nomen dubium)
S. altispinus ULANSKY, 2014a, b (Type)
Genus: Thyreosaurus ZAFATY, OUKASSOU, RIGUETTI, COMPANY, BENDRIOUA, TABUCE, CHARRIERE & PEREDA-SUBERBIOLA, 2024
= Genus: Nova ZAFATY, OUKASSOU, PEREDA-SUBERBIOLA, RIGUETTI, COMPANY, BENDRIOUA, TABUCE & CHARRIERE, 2023
T. atlasicus ZAFATY, OUKASSOU, RIGUETTI, COMPANY, BENDRIOUA, TABUCE, CHARRIERE & PEREDA-SUBERBIOLA, 2024 (Type)
= Species: Nova ZAFATY, OUKASSOU, PEREDA-SUBERBIOLA, RIGUETTI, COMPANY, BENDRIOUA, TABUCE & CHARRIERE, 2023
Genus: Yanbeilong JIA, LI, DONG, SHI, KANG, WANG XU & YOU, 2024
Y. ultimus JIA, LI, DONG, SHI, KANG, WANG XU & YOU, 2024 (Type)
Gen. sp indet.
Family: HUAYANGOSAURIDAE Galton, 1990
Parafamily: HUAYANGOSAURIDAE Galton, 1990 (sensu Olshevsky, 1998)
= Subfamily: HUAYANGOSAURINAE Dong, Tang & Zhou, 1982
= Family: HUOYANGOSAURIDAE Dodson & Dawson, 1991 (sic)
Genus: Chungkingosaurus DONG, ZHOU & ZHANG 1983
= Chunkingosaurus HAUBOLD, 1990 (sic)
= Chungqingosaurus YANG, X.-L. & YANG D., H. 1987 (sic)
C. jiangbeiensis DONG, ZHOU & ZHANG 1983 (Type)
= Chungkingosaurus magnus ULANSKY, 2014a, b (nomen dubium)
= Chungkingosaurus giganticus ULANSKY, 2014a, b (nomen dubium)
Genus: Gigantspinosaurus OUYANG, 1992
G. sichuanensis OUYANG, 1992 (Type)
G. sp
Genus: Huayangosaurus DONG, TANG & ZHOU 1982
= Huangosaurus GALTON, 1986 (sic)
H. taibaii DONG, TANG & ZHOU 1982 (Type)
Genus: Regnosaurus MANTELL, 1848 (nomen dubium)
R. northamptoni MANTELL, 1848 (Type)
= Hylaeosaurus northamptoni(MANTELL, 1848) OWEN, 1858 (nomen dubium)
Genus: Tuojiangosaurus DONG, LI, ZHOU, & ZHANG 1977
= Taojiangosaurus DONG, ZHOU & SHANG, 1983 (sic)
= Tiejiangosaurus DONG, LI, ZHOU & ZHANG, 1977 (sic)
= Tuajiangosaurus DONG, ZHOU & ZHANG 1983 (sic)
= Tueojiangosaurus DONG, LI, ZHOU & ZHANG, 1977 (sic)
= Tuojiangosaurus [ANONYMOUS] 1977 (nomen nudum)
= Tuojiangsaurus YANG, X.-L, & YANG D.-H., 1987 (sic)
= Tuojingosaurus GALTON, 1981 (sic)
= Tuojiongosaurus DONG, LI, ZHOU & ZHANG, 1977 (sic)
T. multispinus DONG, LI, ZHOU & ZHANG 1977 (Type)
Genus: YingshanosaurusZHU, 1994
= Yingshanosaurus ZHOU 1984 (nomen nudum)
= Yinshanosaurus DONG, 1992 (sic)
= Yunshanosaurus (sic)
Y. jichuanensis ZHU, 1994 (Type)
= Yingshanosaurus jichuanensis ZHOU 1984
Bibliography
Family: STEGOSAURIDAE Marsh, 1877
= Family: STEGOSAURIDIDAE Nopcsa, 1917 (sic)
= Family: STEGOSAUROIDAE Marsh, 1877 (sensu Hay, 1930)
= Superfamily: STEGOSAUROIDEA Marsh, 1877 (sensu Hay, 1901 (partim)
Genus: Hesperosaurus CARPENTER, MILES & CLOWARD, 2001
= Genus: Nova CARPENTER & MILES, 1997
H. mjosi CARPENTER, MILES & CLOWARD, 2001 (Type)
= Stegosaurus mjosi (CARPENTER, MILES & CLOWARD, 2001) MAIDMENT, NORMAN, BARRETT & UPCHURCH, 2008
Genus: Jiangjunosaurus JIA, FORSTER, XU & CLARK, 2007
J. junggarensis JIA, FORSTER, XU & CLARK, 2007 (Type)
Genus: Mongolostegus TUMANOVA & ALIFANOV, 2018
M. exspectabilis TUMANOVA & ALIFANOV, 2018 (Type)
= Wuerhosaurus mongoliensis ULANSKY, 2014a, b (nomen dubium)
Genus: Paranthodon NOPCSA, 1929
= Anthodon OWEN, 1876 (partim)
= Paracanthodon von HUENE, 1956 (sic)
P. africanus (BROOM, 1910) COOMBS, 1971 (nomen scruptum) emended OLSHEVSKY, 1978 (Type)
= Palaeoscincus africanus BROOM, 1910
= Anthodon serrarius OWEN, 1876 (partim)
= Paranthodon oweni NOPCSA, 1929
Note: NOPCSA, 1929 erected the genus Paranthodon oweni, but that and BROOM, 1910 genus Palaeoscincus africanus are one and the same specimen. In COOMBS thesis (1971) he corrected the species name, but it wasn’t until OLSHEVSKY, 1978 who officially published the COOMBS correction..
Genus: Nova? ANONYMOUS, 1991
Genus Nova? ANONYMOUS, 1995
Gen. sp indet.
Incertae sedis
Genus: Lexovisaurus non HOFFSTETTER 1957
L.? vetustus or new genus BONEHAM & FORSEY, 1992
Genus: Omosaurus non OWEN, 1875 non LEIDY, 1856
O. phillipsi SEELEY, 1893 (Type) (nomen dubium)
= Dacentrurus phillipsii (SEELEY, 1869) (nomen dubium) HENNING, 1915
Subfamily: DACENTRURINAE Mateus, Maidment & Christiansen, 2009
= Subfamily: OMOSAURINAE Lydekker, 1999
= Family: OMOSAURIDAE Lydekker, 1999
Genus: Dacentrurus LUCAS, 1902
= Dacentrosaurus DONG, 1990 (sic)
= Dacentrurosaurus HENNING, 1925 (sic)
= Omosaurus OWEN, 1875 non LEIDY, 1856 (Omosaurus perplexus, Eucrocopoda, Parasuchia)
= Osmosaurus GALTON, 1980 (sic)
D. armatus (OWEN, 1875) LUCAS, 1902 (Type)
= Omosaurus armatus OWEN, 1875
= Stegosaurus armatus (OWEN, 1875) LYDEKKER, 1890, non MARSH, 1877
= Omosaurus hastiger OWEN 1877
= Omosaurus lennieri NOPCSA, 1911
= Dacentrurus lennieri (NOPCSA, 1911) HENNING, 1915
= Astrodon pusillus LAPPARENT & ZBYSEWSKI 1957
D. sp
Genus: Miragaia MATEUS, MAIDMENT & CHRISTIANSEN, 2009
= Alcovasaurus GALTON, & CARPENTER, 2016
= Natronasaurus ULANSKY, 2014a (nomen nudum)
M. longispinus (GILMORE, 1914) COSTA & MATEUS, 2019
= Stegosaurus longispinus GILMORE, 1914
= Stegosaurus altispinus GILMORE 1914 (sic)
= Kentrosaurus? longispinus (GILMORE, 1914) OLSHEVSKY & FORD, 1993
= Natronasaurus longispinus (GILMORE, 1914) ULANSKY, 2014a (nomen nudum)
= Alcovasaurus longispinus (GILMORE, 1914) GALTON, & CARPENTER, 2016
M. longicollum MATEUS, MAIDMENT & CHRISTIANSEN, 2009 (Type)
Gen. sp indet.
Subfamily: KENTROSAURINAE Olshevsky & Ford, 1993
= Family: KENTROSAURIDAE Olshevsky & Ford, 1993
Genus: Kentrosaurus HENNING 1915
= Centrurosaurus NOPSCA, 1917 (sic)
= Doryphorosaurus NOPCSA, 1916
= Kentrurosaurus HENNING, 1916
K. aethiopicus HENNING 1915 (Type)
= Doryphorosaurus aethiopicus (HENNING, 1915) NOPCSA, 1916
= Kentrurosaurus aethiopicus (HENNING, 1915) HENNING, 1916
Genus: Lexovisaurus HOFFSTETTER 1957
= Lexousaurus DONG, CHANG, LI & SHOUT 1978 (sic)
L. durobrivensis (HULKE, 1887) HOFFSTETTER 1957 (Type)
= Omosaurus durobrivensis HULKE, 1887
= Stegosaurus durobrivensis (HULKE, 1887) HULKE, 1887
= Dacentrurus durobrivensis (HULKE, 1887) HENNING, 1915
= Omosaurus leedsi SEELEY 1901
L. sp
Genus: Loricatosaurus MAIDMENT, NORMAN, BARRETT & UPCHURCH, 2008
L. priscus (NOPCSA, 1911) MAIDMENT, NORMAN, BARRETT & UPCHURCH, 2008 (Type)
= Stegosaurus priscus NOPCSA 1911
= Lexovisaurus priscus (NOPCSA, 1911) KUHN, 1964
Subfamily: STEGOSAURINAE Marsh, 1877 (sensu ABEL, 1919)
= Family: HYPSIRHOPHIDAE Cope, 1898
Genus: Hypsirophus COPE, 1878 (nomen dubium)
= Hypsirhophus COPE, 1878 (sic)
= Hypsirrhopus ovn HUENE, 1909 (sic)
H. discurus COPE, 1878 (Type) (nomen dubium)
= Stegosaurus discurus (COPE, 1878) HENNING, 1915 (nomen dubium)
= Hypsirophus seeleyanus COPE 1879 (nomen nudum)
= Stegosaurus seeleyanus (COPE 1879) HENNING, 1915 (nomen nudum)
Genus: Stegosaurus MARSH, 1877
= Sregosaurus GLUT, 1972 (sic)
= Stegasaurus [ANOMYMOUS] 1980 (sic)
S. armatus MARSH 1877 (Type)
S. stenops MARSH, 1887
= Diracodon stenops (MARSH, 1887) BAKKER, 1986
S. ungulatus MARSH 1879
= Stegosaurus angulatus SAUVAGE, 1880 (sic)
= Stegosaurus duplex MARSH 1887
S?affinis MARSH 1881 (nomen nudum)
S. sulcatus MARSH 1887
S. Nova? BAKKER
S. sp
Genus: Wuerhosaurus DONG 1973
= Wuherosaurus SERENO, 1986 (sic)
W. homheni DONG 1973 (Type)
= Stegosaurus homheni (DONG, 1973) MAIDMENT, NORMAN, BARRETT & UPCHURCH, 2008
W. ordosensis DONG, 1993
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Zheng, W., Jin, X., Azuma, Y., Wang, Q, Miyata, K, and Xu, X., 2018, The most basal ankylosaurine dinosaur from the Albian-Cenomanian of China, with implications for the evolution of the tail club: Scientific Reports 8:3711; DOI: 10.1038/s41598-018-21924-7,17pp.
Zhou, S., 1983, A nearly complete Skeleton of Stegosaur from Middle Jurassic of Dashanpu, Zigong, Sichuan: Journal of Chengdu College of Geology, supplement 1, p. 26-32.
Zhou, S., 1984, The middle Jurassic Dinosaurian fauna from Dashanpu, Zigong, Sichuan: Sichuan Scientific and Technological Publishing House, p. 1-52.
Zhu, S., 1994, Record of a fossil stegosaur from Yingshan in the Sichuan Basin: Sichuan Cultural Relics, S1, p. 8-14.
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https://dokumen.pub/the-princeton-field-guide-to-dinosaurs-second-edition-secondnbsped-9781400883141.html
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The Princeton Field Guide to Dinosaurs: Second Edition [Second ed.] 9781400883141
|
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The best-selling Princeton Field Guide to Dinosaurs remains the must-have book for anyone who loves dinosaurs, from amat...
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dokumen.pub
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https://dokumen.pub/the-princeton-field-guide-to-dinosaurs-second-edition-secondnbsped-9781400883141.html
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Table of contents :
CONTENTS
Preface
Acknowledgments
Introduction
History of Discovery and Research
What Is a Dinosaur?
Dating Dinosaurs
The Evolution of Dinosaurs and Their World
Extinction
After the Age of Dinosaurs
Biology
General Anatomy
Skin, Feathers, and Color
Respiration and Circulation
Digestive Tracts
Senses
Vocalization
Disease and Pathologies
Behavior
Brains, Nerves, and Intelligence
Social Activities
Reproduction
Growth
Energetics
Gigantism
Mesozoic Oxygen
The Evolution—and Loss—of Avian Flight
Dinosaur Safari
If Dinosaurs Had Survived
Dinosaur Conservation
Where Dinosaurs Are Found
Using the Group and Species Descriptions
Group and Species Accounts
Dinosaurs
Theropods
Sauropodomorphs
Ornithischians
Additional Reading
Index: Dinosaur Taxa
Formations
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https://journals.plos.org/plosone/article%3Fid%3D10.1371/journal.pone.0080405
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The Basal Nodosaurid Ankylosaur Europelta carbonensis n. gen., n. sp. from the Lower Cretaceous (Lower Albian) Escucha Formation of Northeastern Spain
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[
"Vertebrae",
"Skull",
"Teeth",
"Dinosaurs",
"Cretaceous period",
"Ribs",
"Spine",
"Ischium"
] | null |
[
"Mark A. Loewen",
"Eduardo Espílez",
"Luis Mampel",
"Jelle P. Wiersma",
"James I. Kirkland",
"Luis Alcalá"
] | null |
Nodosaurids are poorly known from the Lower Cretaceous of Europe. Two associated ankylosaur skeletons excavated from the lower Albian carbonaceous member of the Escucha Formation near Ariño in northeastern Teruel, Spain reveal nearly all the diagnostic recognized character that define nodosaurid ankylosaurs. These new specimens comprise a new genus and species of nodosaurid ankylosaur and represent the single most complete taxon of ankylosaur from the Cretaceous of Europe. These two specimens were examined and compared to all other known ankylosaurs. Comparisons of these specimens document that Europelta carbonensis n. gen., n. sp. is a nodosaur and is the sister taxon to the Late Cretaceous nodosaurids Anoplosaurus, Hungarosaurus, and Struthiosaurus, defining a monophyletic clade of European nodosaurids– the Struthiosaurinae.
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Description and Comparisons
Skull
The skull (AR-1-544/10) was lying on its dorsal surface and is moderately well preserved although distorted through compaction (Fig. 7). The palate is crushed in toward the skull roof, resulting in the medial rotation of both maxillae with the posterior teeth displaced into the posterior palate. The sheet-like palatal bones are highly fragmented. The braincase is crushed along the plane of the cranial nerve openings and the fenestra ovalis completely obscures them. Unexpectedly, the right quadrate (Fig. 8 H–J) and associated portion of the palate was dislodged from the skull and subsequently crushed across the ventral side of the basicranium. This gives the impression that these bones had been expelled from inside the skull prior to compaction. Both the left and right nasals were separated from the skull and the premaxillae (whereas possibly present upon discovery) have not been identified.
The skull has a minimum length of 370.3 mm from the anterior end of the maxillae to the rear margin of the squamosals. The skull has a maximum width of 299.1 mm at the orbits and narrows to 203.7 mm at the posterior end of the skull at the squamosals, giving the skull the “pear-shaped” dorsal profile characteristic of derived nodosaurids [70], [71]. Although tapering posteriorly, there is no distinct post-temporal notch as in polacanthids and other nodosaurids [63].
The maxillae (Fig. 7 D–F) are irregularly sculptured externally with a flattened, horizontally oriented buccal recesses that are inset approximately 2 cm. The anterior margin of the maxilla appears to form the posterior margin of a relatively simple naris relative to derived nodosaurids and ankylosaurids. Medially, there is no evidence that the maxilla formed a portion of a secondary palate. The tooth row was arched ventrally with an estimated 22–25 alveoli increasing in size posteriorly as in Edmontonia [72]. In ventral orientation, the tooth rows are only moderately deflected medially, such that the palate would not have had a pronounced hourglass appearance typical of derived nodosaurs such as Pawpawsaurus, Edmontonia, and Panoplosaurus [73]–[75]. However, it is not dissimilar from that of the primitive nodosaurid Silvisaurus [76], [77].
The nasals (AR-1-133/10, AR-1-639/10) are relatively large and subrectangular, tapering somewhat anteriorly (Fig. 8 A–D). Both nasals extend laterally from their relatively straight, unfused midline suture before flexing down to a sutural contact with the maxillae that extends for most of their length. When rearticulated onto the skull, they appear to fit well, despite the skull's distortion. Most ankylosaurs have fused nasals except the nodosaurids Silvisaurus [76], [77] and Niobrarasaurus [78], although the nasals are unknown in European nodosaurids [24], [32], [33]. A distinct tongue-like process projects from the nasal's posterior margin and would have overlapped the frontals. The external surface is lightly textured and the internal surface is relatively smooth, suggesting the narial passage was large and simple, rather than convolute as in derived nodosaurids and ankylosaurids [79], [80].
The orbits are somewhat crushed and the sutures of the bones surrounding them are obscured by fusion. The orbits are subrectangular in shape, are slightly more elongate anteoposteriorly and are directed anterolaterally. The prominent and evenly rounded suborbital horn is formed mostly from the quadratojugal posterior to the ventral margin of the orbit, as in most derived ankylosaurs [81], [82] and unlike that in polacanthids such as Mymoorapelta, Gargoyleosaurus, and Gastonia where the suborbital horn is below the orbit and is formed exclusively by the jugal [83]–[85]. The suborbital horn appears to be unornamented and hides the head of the quadrate in lateral view.
The lateral wall of the skull extends posteriorly behind orbit with a dorsoventally wide posterior notch, such that the lower temporal opening is just visible in lateral view. There is no lateral wall of skull behind the orbits in polacanthids [70], [81] and most nodosaurids other than Peloroplites [86], Silvisaurus [76], Struthiosaurus transylvanicus [22], [23] and one specimen from the Dinosaur Park Formation assigned to Edmontonia (ROM 1215) [88], although in these taxa the lower temporal opening is still visible in lateral view as in Europelta. The lower temporal opening is completely obscured in lateral view in Cedarpelta [84], [86], Shamosaurus [89]–[91], Gobisaurus, [92] Zhongyuansaurus [93] and all derived ankylosaurids.
Although the palate is fragmented and crushed along the internal surface of the skull roof, the fragments of the vomer suggest it did not extend ventrally to the level of the tooth row. Additionally, the broad sheet-like pterygoids appear to have been flexed nearly dorsally against the anterior portion of the basicranium as in nodosaurids and not like the open transversely oriented pterygoids characteristic of ankylosaurids or polacanthids [94].
The posterolateral margin of the pterygoid is fully fused to the quadrate. There is a sutural contact between the straight, nearly vertical quadrates and the quadratojugal laterally. The quadrates are wide transversely and thin rostrocaudally as compared to the mediolaterally narrower quadrates of other ankylosaurs [82]. The contact with the squamosal is also transversely wide, unlike the narrow, rounded contact seen in many ankylosaurs such as Mymoorapelta (Kirkland, pers. obs.) and Cedarpelta [63], [86]. The mandibular articulation is proportionally wider than in any other ankylosaur examined as a part of this study and the medial condyle larger than the lateral condyle. The ratio of mediolateral quadrate width to dorsoventral quadrate length is 0.77 (94 mm/122 mm). The anteropostior length of the quadrate condyle is 31 mm. There is no fusion between the quadrates and the paroccipital processes.
Vertical compaction has obscured the posterior view of the skull, in particular the foramen magnum and the supraoccipital. However, even with compaction it is apparent that in occipital view the skull was subrectangular and wider than tall as in Gargoyleosaurus, Gastonia, and most other derived anklylosaurs, and unlike the narrow, highly arched occipital region of Struthiosaurus [22]. The paroccipital processes extend horizontally lateral to the foramen magnum and then flare dorsoventrally by approximately 100% of their minimum widths. They angle posteriorly at about 30 degrees when viewed ventrally (Fig. 7 F). In morphology and orientation, they are most similar to those in Gargoyleosaurus [95] although ventral twisting is not present. In most other ankylosaurs, the paroccipital processes extend straight laterally [81], [96] or may be flexed ventrally as in Gastonia [83]. A triangular wedge of bone of unknown identity is fused to the anterior ventrolateral margin of the paroccipital, separating it from the quadrate.
The subspherical occipital condyle (Fig. 7 B, F) has a width of 59.4 mm and height of 46.5 mm and lacks a distinct neck to separate it from the rest of the basicranium. Although no cranial sutures are visible, the occipital condyle does appear to be composed exclusively of the basioccipital. It is similar in overall morphology to that of the basal ankylosaurid Cedarpelta [88] except that the occipital condyle angles somewhat ventrally, but not as much as in more derived nodosaurids [71], [82]. The ventral surface of the relatively elongate basioccipital is broadly convex. Again, as in Cedarpelta [88], there are no distinct, separate basal tubera between the basioccipital and the short basisphenoid, but instead there is a prominent transverse flange extending across the ventral surface of the basicranium along the line of this suture. The pterygoid processes appear to be short, but are completely obscured by crushed pterygoids bone fragments that wall off the anterior part of the braincase as in most nodosaurids.
The skull roof (figs. 7 C, 9 A) is roughened texturally by remodeling of the bone surface as in Cedarpelta, the nodosaurids Sauropelta and Peloroplites, and the shamosaurine-grade ankylosaurids Shamosaurus and Gobisaurus [81], [86], [88]. Europelta differs from these specimens in that some of the margins of the scale impressions on the skull roof are visible, as seen in Edmontonia, Panoplosaurus and Struthiosaurus [22], [77]. These scale margins are represented by shallow grooves that are difficult to see relative to the textured surface of the skull and the cracks in the bone due to compaction. These grooves are particularly evident along the lateral margins of the skull roof above the orbit. An extensive median scale appears to have covered much of the central portion of the skull between and posterior to the orbits on the frontals and parietals as other nodosaurids [63], [82]. There does not appear to be any distinct nuchal ornamentation. The skull is thickened above the orbit, but there is not a distinct supraorbital boss, a condition similar to Peloroplites, Cedarpelta, Shamosaurus, and Gobisaurus [86], [88]–[90], [92]. Narrow grooves along the margin of the skull in this area above the orbits suggest that a particularly robust pair of scales were present in this area as indicated by a deep groove bisecting this ornamented area directly above the orbit. Weak grooves delineate a small scale without underlying ornamentation separating the posterior supraorbital scale from the squamosal horn forming the posteriolateral margin of the skull roof. The squamosal horn is ornamented by narrow grooves radiating from its apex onto the skull roof. Grooves on the anterolateral sides of the fronto-parietal scale appear to delineate two scales between the anterior supraorbital scales. Unfortunately, no distinctive scale boundaries are recognizable on the nasals, although the dorsal surfaces of the nasals are textured. Several elongate scales rimmed the lateral raised margin around the orbit.
In dorsal view, the posterior margin of the skull is concave, whereas it is nearly straight or convex in all other nodosaurids. This reflects the posterior angulation of the paraoccipital processes and the squamosal horns. Interestingly, the occipital condyle is barely visible, though not completely obscured in dorsal view. There is no evidence of any distinct nuchal sculpturing. The skull roof is relatively flat but a slight dome may have been present prior to crushing. However, it is clear that the skull roof is not as highly domed as in many other nodosaurids, such as Struthiosaurus [22], [26].
Attempts were made to image the skull using X-ray photography and CT scanning. The abundance of pyrite present in the skull (Fig. 4E) presents a strong limitation in the use of these techniques as pyrite is opaque to X-rays.
Mandible
A small dentary fragment extending for only four complete alveolae (AR-1-133/10) was preserved from the holotype skeleton (Fig. 8 E–G). However, a robust left dentary and splenial are preserved together (AR-1-3698/31) from the paratype specimen (Fig. 10 A–E). The splenial is not in its posteriomedial position relative to the dentary, but is fused across the posterior portion of the tooth row transversely. Additionally, an isolated left angular with a distinct highly sculptured scale along its ventral margin (AR-1-2945/31), was recovered (Fig. 10 F, G).
The dentary is 184.7 mm long with a minimum of 21 tooth positions, with no possibility of more than two unpreserved alveoli as determined by the position of the suture with the angular and surangular. As with the maxillary teeth, the alveoli are more than twice as large posteriorly. There is only 1.5 cm between the anteriormost alveoli and the symphysis, suggesting that there may have been premaxillary teeth as at least nine anterior teeth would have been positioned to oppose the premaxilla. The primitive ankylosaurs Sarcolestes [34], [98], Gargoyleosaurus, [85], Silvisaurus [76], Animantarx [97], Sauropelta [99], Anoplosaurus [17], Hungarosaurus [33] and Struthiosaurus [22] have a short anterior diastema, and thus a narrow predentary, whereas this diastema is longer in ankylosaurs with wide predentaries. However, the symphysis in Europelta is robust and dorsoventrally deeper (45.0 mm deep and 29.00 mm across) than in ankylosaurs [82], and is most similar to the deep symphysis of Hungarosaurus [32], further suggesting a reduced predentary with a rudimentary ventral process. The symphysis is marked by two deep anteroposteriorly directed grooves. A row of foramina extends posteriorly on the lateral surface of the dentary from just dorsal to the buccal recess to the notch for the surangular, whereas nutritive foraminae are not clearly visible ventral to the alveolae on the medial side of the dentary as in other ankylosaurs. The recessed tooth row is deflected medially and forms a convex arch in lateral view. The dentary of Hungarosaurus is deeper dorsoventrally than that of Europelta [33].
The splenial (Fig. 10 A-D) is a thin bone with a convex ventral margin 156.6 mm long that contacts the angular. It has the appearance of an obtuse triangle in medial view. There is large, well-developed intermandibular foramen (7 mm long and 5.3 mm wide) 50 mm from its anterior end.
The angular (Fig. 10 F, G) has a maximum length of 175 mm. The lateral margin is highly rugose, because the bone is textured and remodeled to support a large scale, extending about 10–12 mm ventral to the ventral margin of the angular for most of its length. A distinct ridge marks the dorsal limit of the mandibular ornament medially, where it is in contact with the ventral margin of the splenial. Dorsal to this contact the bone is smooth. The ventral extent of the textured bone supporting the mandibular scale is similar to that observed in ankylosaurids such as Euoplocephalus [95] and Minataurasaurus [100], rather than the more lateral orientation found in Gargoyleosaurus [93] and in nodosaurids like Sauropelta [99] and Panoplosaurus [101].
Teeth
A large number of teeth are preserved from both the holotype AR-1/10 (20+) and the paratype AR-1/31 (15+) although many have drifted away from the alvaeolae. We assume that the teeth associated with the holotype pertain to the maxilla (several are preserved in the palate and in the maxilla) and those of the paratype pertain to the dentary (several are preserved in the dentary). In general, the cutting surfaces of the teeth are not well preserved, but a few exceptions exist. Wear facets were not observed on any of the teeth. The roots for both dentary and maxillary teeth are swollen lingually, are three to four times the length of the crowns, and are subquadrate in cross-section. One small tooth (AR-1-343/10) is more highly asymmetrical mesiodistally and may represent a premaxillary tooth (Fig. 11 L, M).
The isolated maxillary teeth (Fig. 11 A–K, N–FF) have a weakly developed labial cingulum and a strongly developed lingual cingulum. The best preserved right tooth AR-1-324/10 is 11.50 mm wide, 9.99 mm tall with seven to eight mesial denticles and five to six distal denticles (Fig. 11 A–E). A large right tooth AR-1-564/10 is 17.23 mm wide and 12.95 mm tall with eight to nine mesial denticles and ∼six to seven distal denticles (Fig. 11 V–Z).
The isolated dentary teeth (Fig. 11 GG–FFF) are identical to the maxillary teeth and have a weak lingual cingulum and a strongly developed labial cingulum. The best preserved tooth AR-1-3700/31 is 14.03 mm wide and 12.69 mm tall with eight to nine mesial denticles and six to seven distal denticles (Fig. 11 LL–PP). The largest dentary tooth AR-1-3650/31 is 16.58 mm wide and 13.50 mm tall (Fig. 11 GG–KK).
With their relatively large size and well-developed cingula, the teeth of Europelta are most comparable to those of other nodosaurids [72]. They similar to the teeth of Cedarpelta, Sauropelta [34], [97], [102], Edmontonia and Panoplosaurus [72], but are not as high crowned as in the Jurassic ankylosaurs Sarcolestes and Priodontognathus [103], the Jurassic polacanthids Gargoyleosaurus [93] and Mymoorapelta (Kirkland, pers. obs.), the nodosaurids Peloroplites [84] or Hungarosaurus [33]. Additionally, the large teeth of Gobisaurus are more inflated labiolingually than in Europelta and other ankylosaurs. The teeth of Gastonia and putative Polacanthus teeth are also inflated, but are smaller proportionally [83], [103]. The teeth of Europelta differ from an isolated tooth from the Cenomanian of France which is about half the size, and proportionally is longer mesiodistally with more deeply divided denticles forming ridges on the labiolingual surfaces of the tooth [104]. Likewise, lower Cenomanian teeth assigned to “Acanthopholis” have more deeply divided denticles in what is a proportionally taller tooth [17]. The teeth of Struthiosaurus languedocensis [31] from the lower Campanian of France also differ in size and in having longer, lower tooth crowns.
Axial skeleton
There are numerous ribs and vertebrae preserved from the holotype (AR-1/10) and the paratype specimen (AR-1/31). Vertebral measurements are presented in Table 1.
The complete atlas (AR-1-649/10) from the holotype has a total width of 195.6 mm (Fig. 12 A–F). The neural arch is divided dorsally with the left side fused to the centrum and the right side unattached. The anterior face of the atlantal intercentrum is 73.7 mm wide by 71.7 mm tall and its posterior face is 99.9 mm wide by 61.2 mm tall with a length of 62.0 mm. The axis is not present in either associated skeleton.
There are five post-axis cervical vertebrae (AR-1-431/10, 449, 533, 637, 650) preserved from the holotype skeleton (Fig. 12 I– II) and five from the paratype skeleton; of which four are illustrated (AR-1-3586/31, 3632, 3671, and 3676) (Fig. 13). Overall, they are typical of most other described ankylosaur cervical vertebrae. The centra are amphicoelus, wider than tall, anterorposteriorly short, and medially constricted. Anterior and mid-cervical vertebrae have the anterior faces of the centra dorsally elevated relative to the posterior faces. This is in contrast to the posterior cervical centra which have horizontally aligned faces. The ventral sides of the anterior centra are characterized by two anteroposteriorly-oriented paired fossae separated by a low keel (Figs.12, N, T, Y, EE, II, 13, F), as observed in the primitive nodosaurid Animantarx [97]. The dorsal ends of the neural spines are expanded transversely. AR-1-638/10 may either be the last cervical vertebra or the first dorsal vertebra based on the position of the parapophyses.
There are two complete cervical ribs preserved for the holotype. AR-1-450/10 is a relatively anterior cervical rib (Fig. 12 G, H) and AR-1-4452/10 is a posterior cervical rib. There is no evidence of fusion of cervical ribs to the cervical vertebrae as in the ankylosaurid Saichania [105], [106] or Ankylosaurus [107]. The cervical ribs are Y-shaped overall and much like the cervical ribs of other ankylosaurs such as Silvisaurus [76], [78], [82].
Several amphiplatan to amphicoelus dorsal vertebra are preserved: eight for the holotype AR-1/10 and nine for the paratype AR-1/31. The diapophyses originate at the level of the post-zygopophyses at the dorsal extent of the neural canal. The more anterior vertebrae have large cylindrical amphiplatan centra which lack a constricted ventral keel with circular neural canals and fused ribs (AR-1-448/10, 478, and 535). The broad transverse processes are T-shaped in cross-section and angled dorsally, unlike the laterally directed transverse processes in Polacanthus [10], [38]. Two dorsal vertebrae from the holotype appear to be pathological with the centra overgrown by about 0.5 cm of lumpy reactive bone (Figs. 14, G–K, W–BB). One of these pathologic vertebrae (AR-1-535/10) has fused ribs (Fig. 14 G–K) although the other (AR-1-430/10) does not (Fig. 14 W–BB). Two additional dorsal vertebrae (AR-1-478/10, 448) with fused ribs are not pathologic (Fig. 14 L–V). More posterior dorsal vertebrae have shorter, taller, more medially constricted centra, laterally compressed neural canals, more dorsally directed transverse processes, and lack fused ribs (AR-1-155/10, 322, and 556). The neural spines are thin and rectangular with narrowly expanded dorsal ends as in Sauropelta [99]. The neural spines are oriented dorsally as opposed to the posteriorly inclined neural spines of some other ankylosaurs such as Sauropelta [97]. None of the paratype vertebrae (AR- 1-3489/31, 3633, 3662, 3672, 3673, 3674, 3675, 3677 and 3704) have fused ribs (Fig. 15), suggesting that this character is ontogenetic because the paratype AR-1/31 represents a somewhat smaller (and presumably younger) individual than the holotype AR-1/10. More expanded neural spines are present in Shamosaurus [91].
There are a number of rib fragments preserved with AR-1/10, but there are only three (AR-1-331/10, 333, 476) relatively complete ribs (Fig. 16). As with most other ankylosaurs, the ribs are sharply arched and L-shaped in cross-section proximally in anterior ribs and broadly arched and T-shaped in cross-section proximally in more posterior ribs.
The sacrum is not preserved in AR-1/10 other than an anteriormost centrum (AR-1-154/10) of the synscacrum (Fig. 17 W, X). However, for the paratype, AR-1-3466/31, there is a largely complete but fragmented synsacrum (Fig. 17 A–V) that includes an interpreted anteriormost synsacral centrum (AR-1-3451/31), more of the anterior synsacrum composed of two dorsal centra (AR-1-3450/31), four sacral vertebrae with the sacral ribs from the left side (AR-1-3446/31), two sacral ribs from the right side (AR-1-3452/31, 3460), and one caudosacral vertebra (AR-1-3512/31). Given that at least one intermediate and one anterior fused synsacral dorsal vertebra are missing, the vertebral formula for the synsacrum would be five or more dorsosacral vertebrae, four sacral vertebrae, and one sacrocaudal vertebra. The entire synsacrum would have been over 50 cm long and measures about 44 cm across the sacral ribs. The middle section of the preserved dorsal synsacrum thins anteriorly from about 7 cm wide to about 5.5 cm wide. It then expands again anteriorly as indicated by the anteriormost centrum of the synsacrum. This differs from the sacrum of Euoplocephalus [108] and Saichania [106] in which each centrum making up the synsacrum is constricted medially. The sacrum is distinctive in being more strongly arched anteroposteriorly than other described ankylosaur sacra. The neural spines are dorsoventrally shorter than the height of the centra and are fused into a vertical sheet of bone along the length of the sacrum. The caudosacral neural spine is longer and unexpanded, transitional in form between the sacral neural spines and those of the proximal caudal vertebrae. The neural spines are broken off the anterior end of the synsacrum. The ventral side of the sacrum and anterior synsacrum is longitudinally depressed. The distal ends of the sacral ribs are expanded and the most robust medial sacral rib is about 50% taller (9.4 cm) than wide (6 cm) at its attachment with the ilium. There is no sign of expansion of the dorsal termination of the neural spine on the sacrocaudal vertebra. Additionally, the caudal rib is reduced compared to the sacral ribs.
The sacrum of Struthiosaurus languedocensis [31] is similar overall, but based on the description is not so strongly anteroposteriorly arched as in Europelta. Similarly, the sacrum of Hungarosaurus, as exhibited at the Hungarian Natural History Museum, appears to be moderately arched. The moderate angulation of the faces of the sacral centra (somewhat wedge-shaped in lateral view) in Anoplosaurus [17] indicates that a moderately arched sacram may have been present in this taxon as well. Among North American nodosaurids, we have observed only a moderate anteroposteriorly arching of the synsacrum of Silvisaurus, which appears to be restricted to the posterior part of the sacrum and two sacrocaudals. In other ankylosaurs, the downward flexure of the tail from the hips is taken up in the proximal caudal vertebrae as in Mymoorapelta [84], [109] and Euoplocephalus [70], [82].
Only three proximal caudal vertebrae (AR-1-562/10, 635, 636) are present (Fig. 18 A–F, J–O, V–AA). The proximal-most caudal vertebrae are not preserved for the holotype. The preserved vertebrae probably represent caudal vertebrae positions in the interval of about 3–7. The centra are anteroposteriorly shorter than dorsoventrally tall and somewhat wedge-shaped in anterior and posterior views. The posterior chevron facets are well developed. The neural spines are inclined posteriorly and the dorsal ends of the neural spines are only slightly expanded transversely as in Gargoyleosaurus [95] and some other ankylosaurs such as Cedarpelta [86], Edmontonia [110], Hungarosaurus [32] and Euoplocephalus [70], [82]. The neural spines are strongly expanded in most polacanthids such as Mymoorapelta [84], [109], Gastonia [83], and Polacanthus [10], and some North American nodosaurids such as Sauropelta [99], and Silvisaurus [76]. The neural spine of AR-1-562/10 is broken, erroneously giving it the appearance of being strongly inclined posteriorly. The caudal ribs (transverse processes) in Europelta originate high on the sides of the centrum and angle ventrally proximal to flexing laterally, giving them a dorsally concave profile in anterior view like Hungarosaurus, Struthiosaurus, and Peloroplites, and unlike the ventrally flexed caudal ribs of many polacanthids [10], [84], [109] and the caudal vertebra assigned to “Acanthopholis” [17] or straight caudal ribs of Gargoyleosaurus [95], Cedarpelta, Peloroplites [86], and Edmontonia [87]. The proximal caudal ribs of Hylaeosaurus differ in being swept back posteriorly [111]. The lateral terminations of the caudal ribs do not expand dorsoventrally as they do in Peloroplites [86] and Struthiosaurus, which actually appear to bifurcate [25], [26].
Additionally, there are four chevrons preserved from about the same region of the tail (AR-1-560/10, 561, 569, and 4451) of which three are illustrated (Fig. 18 G–I, P–U). The proximal chevrons are approximately as long as the neural spines as in most other ankylosaurs. They are relatively straight and expanded into teardrop shapes distally in lateral view. Unlike in many ankylosaurs, there is no fusion of proximal chevrons to their respective caudal vertebrae as in Pinacosaurus and Saichania [105], [106], Ankylosaurus [107], [112], and Edmontonia (ROM 1215) [87].
Several more distal caudal vertebrae are preserved in the paratype. The two most proximal of these (AR-1-3348/31, AR-1-3717/31) have centra of nearly equal height, width, and length, with a ventral groove, and caudal ribs shorter than the diameter of the centrum that extend laterally and angle posteriorly (Fig. 19 A–J). The chevron facets are well developed with the posterior facets more strongly developed than the anterior facets. The neural spines are not developed and the zygapophyeses only extend a short distance beyond the anterior and posterior margins of the centra. These vertebrae are interpreted to represent mid-caudal vertebrae. Two more distal mid-caudal vertebrae (AR-1-3616/31, AR-1-3716/31) are similar in morphology except that the caudal ribs are reduced to anteroposteriorly directed ridges on the lateral margins of the centra (Fig. 19 K–N). Their neural spines incline posteriorly, merging with the postzygapophyses as posterior processes extending laterally past the faces of the centra to overlie and articulate between the paired prezygapophyses of the immediatly distal vertebra. This morphology is retained in the distal caudal vertebra. More distally, as in AR-1- 2950/31, 3206, 3243, 3265, 3478, and 3615, the caudal ribs are lost and the centra become more elongate (Fig. 19 O–FF). Unlike many ankylosaurs, the faces of the centra maintain a well-rounded to heart-shaped surface distally down the caudal series [82]. For many of these vertebrae, ventrally anteroposteriorly elongated skid-shaped (inverted T) chevrons are fused to the posterior chevron facets. Fusion of distal chevrons to their respective vertebrae is widespread among ankylosaurs [84], [106], [110] although it is not present in some, such as Nodosaurus [113]. One pair of distal caudal vertebrae is fused by their mutually shared chevron (Fig. 19 GG–II) such as has been documented in Mymoorapelta [84]. The most distal four caudal vertebrae (Fig. 19 JJ–LL) and their chevrons are fused together in AR-1-3204/31 to form a tapering, terminal rod of bone at the end of the tail somewhat similar to that of Sauropelta [71].
Pectoral Girdle
Parts of the right scapulocoracoid are preserved. A portion of the distal scapular blade (AR-1-429/10) is preserved with a portion of the distal ventral margin missing with a curved section broken away. There is no evidence of any distal expansion of the scapular blade as in many nodosaurids [94].
The coracoid (AR-1-657/10) is preserved with only the most proximal portion of the scapula fused on (Fig. 20 D–H). It appears to have been sheared off just dorsal to the suture between the coracoid and the scapula, perhaps in the process of removing the overlying coal seam. The coracoid is relatively equidimensional (201.3 mm long by 186.5 mm tall) relative to the elongate coracoids characteristic of many other nodosaurids [114] such as Peleroplites [86], Texasites [77], [115], and Animantarx [97]. The medial surface is concave and the lateral surface is convex giving it a bowl-shaped appearance. The ventral margin is evenly convex as in many polacanthids and nodosaurids and there is no anteroventral process as in all ankylosaurids, including Shamosaurus [91], [94]. The articular surface of the ventrally directed glenoid is wide, bounded by a flange that extends beyond the medial surface of the coracoid.
Both xiphisternal plates are preserved (Figure 20I–L). The best preserved xiphisternal is approximately 350 mm long. They appear to be arcuate flat bones. Xiphisternal plates are only known in a few nodosaurids, but those of Europelta, whereas similar in overall shape to other nodosaurid xiphisterna, are not fenestrate or scalloped along their margins as in North American nodosaurids for which they are known [82], [87], [116].
Forelimb
Parts of both humeri are preserved. The right humerus (AR-1-655/10) is represented by the proximal end (Fig. 21 A–D). It is 249.2 mm wide with a well-developed proximal head 91.9 mm wide that extends onto the posterior side of the humerus. Distinct notches separate both the laterally directed deltopectoral crest as in nodosaurids such as Sauropelta [70], [71], [99] and the internal tuberosity from the humeral head. The deltopectoral crest extends lateraly from the humerus and is not flexed anteriorly as in polacanthids and ankylosaurids [94].
The left humerus (AR-1-327/10) is represented by a midshaft for which both the proximal and distal ends appear to have rotted off and the core of the shaft has rotted away (Fig. 21 E–H). The shaft is deeply waisted relative to the proximal and distal ends. Although relatively uninformative, enough of this humerus is preserved to indicate that the deltopectoral crest would have made up less than 50% of the length of the humerus as in nodosaurids [71], [117] and in the basal ankylosaur Mymoorapelta (Kirkland, pers. obs.) compared to the longer deltopectoral crests of ankylosaurids [70], [71]. Overall, the humerus of Europelta is similar in proportions to Niobrarasaurus [118], [119]. The wide proximal end of the humerus figured by Ősi and Prondvai [120] as cf. Struthiosaurus is similar to that of Europelta, whereas the humerus of co-occuring Hungarosaurusis is more slender proportionally.
Among the nine unguals preserved for AR-1/31, one specimen (AR-1-3711/31) may represent a manual ungual. It is more equidimensuional than the other eight more elongate unguals.
Pelvic Girdle
The right ilium of AR-1/10 is fused with its ischium and pubis (AR-1-479/10) which are flexed medially due to compaction (Fig. 22 A–D). The acetabulum is completely enclosed as in all derived ankylosaurs [70], [71], [82], [94], [108]. Only Mymoorapelta is known to retain an open acetabulum [84], [109]. The acetabulum is directed verntrally and is situated medially near the contact of the ilium with the sacrum so that the ilium extends far out beyond the acetabulum laterally for a distance nearly equal to its width. The lateral and anterior margins of the laterally oriented ilium are broken away. The prepubic portion of the ilium diverges from the midline of the sacrum at about 30 degrees and is thickened ventrally along its midline. Large, fairly equi-dimensional, closely appressed osteoderms (7-10 cm in diameter) cover the dorsal surface of the ilium posterior to and medial to the acetabulum. As discussed below, this morphology of sacral armor compares well with “Category 3” pelvic armor of Arbour and others [121]. Anteriorly, the smooth dorsal surface of the ilium is exposed. The pubis is fully fused to the anterior margin of the ischium with no visible sutures; its presence is indicated by a slot-shaped foramen along the anterior side of the ischium. This foramen represents the obturator notch between the postpubic process and the main body of the pubis as in Scelidosaurus and stegosaurs [122]. The distal end of the ischium is broken away. Additionally, AR-1-129/10 is a poorly preserved, proximal left ischium with the pubis fully fused to its anterior margin (Fig. 22 E, F).
Beyond some relatively uninformative fragments of the ilium (Fig. 23 A–C), AR-1/31 includes both the right (AR-1-3648/31) and the left (AR-1-3649/31) ischia with fully fused pubes (Fig. 23 D–M). Both exhibit the slot-shaped foramen along the anterior side of the ischium formed by the obturator notch. The proximal ends appear enrolled such that the anterior and posterior margins are nearly parallel due to compaction. Both display an anterior kink at their distal end as in Cedarpelta [86], [88], but overall are straight-shafted as in the Ankylosauridae [70], [82], [123] and the other European nodosaurids Struthiosaurus [31] and Hungarosaurus [32]. The distal end of the left ischium is the best preserved and measures 299.9 mm long along its anterior margin, including the fully fused pubis forming an ischiopubis. Given the asymmetry of the proximal end of the fused ischium and pubis and the position of the obturator foramen, it appears that the pubis still makes up some of the acetabular margin. The contact between the ilium and the fused ischiopubis is straight with about one-fourth to one-third of the acetabulum formed by the fused ischiopubis.
A straight ischium has been considered to be the primitive character state for ankylosaurs, with the bent ischium of Polacanthus and nodosaurids, a derived character [63], [82], [83], [94], [114], [123]. It is possible that as opposed to being primitive, a straight ischium may be secondarily acquired in the ankylosaurids and European nodosaurids. The only known ischium from the Jurassic ankylosaur (Mymoorapelta) is bent, a trait that is also observed in some stegosaurs such as Kentrosaurus [124]. Stegosaur ischia, even when straight, have an angular thickening near the mid-point of the posterior margin [124] that is shared by the polacanthids Mymoorapelta (Kirkland pers. obs.) and Gastonia [83]. Europelta is the oldest known ankylosaur preserving a straight ischium. The slight kink in the distal end of the ischium of Europelta suggests the straight ischium in European nodosaurids and ankylosaurids is achieved by shortening the ischium distal to the bend.
Hindlimb
The right femur, tibia, and fibula were closely associated (Fig. 24 A–F). The robust right femur (AR-1-3244/31) is 502.9 mm long and 178.9 mm wide at the proximal end and has been flattened anteroposteriorly, with the most distortion to the mid-shaft region. The femoral head is distinct with much of its articular surface directed dorsally and only somewhat medially. It forms an angle of about 115° with the long axis of the femur. The femoral head is directed more dorsally under the ilium in polacanthids [7], [12], [82], [95], [125], and several nodosaururids. In addition, the femoral head of Europelta is expanded such that it overhangs the femoral shaft both anteriorly and posteriorly. The greater trochanter is well demarcated from the femoral head by a constriction across the proximal end of the femur, and the anterior trochanter forms a ridge ventral to the greater trochanter that is fully fused to the femur. The robust fourth trochanter overlaps the midpoint of the femoral shaft and its midpoint is located proximal at the midpoint of the femur. Polacanthids and nodosaurid ankylosaurs have this configuration, whereas in ankylosaurids the fourth trochanter is distal to the middle of the shaft [63], [82], [95], [120], [125]. The distal end of the femur is flattened and forms a planar articular surface relative to the straight femoral shaft. The intercondylar notch is not expressed ventrally, and is better developed posteriorly than anteriorly
The right tibia (AR-1-3237/31) and fibula (AR-1-3238/31) were closely associated (Fig. 6) and post-depositionally compressed. Compression has distorted the distal end of the tibia such that the wide posterior surface is twisted counterclockwise in line with the wide lateral side of the anterior end relative to the orientation of the proximal and distal ends of the tibia in most other ankylosaurs, such as Mymoorapelta [84] (Kirkland, pers. obs.). The fibula was taphonomically displaced ventrally and with the ventral end rotated posteriorly relative to its position in life with the tibia.
The tibia is 458.8 mm long and robust for its entire length (Fig. 24 G–K, Q) as in Cedarpelta [86]. The proximal end is 169.2 mm wide by 93.1 mm wide and its distal end is 146.8 mm wide by 70.2 mm. It is significantly more narrowly waisted in Mymoorapelta [84], Gastonia [83], Polacanthus [7], [12], [18], Sauropelta [69], [71], [99], [108], Peloroplites [86], and in Zhejiangosaurus [126] and ankylosaurids like Saichania [106]. The cnemial crest is broadly rounded. The even curvature of the distal end of the tibia suggests that the astragalus was fully fused to it with no evident sutural contact as in most ankylosaurs [63], [82], [121]. The astragalus is not fused to the distal end of the tibia in Mymoorapelta [84], Gastonia [83], Hylaeosaurus [11], and Peloroplites [86].
Generally, ankylosaurids have tibiae that are less than two-thirds the length of their femora, as opposed to nodosaurids which have proportionally longer lower leg elements [127]. With a tibia to femur ratio of 0.91, Europelta has the proportionally longest tibia of any ankylosaur for which this ratio is known. Both Cedarpelta and Peloroplites have relatively longer tibiae than other ankylosaurs [86], with a tibia to femur ratio of 0.82 in both. Peloroplites differs in its proportionally more narrowly waisted tibial shaft.
The fibula is 395.5 mm long (Fig. 24 L–P, R) and laterally flattened. The proximal end is not expanded anteroposteriorly, such that the slender fibula changes little in size and shape from the proximal to distal end. In lateral view, the proximal end is rounded and the distal end is concave. In cross-section, it is flattened medially and convex laterally. It is longer relative to the tibia than in most other ankylosaurs [108].
A calcaneum (AR-1-3289/31) was identified in association with the lower right leg of AR-1/31. It is laterally compressed, convex laterally and concave medially (Fig. 24 S, T). Its dorsal margin is flattened where it would articulate with the fibula. Calcanea are practically unknown in ankylosaurs, but one has been identified in the juvenile specimen of the derived ankylosaur Anodontosaurus [128]. The type of Niobrarasaurus coleii preserves an articulated lower hind limb, with an astragalus fully fused with the tibia and possessing an articulation with the distal end of the fibula and an unfused calcaneum of similar morphology to that of Europelta [118]. The calcaneum is fully fused to the distal end of the fibula in Saichania [106].
A number of metatarsals and phalanges are associated with AR-1/31. The metatarsals have subrectangular proximal ends, indicating that they were closely articulated in a well-integrated pes in life (Fig. 25 A–W). The pedal phalanges (Fig. 25 X–JJJ) are short, as in other ankylosaurs. There are eight relatively large, elongate, spade-like unguals (Fig. 25 KKK–WWWW) of a morphology similar to pedal unguals in other ankylosaurs in which the unguals are nearly as long as the digits[82], which indicates that portions of both feet are present in AR-1/31. We interpret that the pes of Europelta possesses four pedal phalanges as in most other nodosaurids [80]. Liaoningosaurus has three digits on the pes. The eight similar unguals are interpreted as pedal unguals and the smallest ungual (Fig. 25 XXXX–BBBBB) is interpreted as an isolated manual ungual. The overall proportions of the preserved pedal elements are similar to those of Niobrarasaurus [119], which also has pedal unguals nearly as large as its metatarsals.
Armor
There was an abundance of dermal armor recovered with both AR-1/10 and AR-1/31. On comparison with the quarry maps, none of the osteoderms appears to be preserved in situ with any of the skeletal elements or with each other, and there is no fusion between any of the osteoderms recovered. Therefore, the armor has been divided into several broad morphotypes for the purpose of description and comparison to armor described for other ankylosaurs. Although morphotypes and terminologies have been proposed [129], [130], no system fits for all armor types in all ankylosaurs. A number of researchers have divided armor into types as in Type 1, 2, etc. [131]; for this discussion the armor types are alphabetized to ensure minimal confusion with previous descriptions. The term osteoderm is used to describe relatively larger dorsal and lateral armor elements with the presence of an external keel or tubercle, whereas the term ossicle describes relatively smaller dermal armor lacking a keel, in the sense of Blows [130]. It is recognized that a consistent methodology for describing armor is achievable, but must be done within a phylogenetic framework to be of maximum utility.
Osteoderm surface texture may be broadly useful in differentiating ankylosaurids from nodosaurids [132], [133]. The vast majority of the osteoderms examined in Europelta has a moderately rugose texture with sparse pitting more in keeping with nodosaurids and basal ankylosaurids rather than more derived ankylosaurids. Whereas histological studies have proven useful in the study of thyreophorans [132], [134], [135], that is beyond the scope of this study.
It is noteworthy that no portions of distinct cervical rings were recovered, although cervical vertebrae are known for both skeletons of Europelta. Additionally, only one spine from the cervical or pectoral region was tentatively identified. We postulate that these elements were lost through the process of coal removal or may have been taphonomically removed from the skeletal associations. Only the discovery of additional specimens of Europelta can further reveal the presence of cervical half-rings.
Type A armor.
An isolated fragmentary spine (AR-1-128/10), possibly from the cervical or pectoral region, is recognized from the holotype (Fig. 26 A–D). It appears to represent only the anterior half and may have been cut in two as the overlying coal was removed. This sharp, broken margin reveals an asymmetric, Y-shaped cross-section. The base flares more and is is less excavated than in a Type 2 caudal plate, suggesting that it was positioned on a broad flank of the body. From the possible anterior margin, the spine slopes posteriorly 15 cm to the broken margin in a gradual arc. There is no indication that the spine could not have been longer. The spine is compressed as in the cervical spines of Sauropelta [77], [99] and Edmontonia [110], [136], and the pectoral spines of Gastonia [83] and Polacanthus [7], [10]. The base is asymmetrical in a manner similar to the elongate osteoderms in Mymoorapelta [84], with one side of the base extending lower anteriorly and the other posteriorly. There is no evidence of a basal plate incorporated into fusion of the cervical half-ring as in mature ankylosaurs like Mymoorapelta [84] Gargoyleosaurus [85], [95], Gastonia [83], Polacanthus [10], [130], and Sauropelta [77], [99]. This may relate to the anchoring of larger elements into the dermis in Gastonia and Polacanthus [130]. We tentatively interpret AR-1-128/10 as a pectoral spine. However, if the complete element extends beyond the break for more than twice the length of the preserved portion, it would fall into the category of Type B armor, although that is unlikely because it is more massive form than the Type B elements.
Type B armor.
Dorsoventrally compressed, hollow, asymmetric-based plate-like osteoderms with sharp anterior and posterior edges and lateroposteriorly directed apices are identified for AR-1/10 (Fig 26 E–J) and AR-1/31 (Fig. 27 A–L). Similar large osteoderms have been described as caudal plate ostederms in Mymoorapelta [84], [109], Gargoyleosaurus [85], [95], Gastonia [83], and Polacanthus [8]-[10], [38], [130]. Similar, more anterorposteriorly symmetrical caudal plate osteoderms are also known in Minmi [137], [138] and several Asian ankylosaurids [131]. The few plate-like osteoderms of this morphology that are identified in Europelta are mediolaterally shorter and anteroposteriorly longer with a more posteriorly swept apices. Two pairs of similar plates are known for the holotype of Sauropelta (AMNH 3032), with one of the larger plates being illustrated [99]. One plate from the Yale collections of Sauropelta has a unique double apex (YPM 5490). Given the rarity of Type B armor in Sauropelta and Europelta we hypothesize that caudal plates in these nodosaurids ran down the sides of the tail but decreased in size more rapidly, such that long-keeled osteoderms of Type E morphology made up the lateral armor down most of the length of the tail. It is also possible that these large plate-like osteoderms were on the lateral margin of the sacrum as has been documented by Carpenter and others [106] in Saichania. Struthiosaurus preserves several osteoderms of this morphology that have been reconstructed as in Polacanthus as being medial, dorsally-projecting caudal osteoderms [25], [26]. The relative rarity of these plate-like osteoderms suggests that they were restricted to the base of the tail as well.
Type C armor.
Both AR-1/10 (Fig. 28 A–H, O, P) and AR-1/31 (Fig. 29 A– F) preserve fairly large (∼15–25 cm long) subrectangular to subtrapezoidal, solid osteoderms with low, evenly developed keels running down the long axis of the osteoderm either medially or to one side of the mid-line. Their distal and medial surfaces are subparallel and the entire plate may be slightly flexed across the short axis perpendicular to the crest. The straight, longer margins of these plates appear to have been tightly affixed but not fused to adjoining osteoderms. Armor of Type C morphology is not common but is most similar to medial cervical osteoderms of half-rings, and most distinctively, across the mid-line of the pectoral region in some nodosaurids such as Stegopelta [138], Niobrarasaurus [140], [141], Panoplosaurus [74], [101], and Edmontonia [74], [110].
Type D armor.
Both AR-1/10 and AR-1/31 preserve large (∼10-20 cm long) asymmetric, diamond (Fig. 28 I–N, Q–T; Fig. 29 M–P) to tear-drop shaped (Fig. 29 G–L, Q, R, U,V) osteoderms with a long keel rising to an apex medially to posteriorly and in some specimens extending past the posterior margin of the base. They are distinguished from Type E osteoderms because they are wider than 50% of their length. The wider osteoderms are thinner and more solid than the narrower osteoderms with small pockets under the apices. The more diamond-shaped forms may be more closely appressed to each other in anterior bands similar to Type C armor.
Type D Armor is widely known in the nodosaurids such as Sauropelta [99], Panoplosaurus [101], and Edmontonia. Gastonia is documented to have similar armor [142], although more solid in cross section with less basal excavation, which occurs in oblique rows anterior to the sacrum with each osteoderm separated by a single row of small Type H ossicles. This pattern is similar to the dorsal dermal ornamentation documented for the ankylosaur Tarchia by Arbour and others [130], except that in Tarchia most of the intermediate scales lacked ossified cores. Similar armor is known from the lateral sides of the legs in some ankylosaurs such as Saichania [106].
Type E armor.
Both AR-1/10 and AR-1/31 preserve large (10-15 cm long) moderately asymmetric osteoderms more than twice as long as wide with a long keel higher on the assumed posterior end (Fig. 28 Q, R, II–NN; Fig. 30 G–FF). These osteoderms have proportionally more deeply excavated bases than Type D armor, have chevron-shaped cross-sections, and are distinguished from Type D armor by their width being less than 50% of the length. Type E armor is gradational with Type D armor (Fig. 28 S–T; Fig 29 A–F) and may represent lateral or distal armor from the trunk of the body and along the sides of the tail. This armor type is present in Sauropelta [99] and Texasetes [115]. Similar armor is present on the sides of the limbs in Scelidosurus and Saichania [106].
Type F armor.
Medium to large (∼5-15 cm long) oval to circular osteoderms of low profile with a median keel extending into an apex near or overhanging the posterior margin of the osteoderm are represented in both AR-1/10 (Fig. 28 U–Z) and AR-1/31(Fig. 29 W–VVV). The basal surface of the osteoderm is generally solid except for a small pocket under the apex, reminiscent of Type D armor. Less commonly, the base may be more extensively excavated. Armor of this morphology is abundant in many nodosaurids and makes up the major elements of the armor of Sauropelta anterior to the sacrum in AMNH 3036 [142] and is present in Panoplosaurus [101]. These osteoderms may reside within more expansive spaces among the larger dorsal armor as in Edmontonia (AMNH, 5665) and the polacanthids [81], [82], [93], [107], or may be major armor elements on the posterior portion of the sacrum as in Sauropelta (AMNH 3036). They may also lie on the tail between the Type B caudal plate-like osteoderms, or could be arranged along the lateral side of the limbs as in Saichania [106].
Type G armor.
One piece (AR-1-192/10) of flat, oval to subtriangular armor (AR-1-192/10) from AR-1/10 is about 12 cm long and 7 cm wide and is about 0.5 cm thick throughout (Fig. 28 AA, BB). A pair of similar, osteoderms from the Sauropelta specimen AMNH 3032 was curated with a note from the collector, Barnum Brown, stating that these distinct osteoderms were associated with the forelimbs. Therefore, we suggest a similar position for Type G armor in Europelta.
Type H armor.
Small (∼1-4 cm long) solid ossicles are abundant, with 71 examples from both AR-1/10 (Fig. 31) and AR-1/31 (Fig. 32) illustrated. These ossicles range in shape from round, to oval and even irregularly shaped, and are probably filling in the spaces between larger osteoderms. Small interstitial ossicles are not known for every ankylosaur taxon, but appear to be present in many nodosaurid taxa such as Sauropelta [99], [143] and Edmontonia [74], [136], in polacanthid ankylosaurs such as Gastonia [83] and in some ankylosaurids such as Tarchia [131], in which epidermal scales interstitial to osteoderms do not preserve deeper, interstitial ossicles. Their absence may be real, in that they never form deep to the epidermal scales, taphonomic, in that they are selectively transported away because of their small size and low density, or ontogenetic; in that they only ossify late in ontogeny. The surface texture of Gastonia ossicles is smoother than those of Europelta.
Sacral armor.
Armor is present on the posterior margin of the ilium AR-1-479/10. It is composed of large, subequal-sized (7-10 cm) osteoderms that are tightly sutured together (Fig. 22 A) as in the poorly known Stegopelta [139], Nodosaurus [113], Aletopelta [127], and Glyptodontopelta [132], [144]. These low-relief ossicles lack a central apex or keel. The boundary between the margins of the osteoderms and the area devoid of osteoderms on the ilium is sharply demarcated along the margins of unbroken osteoderms, suggesting the armor was not coossified as in Aletopelta [127] and unlike the fully fused sacral armor in the polacanthids Polacanthus and Gastonia [63], [83]. This form of pelvic armor fits that of Arbour and others' Category 3 pelvic armor [121].
Additionally, there is a unique osteoderm AR-1-653/10 that has a large, posteriorly-curved, plate-like keel extending out from the surface that, considered in isolation, is comparable in size and morphology to Type B armor (Fig. 26 K–N). The base is smooth and gently convex, suggesting it may have been closely appressed to the more anterior portion of the ilium. In overall morphology, this large osteoderm is comparable to the spine-bearing armor plate-like osteoderm identified in Hungarosaurus and interpreted to be present in Struthiosaurus [33].
Unique armor pieces.
Some irregularly shaped armor specimens are not represented by more than one element among this material or in the armor from other taxa. At this time, we can offer no positional interpretation of this armor. AR-1-447/10 is an irregular mass of what we interpret as an osteoderm, although it could be sacral armor (Figure 28 CC–FF). AR-1-438/10 is a small, cap-shaped shaped with a small excavation in the center of the external surface (Fig. 28 OO, PP). Two small, deeply basally excavated, oval osteoderms (Fig. 30 GG–JJ) were collected from AR-1/31(AR-1-3239/31, 3721). These osteoderms lack the external excavation.
Discussion
Europelta (Fig. 33) can be distinguished from any of the ankylosaurs assigned to the Polacanthidae (sensu Kirkland's Polacanthinae [83] and Carpenter's Polacanthidae [63] from the Upper Jurassic and Lower Cretaceous as defined by Yang and others [64]; see Terminology) by its rounded, tear-drop shaped skull and a suborbital horn developed on the posterior portion of the jugal and the quadratojugal posterior to the orbit, as opposed to a triangular-shaped skull that is widest at the posterior margin and a suborbital horn developed exclusively on the jugal (as seen in polacanthids). Post-cranially, it can also be distinguished from polacanthids, by its elongate lower hind limbs, the apparent rarity of cervical, pectoral, and thoracic spines, and reduction in the number of caudal plate-like osteoderms. Likewise, it has an abundance of Type D, asymmetric, tear-drop shaped osteoderms like those observed in many nodosaurids and absent in all polacanthids.
Europelta is also distinguished from derived ankylosaurids by its weakly ornamented teardrop-shaped skull in which the lower temporal opening is visible in lateral view. The absence of a tail club also distinguishes the taxon from these ankylosaurids. More basal “shamosaurine grade” ankylosaurids [63], [86] are more similar to Europelta, but also have the lower temporal openings completely obscured laterally by expanding the lateral margin of their skulls. “Shamosaurine grade” ankylosaurids also possess skulls that are approximately as wide mediolaterally between the orbits as they are across the posterior margin.
Europelta shares a number of derived characters with nodosaurids [71], [72], [83], [94], [114]. It has a tear-drop shaped skull that is longer than wide with its greatest width dorsal to the orbits, whereas the short, boxy skulls of Minmi and all anklosaurids are essentially as wide at the posterior edge of the skull, as are the elongate skulls of “shamosaurine-grade” ankylosaurids. Grooves in the remodeled textured skull roof define epidermal scale impressions, with the largest covering the frontoparietal area. Although poorly preserved, the laterally extensive pterygoids are pressed up against the anterior face of the braincase. All known nodosaurid scapulae have a prominent acromion process extending on to the blade of the scapula that terminates in an expanded knob. Unfortunately, this portion of the scapula is as yet unknown in Europelta.
Some character states considered typical of nodosaurids are absent in Europelta. Instead of having a distinct hourglass-shaped palate typical of nodosaurids [70], [71], [82], [83], [114], the upper tooth rows show less lateral emargination and diverge posteriorly. This is also true of Silvisaurus, which also shares an expanded lateral wall of the skull [76], [77]. The coracoid of Europelta is nearly as long as it is tall, whereas in other nodosaurids, for which the corocoid is known, it is expanded anteriorly and longer than tall [71], [72], [83], [94], [114].
The only other Early Cretaceous nodosaurid to have large cranial scales as in Europelta is Propanoplosaurus, known only from an embryonic to hatchling specimen from the base of the Potomac Group of Maryland [145]. However, only the anterior cranial scales are well defined in Propanoplosaurus, whereas only the posterior scale pattern in Europelta. The unusual preservation and extremely small size of Propanoplosaurus lead us to suspect that the fossil preserves the actual scales overlying the skull and not the remodeled skull roof, because this is such a young specimen and remodeling of the cranial bones is not expected to have occurred so early in ontogeny [129], [146].
Additionally, a number of important characters traditionally used to define nodosaurids are not known in Europelta, as yet, because of the missing anteroventral half of the scapula and the absence of premaxilla and surangulars. Thus, the presence absence of premaxillary teeth, if the tooth row joined the margin of premaxillary beak, the morphology of the naris, the height of the coronoid process, and the morphology of the acromion process are unknown for Europelta.
Europelta is distinguishable from European nodosaurids from the Albian through the Cenomanian. The juvenile Anoplosaurus from the Albian Gault Clays of southern England differs in a number of characters, such as possessing a proportionally longer coracoid, a narrower proximal end of the humerus, and a femur with a separate anterior trochanter [17] although the latter two characters are consistent with the juvenile nature of Anoplosaurus. No pectoral spines of the morphology described for “Acanthopholis” from the Cenomanian Lower Chalk in southern England by Huxley [13] are known in Europelta. Additionally, the tall teeth assigned to “Acanthopholis” are distinct in the long apicobasal ridges extending from the denticles to the root on medial and lateral faces of the teeth, and in the presence of caudal ribs that extend laterally and flex ventrally, whereas the caudal ribs in Europelta extend ventrolaterally and flex laterally [16], [17]. Europelta is like other Late Cretaceous European nodosaurids in having a short symphysis for the predentary, a mediolaterally wide and anteroposteriorly thin quadrate, an anteroposteriorly arched sacrum, and a straight ischium [21], [32].
The domed skull and elongate cervical vertebrae in Struthiosaurus clearly distinguish it from Europelta. Likewise, Hungarosaurus also has more elongate cervical vertebrae [32]. Both Hungarosaurus and Struthiosaurus possess a pair of spines on the anterior portion of the pelvis [33], whereas we interpret the presence of a pair of upright plate-like armor elements in this position in Europelta (Fig. 33).
The lateral wall of the skull in most North American nodosaurids is typically narrow [82], whereas in Europelta it is relatively wider, although a broad notch along its posterior margin permits the caudal margin of the lower temporal opening to be observed in lateral view. This morphology in Europelta is similar to that in the nodosaurids Silvisaurus [76], [77] and Peloroplites [86]. Although, the skull of Struthiosaurus transylvanicus is highly reconstructed [22], it appears that the lateral wall of the skull is expanded laterally, whereas not completely obscuring the lower temporal opening. This character state is not known in other species of Struthiosaurus, but appears to be moderately developed in Hungarosaurus [32].
Comparisons of Europelta with the Asian”nodosaurids” Zhongyuansaurus [93] and Zhejiangosaurus [126] from the lower Upper Cretaceous of China hinges partially on the question of whether those taxa have been validly referred to Nodosauridae. Carpenter and others [86] noted that the skull of Zhongyuansaurus is morphologically similar to that of a “shamosaurine-grade” (like Shamosaurus and Gobisaurus) ankylosaurids and was the first shamosaurine-grade ankylosaurid documented to not have a tail club. However, its distal tail is modified into a stiffened structure of the same morphology as the “handle” of the tail club in more derived ankylosaurids [147], [148]. Zhejiangosaurus was assigned to the nodosaurids based on characteristics of the femur and sacrum, together with the lack of a tail club [126]. We hypothesize that it lacked a knob as in basal ankylosaurids, polacanthids and nodosaurids because ankylosaurids with a full tail club have distal free caudal vertebrae bearing caudal ribs at the base of the handle. Most of the distal caudal vertebrae of Zhejiangosaurus have raised ridges on the sides of the centra as in the distal vertebrae of polacanthids and nodosaurids. Additionally, whereas the position of its most proximal preserved caudal vertebrae is not known, morphologically, they do not appear to represent the most proximal caudal vertebrae. Thus, while Zhejiangosaurus' 13 preserved caudal vertebra are more than the number of free caudals preserved in most ankylosaurs with tail clubs (10 in Saichania [106] and Dyoplosaurus [148]), the total number of free caudals in its tail would appear to be more than the 14 in Tarchia [130] and 15 in Pinacosaurus [129]. Unlike nodosaurids, Zhejiangosaurus has an exceedingly low ratio of femur to tibia length of 0.46 similar to that of with ankylosaurids and polacanthids rather than nodosaurids. Dongyangopelta [149] was described as a second nodosaurid from the same area and stratum as Zhejiangosaurus, which was found to be its sister taxon in their phylogeny [149]. With few overlapping elements, we feel that the proposed differences between these taxa may be due to preservation, individual variation, or ontogeny. Additionally, given the presence of a pelvic shield and numerous caudal plate-like osteoderms in Dongyangopelta, we suggest that both specimens may pertain to the same taxon and represent the first polacanthid described from Asia. Given the recent description of the polacanthid Taohelong from the upper portion of the Lower Cretaceous of Gansus Province in western China [64], this hypothesis has added support. We also do not think that the partial ankylosaur skull reported from the lower Upper Cretaceous of Hokkaido, Japan [150] can be diagnosed as a nodosaurid with any confidence at this time, due to the incomplete nature of the specimen. Thus, we do not presently recongnize the presence of true nodosaurids in Asia.
In his seminal paper defining a bipartite division of the Ankylosauria into Ankylosauridae and Nodosauridae, Coombs [71] hypothesized that Acanthopholis (as a nomen dubium in which he would have included Anoplosaurus) and Struthiosaurus might represent a separate lineage of European nodosaurids. Unlike Hylaeosaurus (in which he included Polacanthus), these taxa had a well-developed supraspinus fossa developed anteriorly on the scapula as did all North American nodosaurids. This European lineage was hypothesized based on their small body size, presence of premaxillary teeth, and their possessing an unfused scapula and corocoid. Although, none of the characters are valid in defining such a group, our research on Europelta has resulted in supporting the taxonomic hypothesis of Coombs [71], [72] as correct, just for the wrong reasons.
Relationships to Other Taxa
We use Struthiosaurinae to define the clade of European nodosaurs. Nopcsa [25] proposed Acanthopholidae as a family of relatively lightly built thyreophorans, that included Acanthopholis ( = Anoplosaurus), Polacanthus, Stegopelta, Stegoceras, and Struthiosaurus. In 1923, he divided the Acanthopholidae into an Acanthopholinae and a Struthiosaurinae without comment [69]. Subsequently, he relegated the Acanthopholidae to a subfamily of the Nodosauridae, in which he also included Ankylosaurus and restricted the Acanthopholinae to Acanthopholis, Hylaeosaurus, Rhodanosaurus, Struthiosaurus, Troodon [26], [151]. This artificial grouping included a polacanthid ankylosaur [72], [83], a pachycephalosaur [152] and Acanthopholis, now considered a nomen dubium [17], [82]. Thus, the term Acanthopholinae is not acceptable for this newly recognized clade of nodosaurids. Thus, Struthiosaurinae is the next published term available to use for this clade and is derived from the first described and youngest member of this clade. Struthiosaurinae is defined as the most inclusive clade containing Europelta but not Cedarpelta, Peloroplites, Sauropelta or Edmontonia.
In order to determine the systematic position of Europelta, it was found that previous cladistic analyses [71], [72], [82], [83], [114], did not include many of the character states that we identify as significant in our research on Upper Jurassic and Lower Cretaceous ankylosaurs. A major weakness of these analyses is the limited recognition of postcranial skeletal and dermal characters that restricts the testing the phylogenetic relationships for taxa for which skulls are either poorly known or not known at all.
We present a character based definition of Struthiosaurinae as: nodosaurid ankylosaurs that share a combination of characters including: narrow predentary; a nearly horizontal, unfused quadrate that is oriented less than 30° from the skull roof, and mandibular condyles that are 3 times transversely wider than long; premaxillary teeth and dentary teeth that are near the predentary symphysis; dorsally arched sacrum; an acromion process dorsal to midpoint of the scapula-coracoid suture; straight ischium, with a straight dorsal margin; relatively long slender limbs; a sacral shield of armor; and erect pelvic osteoderms with flat bases. This suite of characters unites Europelta with the European nodosaurids Anoplosaurus, Hungarosaurus and all species assigned to Struthiosaurus. This clade of European nodosaurids has not been previously recognized. Europelta represents the earliest member of the European clade Struthiosaurinae.
Biogeogeographic Implications
The near simultaneous appearance of nodosaurids in both North America and Europe is worthy of consideration (Fig. 34). Europelta is the oldest nodosaurid known in Europe, it derived from strata in the lower Escucha Formation that is dated to early Albian. The oldest nodosaurid from western North America is Sauropelta, which in the lower part of its range is in the lower Albian Little Sheep Mudstone Member (B interval) of the Cloverly Formation in northern Wyoming and southern Montana [99], [153] with an ash bed 75 meters above the base near the top of the member providing an age of 108.5±0.2 Ma [154]. Nodosaurid remains from eastern North America appear to be older. Teeth of a large nodosaurid Priconodon crassus are known from the Arundel Clay of the Potomac Group [77], [155], which palynology dates as near the Albian-Aptian stage boundary [156]. The hatchling Propanoplosaurus is from the base of the underlying Patuxent Formation of the Potomac Group of Maryland, which has been dated as late Aptian [157], [158], making Propanoplosaurus the oldest known nodosaurid. Polacanthid ankylosaurs characterize pre-Aptian faunas in both Europe [11], [12], [37]-[39] and North America [70], [95], [159]. We have not been able to document a specific example of Polacanthus in the Lower Aptian Vectis Formation of the Wealden Group, although Polacanthus has been reported to occur in those strata [10]-[12], [82], [160]. However, polacanthids are present in the lower Aptian Morella Formation of northeastern Spain [40]. Blows [10] illustrated a block with ankylosaur dorsal vertebrae with the uninformative ventral portion of a pelvic shield fragment and noted it as being from Charmouth, suggesting that there were upper Albian polacanthids in England [160]. However, the specimen NMW 92.34G.2 was actually found on the beach further west at Charton Bay and may have come from either the Aptian (Lower Greensand) or Albian (Upper Greensand). Only preparation of the dorsal surface of the pelvic shield would reveal if the specimen is a polacanthid or nodosaurid. A large polacanthid (BYU R254) occurs in the Poison Strip Sandstone Member of the Cedar Mountain Formation [156]. It is not a nodosaurid close to Sauropelta as reported by Carpenter and others [97], but a polacanthid that was initially described as cf. Hoplitosaurus [161]. These rocks have been dated as lower to middle Aptian by laser ablation of detrital zircons and by U-Pb dating of early diagenetic carbonate [162]. A fragmentary large nodosaurid with massive cervical spikes that may be referred to as cf. Sauropelta (DMNS 49764) has been recovered from the overlying Ruby Ranch Member about 20 m up section in the same region [163] in strata interpreted to be of Lower Albian age [162]. Thus, the youngest polacanthids occur in the lower to possibly mid-Aptian and the oldest documented nodosaurids occur in the upper Aptian or lower Albian in both Europe and North America with no discernible stratigraphic overlap (Fig. 34). Why this faunal discontinuity occurs is unknown. There are no documented significant changes in sea level or shifts in geochemical indicators to suggest a geological or environmental change that would affect ankylosaurs on both continents at approximately the same time [164]. However, the OAE1a or “Sella” organic burial episode near the base of the Aptian was followed by a positive carbon isotope excursion that may have precipitated longer-term environmental effects that would result in the turnover of ankylosaurs in the “middle” Aptian [165]. In North America, “medial” grade iguanodonts (basal Steracosterna) are replaced by the considerably more primitive basal iguanodont Tenontosaurus at this time, while in Europe the lower Albian more derived iguanodont Proa is phylogenetically close to Iguanodon [43], [159] at the base of Hadrosauriformes [43], documenting different patterns of faunal change for iguanodonts and ankylosaurs. Therefore, a cause for this faunal turnover, which might specifically have affected ankylosaurs, should be sought. Ankylosaurs are low feeders, so perhaps the rapid ongoing radiation of flowering plants at this time [166]-[170] might have driven their diversification. It has been proposed that this floral revolution was linked to a decline in atmospheric CO2 concentrations [171] or, more likely, an increase in CO2 and global warming resulting from massive early Aptian volcanic activity forming the Ontong Java and Manihiki plateaus [172], [173]-[174]. Therefore the rapid domination of shrubby angiosperms may have caused a disruption in the availability of forage to which polacanthids were adapted. Kirkland and others have proposed that North America became isolated from Europe at the end of the Barremian [159], [175]. Certainly the timing of the appearance of nodosaurids on both continents indicates that the origins of the clade preceded the complete isolation of North America and Europe pushing up this date in to at least the “middle” Aptian. The separation of the Nodosauridae into a North American Nodosaurinae and a European Struthiosaurinae by the end of the Aptian, would thus provide a revised date for the isolation of North America from Europe with rising sealevel.
Additionally, whereas there is no definitive evidence for nodosaurids in Asia, apparently polacanthids entered Asia in the later portion of the Early Cretaceous and survived there in isolation into the early Late Cretaceous.
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Dinosaurs list, List of Dinosaurs, Dinosaurs a to z, List of Dinosaurs a
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Dinosaurs » A-Z Dinosaurs List
A-Z Dinosaurs List
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https://s.click.aliexpress.com/deep_link.htm%3Faff_short_key%3DUneMJZVf%26dl_target_url%3Dhttps%253A%252F%252Fwww.aliexpress.com%252Fitem%252F1005005657852691.html%253F_randl_currency%253DGBP%2526_randl_shipto%253DGB%2526src%253Dgoogle
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淘！我喜欢
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https://en.wikipedia.org/wiki/List_of_Asian_dinosaurs
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List of Asian dinosaurs
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https://en.wikipedia.org/static/favicon/wikipedia.ico
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https://en.wikipedia.org/static/favicon/wikipedia.ico
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[
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2009-04-25T19:33:56+00:00
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en
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https://en.wikipedia.org/wiki/List_of_Asian_dinosaurs
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Name Year Formation Location Notes Images Abdarainurus 2020 Alagteeg Formation (Late Cretaceous, Santonian to Campanian) Mongolia Inconsistent in phylogenetic placement; could represent an unknown lineage of macronarians[1] Abrosaurus 1989 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China Had unusually large fenestrae Achillobator 1999 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia Its robust build suggests it was not a cursorial animal[2] Adasaurus 1983 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Its sickle claw was markedly reduced compared to other dromaeosaurids Aepyornithomimus 2017 Djadochta Formation (Late Cretaceous, Campanian) Mongolia The first ornithomimosaur named from a dry desert environment Agilisaurus 1990 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China The holotype specimen was discovered during the construction of the museum where it is now housed Albalophosaurus 2009 Kuwajima Formation (Early Cretaceous, Valanginian to Hauterivian) Japan Only known from fragments of a skull Albinykus 2011 Javkhlant Formation (Late Cretaceous, Santonian) Mongolia Preserved in a sitting position not unlike that of modern birds Alectrosaurus 1933 Iren Dabasu Formation (Late Cretaceous, Cenomanian to Santonian) China Had long legs which may be an adaptation to pursuit predation[3] Alioramus 1976 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Possessed an elongated snout with a row of short crests Almas 2017 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Preserved alongside eggshells which may have come from a troodontid[4] Altirhinus 1998 Khuren Dukh Formation (Early Cretaceous, Barremian to Albian) Mongolia Had a distinctive, elevated nasal bone which supported a large nasal cavity Alxasaurus 1993 Bayin-Gobi Formation (Early Cretaceous, Albian) China Most of the skeleton is known, which allowed researchers to connect therizinosaurs to other theropods Ambopteryx 2019 Unnamed formation (Late Jurassic, Oxfordian) China Preserves stomach contents containing gastroliths and fragments of bone, suggesting an omnivorous diet Amtocephale 2011 Bayan Shireh Formation (Late Cretaceous, Turonian to Santonian) Mongolia One of the oldest known pachycephalosaurs Amurosaurus 1991 Udurchukan Formation (Late Cretaceous, Maastrichtian) Russia One specimen may have come from an individual with a limp[5] Analong 2020 Chuanjie Formation (Middle Jurassic, Bajocian) China Originally described as a specimen of Chuanjiesaurus but assigned a new genus due to several morphological differences Anchiornis 2009 Tiaojishan Formation (Late Jurassic, Oxfordian) China Analysis of fossilized melanosomes suggest a mostly gray or black body, white and black patterns on its wings, and a red head crest[6] Anhuilong 2020 Hongqin Formation (Middle Jurassic, Aalenian to Callovian) China Closely related to Huangshanlong and Omeisaurus, all forming an exclusive clade of mamenchisaurids Anomalipes 2018 Wangshi Group (Late Cretaceous, Maastrichtian) China May have been closely related to Gigantoraptor despite its significantly smaller size[7] Anserimimus 1988 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Had powerful forelimbs with uniquely straight, flattened claws Aorun 2013 Shishugou Formation, (Late Jurassic, Oxfordian) China Potentially a basal member of the alvarezsaurian lineage[8] Aralosaurus 1968 Bostobe Formation, (Late Cretaceous, Santonian to Campanian) Kazakhstan Its crest has been interpreted as being arch-shaped as in kritosaurin hadrosaurs, but this cannot be confirmed Archaeoceratops 1997 Xinminbao Group (Early Cretaceous, Barremian) China Had no horns and only the beginnings of a frill Archaeornithoides 1992 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Known from only a partial skull with scratches that may have been created by a small mammal[9] Archaeornithomimus 1972 Bissekty Formation?, Iren Dabasu Formation (Late Cretaceous, Cenomanian to Turonian) China
Uzbekistan? Unlike other ornithomimosaurs, its feet were not arctometatarsalian Arkharavia 2010 Udurchukan Formation (Late Cretaceous, Maastrichtian) Russia Described from a series of vertebrae, several of which were found to not belong to this taxon[10] Arstanosaurus 1982 Bostobe Formation (Late Cretaceous, Santonian to Campanian) Kazakhstan Poorly known Asiaceratops 1989 Khodzhakul Formation, Xinminbao Group? (Late Cretaceous, Cenomanian) China?
Uzbekistan Potentially a leptoceratopsid[11] Asiatosaurus 1924 Öösh Formation, Xinlong Formation (Early Cretaceous, Barremian to Albian) China
Mongolia Two species have been named but both are only known from extremely scant remains Auroraceratops 2005 Xinminbao Group (Early Cretaceous, Aptian) China Known from more than eighty specimens, including complete skeletons Aurornis 2013 Tiaojishan Formation (Late Jurassic, Oxfordian) China If an avialan as originally described it would be one of the oldest members of the group Avimimus 1981 Barun Goyot Formation, Nemegt Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia Bonebed remains indicate a gregarious lifestyle; it may have formed age-segregated herds for lekking or flocking purposes[12] Bactrosaurus 1933 Iren Dabasu Formation, Majiacun Formation? (Late Cretaceous, Cenomanian to Santonian?) China Remains of at least six individuals are known, making up much of the skeleton Bagaceratops 1975 Barun Goyot Formation, Bayan Mandahu Formation, Djadochta Formation? (Late Cretaceous, Campanian to Maastrichtian) China
Mongolia May have been a direct descendant of Protoceratops which it physically resembles[13] Bagaraatan 1996 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Combines traits of several theropod groups, possibly due to being chimaeric[14] Bainoceratops 2003 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Its supposedly diagnostic features may fall within Protoceratops variation[15] Banji 2010 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Vertical striations adorned the sides of its crest Bannykus 2018 Bayin-Gobi Formation (Early Cretaceous, Barremian to Aptian) China Exhibited a transitional hand morphology for an alvarezsaur, having three fingers of roughly equal length with the first being robust Baotianmansaurus 2009 Gaogou Formation (Late Cretaceous, Cenomanian to Turonian) China Large but known from only a few bones Barsboldia 1981 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Possessed elongated neural spines particularly above the hips Bashanosaurus 2022 Shaximiao Formation (Middle Jurassic, Bajocian) China Its skeleton combines traits of stegosaurs and more basal thyreophorans Bashunosaurus 2004 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China Although described as a macronarian, this has yet to be rigorously tested[16] Batyrosaurus 2012 Bostobe Formation (Late Cretaceous, Santonian to Campanian) Kazakhstan Remains originally identified as Arstanosaurus Bayannurosaurus 2018 Bayin-Gobi Formation (Early Cretaceous, Aptian) China Known from a well-preserved, almost complete skeleton Beg 2020 Ulaanoosh Formation (Early Cretaceous to Late Cretaceous, Albian to Cenomanian) Mongolia Its preserved skull has a rugose texture Beibeilong 2017 Gaogou Formation (Late Cretaceous, Cenomanian to Coniacian) China Similar to but more basal than Gigantoraptor.[17] Known from only a single embryo still in its egg Beipiaosaurus 1999 Yixian Formation (Early Cretaceous, Aptian) China Preserves evidence of downy feathers as well as a secondary coat of simpler "elongated broad filamentous feathers" or EBFFs[18] Beishanlong 2010 Xinminbao Group (Early Cretaceous, Aptian to Albian) China Lacked the elongated claws of more derived ornithomimosaurs Bellusaurus 1990 Shishugou Formation (Late Jurassic, Oxfordian) China Known from a bone bed with the remains of seventeen juvenile specimens Bienosaurus 2001 Lufeng Formation (Early Jurassic, Sinemurian) China Potentially synonymous with Tatisaurus[19] Bissektipelta 2004 Bissekty Formation (Late Cretaceous, Turonian to Coniacian) Uzbekistan Analysis of its braincase suggests poor hearing and eyesight but good olfaction and taste; it has been suggested to be a filter feeder[20] Bolong 2010 Yixian Formation (Early Cretaceous, Aptian) China Originally known from only a skull; an almost complete skeleton was described in 2013[21] Borealosaurus 2004 Sunjiawan Formation (Late Cretaceous, Cenomanian to Turonian) China Its caudal vertebrae were distinctively opisthocoelous Borogovia 1987 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Had a uniquely straight and flattened sickle claw, which may have had a weight-bearing function Breviceratops 1990 Barun Goyot Formation (Late Cretaceous, Campanian) Mongolia Only known from juvenile remains but can be distinguished from other protoceratopsids Brohisaurus 2003 Sembar Formation (Late Jurassic, Kimmeridgian) Pakistan Possibly an early titanosauriform Byronosaurus 2000 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Two juvenile skulls were found in an oviraptorid nest and claimed to be evidence for nest parasitism in this taxon, but both their identity and taphonomy have been questioned[4][22] Caenagnathasia 1994 Bissekty Formation (Late Cretaceous, Turonian to Coniacian) Uzbekistan One of the oldest and smallest known caenagnathoids Caihong 2018 Tiaojishan Formation (Late Jurassic, Oxfordian) China Possessed platelet-shaped melanosomes that produced iridesence as in modern trumpeters Caudipteryx 1998 Yixian Formation (Early Cretaceous, Barremian) China Two species are known. At least C. zoui did not have secondary feathers attached to the lower arm Ceratonykus 2009 Barun Goyot Formation (Late Cretaceous, Campanian) Mongolia Several osteological features were described as similar to ornithischians[23] Changchunsaurus 2005 Quantou Formation (Early Cretaceous to Late Cretaceous, Aptian to Cenomanian) China Had wavy enamel on its leaf-shaped teeth that made them more resistant to wear; this feature is also present in hadrosaurs[24] Changmiania 2020 Yixian Formation (Early Cretaceous, Barremian) China Preserved in a curled-up position as if sleeping in a potential burrow Changyuraptor 2014 Yixian Formation (Early Cretaceous, Barremian) China The largest microraptorian dromaeosaurid known. Had tail feathers almost a foot long[25] Chaoyangsaurus 1999 Tuchengzi Formation (Late Jurassic, Tithonian) China Known by a number of alternate spellings (e.g. Chaoyangosaurus, Chaoyoungosaurus) before its formal description Charonosaurus 2000 Yuliangze Formation (Late Cretaceous, Maastrichtian) China May have had a long, backwards-arcing crest similar to that of Parasaurolophus Chialingosaurus 1959 Shaximiao Formation (Late Jurassic, Oxfordian to Kimmeridgian) China Had both large plates and smaller spines, similar to Kentrosaurus Chiayusaurus 1953 Hasandong Formation, Xinminbao Group (Early Cretaceous, Barremian to Albian) China
South Korea Two species have been named, both from teeth. Those of C. lacustris are apparently indistinguishable to those of Euhelopus[26] or Mamenchisaurus[27] Chilantaisaurus 1964 Ulansuhai Formation (Late Cretaceous, Turonian) China Had a particularly hooked claw on its first finger Chingkankousaurus 1958 Wangshi Group (Late Cretaceous, Santonian to Campanian) China Known from only a scapula. Possibly a tyrannosauroid[28] Chinshakiangosaurus 1992 Fengjiahe Formation (Early Jurassic, Hettangian) China Had a U-shaped snout that may have supported fleshy cheeks, an adaptation to bulk feeding Choyrodon 2018 Khuren Dukh Formation (Early Cretaceous, Albian) Mongolia It had an enlarged nose similar to its contemporary, Altirhinus, but it is most likely a separate taxon[29] Chuandongocoelurus 1984 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China A tetanuran of uncertain relationships Chuanjiesaurus 2000 Chuanjie Formation (Middle Jurassic, Bathonian) China One of the more derived mamenchisaurids[30] Chuanqilong 2014 Jiufotang Formation (Early Cretaceous, Barremian to Aptian) China May have been the adult form of the coeval Liaoningosaurus[31] Chungkingosaurus 1983 Shaximiao Formation (Late Jurassic, Oxfordian) China May have possessed at least six thagomizer spikes; the rearmost pair was mounted horizontally, directed outwards and backwards Chuxiongosaurus 2010 Lufeng Formation (Early Jurassic, Hettangian to Pliensbachian) China Potentially a synonym of Jingshanosaurus[32] Citipati 2001 Djadochta Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia Had a distinctive triangular crest. A referred specimen known as the Zamyn Khondt oviraptorid possessed the familiar rectangular domed crest in most depictions of Oviraptor, but likely does not belong to that genus or Citipati[33] Conchoraptor 1986 Barun Goyot Formation, Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Named for a hypothesized diet of shellfish, but this cannot be confirmed Corythoraptor 2017 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Its crest was vertical and rectangular, not unlike that of a cassowary Crichtonpelta 2015 Sunjiawan Formation (Late Cretaceous, Cenomanian) China Originally named as a second species of Crichtonsaurus Crichtonsaurus 2002 Sunjiawan Formation (Late Cretaceous, Cenomanian to Turonian) China Sometimes reconstructed with semicircular osteoderms vaguely similar to the plates of stegosaurs Daanosaurus 2005 Shaximiao Formation (Late Jurassic, Oxfordian to Tithonian) China Phylogenetic position is uncertain as it is only known from the remains of a juvenile Daliansaurus 2017 Yixian Formation (Early Cretaceous, Barremian) China Had an enlarged claw on the fourth toe comparable in size to the sickle claw on its second Dashanpusaurus 2005 Shaximiao Formation (Middle Jurassic, Callovian) China One of the basalmost and earliest known macronarians[34] Datanglong 2014 Xinlong Formation (Early Cretaceous, Barremian to Albian) China Had a uniquely pneumatized ilium similar to megaraptorans Datonglong 2016 Huiquanpu Formation (Late Cretaceous, Cenomanian to Campanian) China Precise dating uncertain Datousaurus 1984 Shaximiao Formation (Middle Jurassic to Late Jurassic, Bathonian to Oxfordian) China One of the rarer sauropods of the Shaximiao, known from only two skeletons and a large, deep skull Daurlong 2022 Longjiang Formation (Early Cretaceous, Aptian) China Preserves remains of an intestinal tract Daxiatitan 2008 Hekou Group (Early Cretaceous, Barremian) China Large and very long-necked Deinocheirus 1970 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Had a suite of unique features, most notably a hump supported by elongated neural spines Dilong 2004 Yixian Formation (Early Cretaceous, Barremian) China Preserves evidence of a coating of simple feathers Dongbeititan 2007 Yixian Formation (Early Cretaceous, Barremian) China A theropod tooth has been found encrusted in one of its ribs[35] Dongyangopelta 2013 Chaochuan Formation (Early Cretaceous to Late Cretaceous, Albian to Cenomanian) China Coexisted with Zhejiangosaurus but could be distinguished based on subtle osteological features[36] Dongyangosaurus 2008 Jinhua Formation (Late Cretaceous, Turonian to Coniacian) China Its phylogenetic placement is uncertain Dzharaonyx 2022 Bissekty Formation (Late Cretaceous, Turonian) Uzbekistan One of the oldest known parvicursorines Dzharatitanis 2021 Bissekty Formation (Late Cretaceous, Turonian) Uzbekistan Originally described as a rebbachisaurid[37] but later reinterpreted as a titanosaur with possible lognkosaurian affinities[38] Elmisaurus 1981 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia One of the most complete caenagnathids known Embasaurus 1931 Neocomian Sands (Early Cretaceous, Berriasian) Kazakhstan Known from only two vertebrae Enigmosaurus 1983 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia Had a large, backwards-pointing pelvis Eomamenchisaurus 2008 Zhanghe Formation (Middle Jurassic to Late Jurassic, Aalenian to Oxfordian) China One of the oldest mamenchisaurids Eosinopteryx 2013 Tiaojishan Formation (Late Jurassic, Oxfordian) China Described as lacking advanced tail feathers and long "hind wings", unlike other paravians, but this may be an artifact of preservation[39] Epidexipteryx 2008 Haifanggou Formation (Middle Jurassic, Callovian) China Supported four long feathers from an abbreviated tail Equijubus 2003 Xinminbao Group (Early Cretaceous, Albian) China A grazer that preserves the oldest known evidence of grass-eating[40] Erketu 2006 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia May have had the longest neck of any dinosaur relative to its body Erliansaurus 2002 Iren Dabasu Formation (Late Cretaceous, Cenomanian) China Had long, curved claws on its fingers Erlikosaurus 1980 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia Preserves the most complete skull known from any therizinosaur Eshanosaurus 2001 Lufeng Formation (Early Jurassic, Hettangian) China Has been suggested to be the oldest known therizinosaur Euhelopus 1956 Meng-Yin Formation (Early Cretaceous, Berriasian to Valanginian) China Originally believed to have lived in a marshy environment Euronychodon 1991 Bissekty Formation (Late Cretaceous, Turonian) Uzbekistan Type species was found in Portugal. The Asian species may represent a form taxon of improperly developed teeth[41] Ferganasaurus 2003 Balabansai Formation (Middle Jurassic, Callovian) Kyrgyzstan Claimed to have two hand claws, but this is disputed[42] Ferganocephale 2005 Balabansai Formation (Middle Jurassic, Callovian) Kyrgyzstan Unusually, its teeth were not serrated Fujianvenator 2023 Nanyuan Formation (Late Jurassic, Tithonian) China Possessed proportionally long legs which may be an adaptation to wading Fukuiraptor 2000 Kitadani Formation, Sebayashi Formation? (Early Cretaceous, Barremian to Aptian) Japan Similarly to Megaraptor, it was originally reconstructed as a dromaeosaur with its hand claw on its foot Fukuisaurus 2003 Kitadani Formation (Early Cretaceous, Barremian) Japan The elements of its skull are so strongly fused that it was unable to chew[43] Fukuititan 2010 Kitadani Formation (Early Cretaceous, Barremian to Aptian) Japan The first sauropod named from Japan Fukuivenator 2016 Kitadani Formation (Early Cretaceous, Barremian to Aptian) Japan Possesses traits of various groups of coelurosaurs, though probably a therizinosaur.[44] May have been a herbivore or omnivore due to its heterodont dentition Fulengia 1977 Lufeng Formation (Early Jurassic, Hettangian to Toarcian) China May have been a juvenile Lufengosaurus Fushanosaurus 2019 Shishugou Formation (Late Jurassic, Oxfordian) China Known from a single femur of immense size Fusuisaurus 2006 Xinlong Formation (Early Cretaceous, Aptian to Albian) China A referred humerus may support an extremely large size for this taxon[45] Gallimimus 1972 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Had a relatively long beak with a rounded tip Gannansaurus 2013 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Its vertebrae were more similar to those of Euhelopus than to other sauropods Ganzhousaurus 2013 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Coexisted with at least seven other oviraptorosaurs, which may have niche-partitioned. It was likely primarily herbivorous[46] Garudimimus 1981 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia Was not as well-adapted to running as later ornithomimosaurs Gasosaurus 1985 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China Discovered as a byproduct of construction work Gigantoraptor 2007 Iren Dabasu Formation (Late Cretaceous, Cenomanian) China The largest known oviraptorosaur, comparable in size to Albertosaurus Gigantspinosaurus 1992 Shaximiao Formation (Late Jurassic, Oxfordian) China Possessed broad, greatly enlarged shoulder spines Gilmoreosaurus 1979 Bissekty Formation?, Iren Dabasu Formation, Khodzhakul Formation? (Late Cretaceous, Cenomanian) China
Uzbekistan? Several fossils preserve evidence of cancer-induced tumors[47] Gobihadros 2019 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia Known from multiple specimens representing different growth stages Gobiraptor 2019 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Possessed a deep jaw that may be an adaptation to crushing bivalves or seeds[48] Gobisaurus 2001 Ulansuhai Formation (Late Cretaceous, Turonian) China Had no tail club but already possessed the stiff tail of derived ankylosaurids[49] Gobititan 2003 Xinminbao Group (Early Cretaceous, Aptian) China Retained the fifth digit of the foot, a basal trait Gobivenator 2014 Djadochta Formation (Late Cretaceous, Campanian) Mongolia The most completely known Cretaceous troodontid Gongbusaurus 1983 Shaximiao Formation (Late Jurassic, Oxfordian) China Only known from a pair of teeth. May be an ankylosaurian[50] Gongpoquansaurus 2014 Xinminbao Group (Early Cretaceous, Albian) China Remains originally named as a species of Probactrosaurus Gongxianosaurus 1998 Ziliujing Formation (Early Jurassic, Toarcian) China The only sauropod with ossified distal tarsals, hinting at its basal position Goyocephale 1982 Unnamed formation (Late Cretaceous, Campanian) Mongolia Had a sloping head with a flat skull roof Graciliceratops 2000 Bayan Shireh Formation (Late Cretaceous, Santonian) Mongolia Possessed a short frill with large fenestrae Graciliraptor 2004 Yixian Formation (Early Cretaceous, Barremian) China A close relative of Microraptor with characteristically slender bones Guanlong 2006 Shishugou Formation (Late Jurassic, Oxfordian) China Two specimens have been discovered, one on top of the other Halszkaraptor 2017 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Originally interpreted as a semiaquatic fish hunter similar to a merganser[51] but this hypothesis has been criticized[52] Hamititan 2021 Shengjinkou Formation (Early Cretaceous, Aptian) China Known from seven caudal vertebrae and associated elements Haplocheirus 2010 Shishugou Formation (Late Jurassic, Oxfordian) China Possessed three long fingers with short claws. Originally described as a basal alvarezsauroid but similarities have been noted with other coelurosaurs[14][53] Harpymimus 1984 Khuren Dukh Formation?/Shinekhudag Formation? (Early Cretaceous, Albian) Mongolia Mostly toothless but retains a few teeth in the dentary Haya 2011 Javkhlant Formation (Late Cretaceous, Santonian to Campanian) Mongolia One specimen preserves a large mass of gastroliths Heishansaurus 1953 Minhe Formation (Late Cretaceous, Campanian to Maastrichtian) China May be a junior synonym of Pinacosaurus[54] Helioceratops 2009 Quantou Formation (Early Cretaceous to Late Cretaceous, Aptian to Cenomanian) China Had a distinctively short lower jaw Hexing 2012 Yixian Formation (Early Cretaceous, Valanginian to Barremian) China Three or four teeth are known, but they are not well-preserved Hexinlusaurus 2005 Shaximiao Formation (Middle Jurassic, Bajocian) China Originally named as a species of Yandusaurus Heyuannia 2002 Barun Goyot Formation, Dalangshan Formation (Late Cretaceous, Maastrichtian) China
Mongolia Fossilized pigments in referred eggshells suggest they were blue-green[55] Homalocephale 1974 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Has been suggested to be a juvenile Prenocephale on account of its flat head,[56] but this is no longer thought to be the case[57] Huabeisaurus 2000 Huiquanpu Formation (Late Cretaceous, Cenomanian to Maastrichtian) China May be closely related to Tangvayosaurus[58] Hualianceratops 2015 Shishugou Formation (Late Jurassic, Oxfordian) China Had a series of bumps around the edge of the beak Huanansaurus 2015 Nanxiong Formation (Late Cretaceous, Campanian to Maastrichtian) China Possessed a distinctive short trapezoidal crest Huanghetitan 2006 Haoling Formation, Hekou Group (Early Cretaceous, Aptian to Albian) China Had ribs 3 metres (9.8 ft) long, which supported one of the deepest body cavities of any dinosaur[59] Huangshanlong 2014 Hongqin Formation (Middle Jurassic to Late Jurassic, Aalenian to Oxfordian) China Known from some bones of the right forelimb Huaxiagnathus 2004 Yixian Formation (Early Cretaceous, Aptian) China One of the largest known compsognathids Huayangosaurus 1982 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China Possessed flank osteoderms and a small tail club in addition to plates and spikes Hudiesaurus 1997 Kalaza Formation (Late Jurassic, Tithonian) China Had a butterfly-shaped process on its vertebra Hulsanpes 1982 Barun Goyot Formation (Late Cretaceous, Campanian) Mongolia Closely related to Halszkaraptor but appears to be more cursorial[60] Ichthyovenator 2012 Grès supérieurs Formation (Early Cretaceous, Aptian) Laos One of its sacral vertebrae was greatly reduced, giving the illusion of a break in its sail or of two separate sails Incisivosaurus 2002 Yixian Formation (Early Cretaceous, Barremian) China Two specimens of different ontogenetic stages are known, both with differing types of feathers[61] Irisosaurus 2020 Fengjiahe Formation (Early Jurassic, Hettangian) China Closely related to Mussaurus[62] Isanosaurus 2000 Nam Phong Formation (Late Triassic, Norian to Rhaetian) Thailand May have actually come from the Late Jurassic[63] Ischioceratops 2015 Wangshi Group (Late Cretaceous, Campanian to Maastrichtian) China Noted for its peculiarly-shaped ischium Itemirus 1976 Bissekty Formation (Late Cretaceous, Turonian) Uzbekistan Originally known from a braincase but abundant new remains were described in 2014[64] Jaculinykus 2023 Barun Goyot Formation (Late Cretaceous, Maastrichtian) Mongolia Was didactyl, with a large first finger and a reduced second finger Jaxartosaurus 1937 Dabrazhin Formation (Late Cretaceous, Santonian) Kazakhstan Not known from many remains but they are enough to tell that it was a basal lambeosaurine[65] Jeholosaurus 2000 Yixian Formation (Early Cretaceous, Aptian) China One specimen is preserved in a curled position Jianchangosaurus 2013 Yixian Formation (Early Cretaceous, Barremian) China Several characters of its teeth and jaws are convergently similar to those of ornithischians[66] Jiangjunosaurus 2007 Shishugou Formation (Late Jurassic, Oxfordian) China Had two rows of circular or diamond-shaped plates Jiangshanosaurus 2001 Jinhua Formation (Late Cretaceous, Turonian to Coniacian) China A potential member of the Euhelopodidae[67] Jiangxisaurus 2013 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Overall similar to Heyuannia but with a thinner, frailer mandible Jiangxititan 2023 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Described as one of the few known lognkosaurs from mainland Asia Jianianhualong 2017 Yixian Formation (Early Cretaceous, Aptian) China Possessed a subtriangular tail frond made of asymmetrical feathers, although it was most likely flightless Jinbeisaurus 2019 Huiquanpu Formation (Late Cretaceous, Cenomanian to Maastrichtian) China A medium-sized tyrannosauroid Jinfengopteryx 2005 Huajiying Formation (Early Cretaceous, Barremian) China May have been capable of some sort of flight[68] Jingshanosaurus 1995 Lufeng Formation (Early Jurassic, Hettangian) China One of the latest-surviving non-sauropod sauropodomorphs Jintasaurus 2009 Xinminbao Group (Early Cretaceous, Albian) China Known from only the rear half of a skull, including a complete braincase Jinyunpelta 2018 Liangtoutang Formation (Early Cretaceous to Late Cretaceous, Albian to Cenomanian) China The oldest ankylosaurid known to have a tail club Jinzhousaurus 2001 Yixian Formation (Early Cretaceous, Aptian) China Its holotype is nearly complete, preserved whole on a single slab Jiutaisaurus 2006 Quantou Formation (Early Cretaceous to Late Cretaceous, Barremian to Cenomanian) China Named based on eighteen vertebrae Kaijiangosaurus 1984 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China Potentially synonymous with other medium-sized Shaximiao theropods Kamuysaurus 2019 Hakobuchi Formation (Late Cretaceous, Maastrichtian) Japan Informally referred to as "Mukawaryu" before its formal description Kansaignathus 2021 Ialovachsk Formation (Late Cretaceous, Santonian) Tajikistan The first non-avian dinosaur described from Tajikistan Kazaklambia 2013 Dabrazhin Formation (Late Cretaceous, Santonian) Kazakhstan Morphologically distinct from other Eurasian lambeosaurines[69] Kelmayisaurus 1973 Lianmuqin Formation (Early Cretaceous, Valanginian to Albian) China One popular book mentions a giant species belonging to this genus,[70] but this referral may be incorrect Kerberosaurus 2004 Tsagayan Formation (Late Cretaceous, Maastrichtian) Russia Potentially a close relative of Edmontosaurus[71] Khaan 2001 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Two morphotypes of chevrons are known, which may be a sexually dimorphic trait[72] Khulsanurus 2021 Barun Goyot Formation (Late Cretaceous, Campanian) Mongolia Contemporary with Parvicursor but can be distinguished by characters of its caudal vertebrae[73] Kileskus 2010 Itat Formation (Middle Jurassic, Bathonian) Russia Uncertain if it possesses the head crest as seen in other proceratosaurids Kinnareemimus 2009 Sao Khua Formation (Early Cretaceous, Valanginian to Barremian) Thailand Potentially one of the oldest ornithomimosaurs Klamelisaurus 1993 Shishugou Formation (Middle Jurassic, Callovian) China Close relatives included several referred species of Mamenchisaurus[74] Kol 2009 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Had a "hyperarctometatarsus" more strongly pinched than other arctometatarsalian taxa. Described as an alvarezsaurid[75] but has been suggested to be related to Avimimus[76] Koreaceratops 2011 Sihwa Formation (Early Cretaceous, Albian) South Korea Possessed elongated neural spines on its caudal vertebrae. Its describers suggest that it was used as a swimming organ,[77] but a later study found it to live in a semiarid environment, making this unlikely[78] Koreanosaurus 2011 Seonso Conglomerate (Late Cretaceous, Campanian) South Korea Had short but powerful forelimbs suggesting it may have been a quadruped[79] Koshisaurus 2015 Kitadani Formation (Early Cretaceous, Hauterivian) Japan Distinguished from other hadrosauroids by the presence of an antorbital fossa Kulceratops 1995 Khodzhakul Formation (Early Cretaceous, Albian) Uzbekistan Only known from fragments of a jaw and teeth Kulindadromeus 2014 Ukureyskaya Formation (Middle Jurassic, Bathonian) Russia An ornithischian that preserves evidence of filaments, suggesting that protofeathers were basal to Dinosauria as a whole Kundurosaurus 2012 Udurchukan Formation (Late Cretaceous, Maastrichtian) Russia May be synonymous with Kerberosaurus[80] Kuru 2021 Barun Goyot Formation (Late Cretaceous, Maastrichtian) Mongolia Had been informally referred to as "Airakoraptor" prior to its formal description Laiyangosaurus 2019 Wangshi Group (Late Cretaceous, Maastrichtian) China Some specimens referred to this edmontosaurin actually belong to kritosaurins and lambeosaurines[81] Lanzhousaurus 2005 Hekou Group (Early Cretaceous, Barremian) China Possessed the largest known teeth of any dinosaur Leshansaurus 2009 Shaximiao Formation (Late Jurassic, Oxfordian to Kimmeridgian) China Its braincase is nearly identical to that of Piveteausaurus[82] Levnesovia 2009 Bissekty Formation (Late Cretaceous, Turonian) Uzbekistan One of the smallest known hadrosauroids[42] Liaoceratops 2002 Yixian Formation (Early Cretaceous, Barremian) China One specimen was found without a skull roof, possibly displaced by a predator to eat its brain[83] Liaoningosaurus 2001 Yixian Formation (Early Cretaceous, Barremian to Aptian) China One specimen has been interpreted as possessing fork-like teeth, sharp claws, and stomach contents including fish, which has been claimed to be evidence of a semi-aquatic, turtle-like lifestyle[84] Liaoningotitan 2018 Yixian Formation (Early Cretaceous, Barremian) China The second sauropod named from the Yixian Formation Liaoningvenator 2017 Yixian Formation (Early Cretaceous, Barremian) China Uniquely preserved with the head curving forwards, differing from the classic theropod "death pose" and the sleeping position of other troodontids Limusaurus 2009 Shishugou Formation (Late Jurassic, Oxfordian) China Multiple specimens from different growth stages are known. Juveniles possessed teeth which were lost and replaced with a beak as adults, suggesting a change in diet[85] Lingwulong 2018 Yanan Formation?/Zhiluo Formation? (Middle Jurassic to Late Jurassic, Aalenian to Oxfordian) China The first confirmed diplodocoid from Asia. Originally considered Early Jurassic, making it the oldest known neosauropod, but this age has been disputed[86][87] Lingyuanosaurus 2019 Jiufotang Formation?/Yixian Formation? (Early Cretaceous, Valanginian to Aptian) China Possessed a mix of basal and derived therizinosaurian traits Linhenykus 2011 Bayan Mandahu Formation (Late Cretaceous, Campanian to Maastrichtian) China Completely monodactyl due to lacking the vestigial second and third fingers of other alvarezsaurids Linheraptor 2010 Bayan Mandahu Formation (Late Cretaceous, Campanian) China Potentially a synonym of Tsaagan[88] Linhevenator 2011 Bayan Mandahu Formation (Late Cretaceous, Campanian) China Had a greatly enlarged sickle claw, comparable in size to those of dromaeosaurids Liubangosaurus 2010 Xinlong Formation (Early Cretaceous, Barremian to Aptian) China Described only as a eusauropod[89] but has since been reinterpreted as a somphospondylian[90] Luanchuanraptor 2007 Qiupa Formation (Late Cretaceous, Maastrichtian) China The first Asian dromaeosaurid found outside the Gobi Desert and northeastern China. May have been closely related to Adasaurus[14] Lufengosaurus 1940 Lufeng Formation (Early Jurassic, Hettangian to Sinemurian) China The rib of one specimen preserves the oldest known evidence of collagen proteins[91] Luoyanggia 2009 Haoling Formation (Early Cretaceous, Aptian to Albian) China Originally believed to date from the Late Cretaceous Machairasaurus 2010 Bayan Mandahu Formation (Late Cretaceous, Campanian) China Its hand claws are elongated and blade-like in side view Mahakala 2007 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Possessed basal traits for a dromaeosaurid. May be a close relative of Halszkaraptor[92] Maleevus 1987 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia Its only purportedly distinguishing trait is also shared with Pinacosaurus[36] Mamenchisaurus 1954 Penglaizhen Formation, Shaximiao Formation, Shishugou Formation, Suining Formation (Late Jurassic to Early Cretaceous, Oxfordian to Aptian) China Several species have been named, but most may not belong to this genus[74] Mandschurosaurus 1930 Grès supérieurs Formation?, Yuliangze Formation (Late Cretaceous, Maastrichtian) China
Laos? One of the first non-avian dinosaurs named from Chinese remains Mei 2004 Yixian Formation (Early Cretaceous, Aptian) China Two specimens are preserved in bird-like sleeping positions[93] Microceratus 2008 Ulansuhai Formation (Late Cretaceous, Turonian) China Originally named Microceratops, although that genus name is preoccupied by a wasp Microhadrosaurus 1979 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Reportedly an unusually small hadrosaurid Micropachycephalosaurus 1978 Wangshi Group (Late Cretaceous, Campanian to Maastrichtian) China Once considered to be a pachycephalosaur, although it is now usually considered to be a ceratopsian[94] Microraptor 2000 Jiufotang Formation (Early Cretaceous, Aptian) China Known from over three hundred fossils.[95] Several are well-preserved enough to reveal fine details such as feather covering and an iridescent black coloration[96] Migmanychion 2023 Longjiang Formation (Early Cretaceous, Aptian) China Its hand combines features of multiple groups of coelurosaurs Minimocursor 2023 Phu Kradung Formation (Late Jurassic, Tithonian) Thailand The first basal neornithischian known from southeastern Asia Minotaurasaurus 2009 Djadochta Formation (Late Cretaceous, Campanian) Mongolia The holotype skull was excavated illegally, which obscured its true provenance until recently Mongolosaurus 1933 On Gong Formation (Early Cretaceous, Aptian to Albian) China Known from only scant remains but has been confidently assigned to Somphospondyli in recent years[90] Mongolostegus 2018 Dzunbain Formation (Early Cretaceous, Aptian to Albian) Mongolia Informally assigned to the genus Wuerhosaurus before its formal description Monkonosaurus 1986 Loe-ein Formation?/Lura Formation? (Late Jurassic, Oxfordian to Kimmeridgian?/Early Cretaceous, Albian?) China Poorly known Monolophosaurus 1993 Shishugou Formation (Middle Jurassic, Bathonian to Callovian) China Possessed a short, rectangular crest running along the midline of the skull Mononykus 1993 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Proposed to have an anteater-like lifestyle, using its unique forearms to break into termite mounds[97] Mosaiceratops 2015 Xiaguan Formation (Late Cretaceous, Turonian to Campanian) China Combined features of different groups of basal ceratopsians Nankangia 2013 Nanxiong Formation (Late Cretaceous, Maastrichtian) China May have specialized in soft foods such as leaves and seeds[98] Nanningosaurus 2007 Unnamed formation (Late Cretaceous, Maastrichtian) China Potentially a basal lambeosaurine Nanshiungosaurus 1979 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Originally misidentified as a sauropod on account of its unusual pelvis Nanyangosaurus 2000 Xiaguan Formation (Late Cretaceous, Turonian to Campanian) China Completely lost the first digit of its hands Napaisaurus 2022 Xinlong Formation (Early Cretaceous, Aptian to Albian) China May be closely related to contemporary Thai iguanodonts Natovenator 2022 Barun Goyot Formation (Late Cretaceous, Maastrichtian) Mongolia Possessed a streamlined body and a long, toothed snout, convergently similar to several groups of aquatic vertebrates Nebulasaurus 2015 Zhanghe Formation (Middle Jurassic, Aalenian to Bajocian) China Only known from a single braincase, but it is enough to tell that it was related to Spinophorosaurus Neimongosaurus 2001 Iren Dabasu Formation (Late Cretaceous, Cenomanian) China Could extend its arms considerably forward due to the structure of its shoulder joint[99] Nemegtomaia 2005 Barun Goyot Formation, Nemegt Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia One specimen preserves traces of damage by skin beetles[100] Nemegtonykus 2019 Nemegt Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia The second alvarezsaurid named from the Nemegt Formation Nemegtosaurus 1971 Nemegt Formation, Subashi Formation? (Late Cretaceous, Maastrichtian) China?
Mongolia Had a long, low skull similar in proportions to those of diplodocoids Ningyuansaurus 2012 Yixian Formation (Early Cretaceous, Aptian) China Preserves small oval-shaped structures in its stomach region which may be seeds Nipponosaurus 1936 Yezo Group (Late Cretaceous, Santonian to Campanian) Russia Discovered on the island of Sakhalin, which was owned by Japan in 1936 but later annexed by Russia Oksoko 2020 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Its third finger was so greatly reduced that it was functionally didactyl Olorotitan 2003 Udurchukan Formation (Late Cretaceous, Maastrichtian) Russia Had a broad, hatchet-shaped crest Omeisaurus 1939 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China Members of this genus are characterized by extremely elongated necks Ondogurvel 2022 Barun Goyot Formation (Late Cretaceous, (Campanian) Mongolia Known from well-preserved remains of the hands and feet Opisthocoelicaudia 1977 Nemegt Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia Walked on its metacarpals due to its complete lack of phalanges Oviraptor 1924 Djadochta Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia Originally mistakenly thought to be an egg-eater Pachysuchus 1951 Lufeng Formation (Early Jurassic, Sinemurian to Pliensbachian) China Considered a phytosaur from its original naming until a redescription in 2012[101] Panguraptor 2014 Lufeng Formation (Early Jurassic, Hettangian to Sinemurian) China The first definitive coelophysoid known from Asia Papiliovenator 2021 Bayan Mandahu Formation (Late Cretaceous, Campanian) China Had a short, subtriangular skull similar to those of Early Cretaceous troodontids Paralitherizinosaurus 2022 Yezo Group (Late Cretaceous, Campanian Japan Had stiffened claws that may have been used to pull vegetation to the mouth[102] Parvicursor 1996 Barun Goyot Formation (Late Cretaceous, Campanian) Mongolia Originally believed to represent a diminutive adult dinosaur, although it was recently reinterpreted as a juvenile[103] Pedopenna 2005 Haifanggou Formation (Middle Jurassic, Callovian) China Known from a single leg with the impressions of long, symmetrical feathers Peishansaurus 1953 Minhe Formation (Late Cretaceous, Santonian to Campanian) China Has been compared to thyreophorans and marginocephalians, but it is impossible to determine which assignment is correct Penelopognathus 2005 Bayin-Gobi Formation (Early Cretaceous, Albian) China Named from a single dentary Phaedrolosaurus 1973 Lianmuqin Formation (Early Cretaceous, Valanginian to Albian) China May have been a dromaeosaurid[104] Philovenator 2012 Bayan Mandahu Formation (Late Cretaceous, Campanian) China Closely related to the contemporary Linhevenator[93] but likely represents a separate taxon[105] Phuwiangosaurus 1994 Sao Khua Formation (Early Cretaceous, Valanginian to Hauterivian) Thailand A large member of the Euhelopodidae[90] Phuwiangvenator 2019 Sao Khua Formation (Early Cretaceous, Barremian) Thailand Combines features of both allosauroids and coelurosaurs[106] Pinacosaurus 1933 Bayan Mandahu Formation, Djadochta Formation (Late Cretaceous, Santonian to Campanian) China
Mongolia May have been capable of bird-like vocalizations[107] Plesiohadros 2014 Alagteeg Formation (Late Cretaceous, Campanian) Mongolia The first hadrosauroid known from the Alagteeg Formation Prenocephale 1974 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Had a distinctively conical dome Probactrosaurus 1966 Dashuigou Formation (Early Cretaceous, Albian) China The closest relative to the Hadrosauromorpha based on the definition of the group[108] Prodeinodon 1924 Öösh Formation, Xinlong Formation (Early Cretaceous, Barremian to Aptian) China
Mongolia Potentially a carnosaur[109] Protarchaeopteryx 1997 Yixian Formation (Early Cretaceous, Aptian) China Usually thought to be a basal oviraptorosaur but one study suggests a basal position within Pennaraptora[14] Protoceratops 1923 Bayan Mandahu Formation, Djadochta Formation (Late Cretaceous, Campanian) China
Mongolia Its remains are so abundant that it has been nicknamed the "sheep of the Cretaceous" Protognathosaurus 1991 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China Originally named Protognathus, but that is preoccupied by an extinct beetle[110] Psittacosaurus 1923 Andakhuduk Formation, Bayin-Gobi Formation, Ejinhoro Formation, Ilek Formation, Jiufotang Formation, Khok Kruat Formation, Öösh Formation, Qingshan Formation, Tugulu Group, Xinminbao Group, Yixian Formation (Early Cretaceous, Barremian to Albian) China
Mongolia
Russia
Thailand Known from hundreds of specimens, many of them well-preserved. Lived in a broad range Pukyongosaurus 2001 Hasandong Formation (Early Cretaceous, Aptian to Albian) South Korea One of its caudal vertebrae has bite marks caused by theropod teeth Qianlong 2023 Ziliujing Formation (Early Jurassic, Sinemurian) China Associated with fossils of leathery eggs, the oldest of their kind in the world Qianzhousaurus 2014 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Has been nicknamed "Pinocchio rex" on account of its elongated snout Qiaowanlong 2009 Xinminbao Group (Early Cretaceous, Albian) China Originally described as a brachiosaurid[111] but has since been reinterpreted as a euhelopodid[112] Qijianglong 2015 Suining Formation (Early Cretaceous, Aptian) China Once believed to date from the Late Jurassic Qingxiusaurus 2008 Unnamed formation (Late Cretaceous, Maastrichtian) China Known from very limited remains Qinlingosaurus 1996 Hongtuling Formation?/Shanyang Formation? (Late Cretaceous, Maastrichtian) China Potentially a titanosaur given its age, but this cannot be confirmed Qiupalong 2011 Qiupa Formation (Late Cretaceous, Campanian to Maastrichtian) China A referred specimen was found in Canada[113] Qiupanykus 2018 Qiupa Formation (Late Cretaceous, Maastrichtian) China May have used its robust thumb claws to crack open oviraptorid eggshells[114] Quaesitosaurus 1983 Barun Goyot Formation (Late Cretaceous, Maastrichtian) Mongolia Potentially a close relative of Nemegtosaurus Ratchasimasaurus 2011 Khok Kruat Formation (Early Cretaceous, Aptian) Thailand Only known from a single toothless dentary Rhomaleopakhus 2021 Kalaza Formation (Late Jurassic, Tithonian) China Possessed a robust forelimb that may be a locomotory adaptation Rinchenia 1997 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Had a tall, domed crest Ruixinia 2022 Yixian Formation (Early Cretaceous, Barremian) China Its last few caudal vertebrae were fused into a rod-like structure Ruyangosaurus 2009 Haoling Formation (Early Cretaceous, Aptian to Albian) China Only known from scant remains but was one of the largest dinosaurs known from Asia Sahaliyania 2008 Yuliangze Formation (Late Cretaceous, Maastrichtian) China Possibly a synonym of Amurosaurus[115] Saichania 1977 Barun Goyot Formation, Nemegt Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia Possessed complicated nasal passages that may have cooled the air it breathed Sanpasaurus 1944 Ziliujing Formation (Early Jurassic, Toarcian) China Historically conflated with the remains of an ornithischian Sanxiasaurus 2019 Xintiangou Formation (Middle Jurassic, Bajocian) China The oldest neornithischian known from Asia Saurolophus 1912 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Type species was found in Canada. The Asian species is distinguished by its larger size and backwards-pointing diagonal crest Sauroplites 1953 Zhidan Group (Early Cretaceous, Barremian to Aptian) China Preserved lying on its back with parts of its armor in an articulated position Saurornithoides 1924 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Its hindlimbs were well-developed even as juveniles, suggesting it needed little to no parental care Scansoriopteryx 2002 Haifanggou Formation (Middle Jurassic to Late Jurassic, Callovian to Kimmeridgian) China Was well-adapted for climbing due to the structure of its hands and feet Segnosaurus 1979 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Turonian) Mongolia One of the first known therizinosaurs. Its relationships were originally obscure Serikornis 2017 Tiaojishan Formation (Middle Jurassic to Late Jurassic, Callovian to Oxfordian) China Possessed simple, wispy feathers similar to those of a Silkie chicken Shamosaurus 1983 Dzunbain Formation (Early Cretaceous, Aptian to Albian) Mongolia The osteoderms on its head were not separated into obvious tiles as with later ankylosaurs Shanag 2007 Öösh Formation (Early Cretaceous, Berriasian to Barremian) Mongolia Shows a mixture of traits of various paravian groups Shantungosaurus 1973 Wangshi Group (Late Cretaceous, Campanian) China The largest known hadrosaurid Shanxia 1998 Huiquanpu Formation (Late Cretaceous, Cenomanian to Campanian) China May be synonymous with Tianzhenosaurus[116] and/or Saichania[36] Shanyangosaurus 1996 Shanyang Formation (Late Cretaceous, Maastrichtian) China Indeterminate but its hollow bones are a synapomorphy for Coelurosauria. One study suggests an oviraptorosaurian position[14] Shaochilong 2009 Ulansuhai Formation (Late Cretaceous, Cenomanian to Turonian) China Had a relatively short maxilla, suggesting a unique ecological role Shenzhousaurus 2003 Yixian Formation (Early Cretaceous, Aptian) China Preserves pebbles in its thoracic cavity which may be gastroliths Shidaisaurus 2009 Chuanjie Formation (Middle Jurassic, Aalenian) China Potentially one of the oldest known allosauroids Shishugounykus 2019 Shishugou Formation (Late Jurassic, Oxfordian) China Its manus combines features of both alvarezsaurians and more basal coelurosaurs Shixinggia 2005 Pingling Formation (Late Cretaceous, Maastrichtian) China Known from a fair amount of postcranial material Shri 2021 Barun Goyot Formation (Late Cretaceous, Maastrichtian) Mongolia Before its formal description, it was nicknamed "Ichabodcraniosaurus" because its holotype lacked a skull Shuangmiaosaurus 2003 Sunjiawan Formation (Early Cretaceous, Albian) China Only known from some parts of a skull Shunosaurus 1983 Shaximiao Formation (Late Jurassic, Oxfordian to Kimmeridgian) China Possessed a small tail club topped by two short spikes Shuvuuia 1998 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Displays several adaptations that may point to a nocturnal, owl-like lifestyle[117] Siamodon 2011 Khok Kruat Formation (Early Cretaceous, Aptian) Thailand May have been closely related to Probactrosaurus[118] Siamosaurus 1986 Khok Kruat Formation, Sao Khua Formation (Early Cretaceous, Barremian to Aptian) Thailand Only known from teeth. Some spinosaurid postcrania from the same area may be referrable to this genus[119] Siamotyrannus 1996 Sao Khua Formation (Early Cretaceous, Berriasian to Barremian) Thailand Has been recovered in a variety of positions within Avetheropoda Sinankylosaurus 2020 Wangshi Group (Late Cretaceous, Campanian) China Only known from an ilium. Described as an ankylosaur but a recent study doubts this interpretation[120] Sinocalliopteryx 2007 Yixian Formation (Early Cretaceous, Barremian to Aptian) China Stomach contents indicate a possible preference for volant prey such as dromaeosaurids and early birds[121] Sinocephale 2021 Ulansuhai Formation (Late Cretaceous, Turonian) China Originally named as a species of Troodon when that genus was thought to be a pachycephalosaur Sinoceratops 2010 Wangshi Group (Late Cretaceous, Campanian to Maastrichtian) China Possessed forward-curving hornlets and a series of low knobs on the top of the frill Sinocoelurus 1942 Kuangyuan Series (Late Jurassic, Oxfordian to Tithonian China One study considered it to be a potential plesiosaur[122] Sinornithoides 1993 Ejinhoro Formation (Early Cretaceous, Aptian to Albian) China Preserved in a roosting position, its head tucked underneath its left wing Sinornithomimus 2003 Ulansuhai Formation (Late Cretaceous, Turonian) China Formed age-segregated herds as evidenced by a concentration of juvenile skeletons[123] Sinornithosaurus 1999 Yixian Formation (Early Cretaceous, Barremian to Aptian) China One specimen has disloged teeth, leading to suggestions it was venomous[124] Sinosauropteryx 1996 Yixian Formation (Early Cretaceous, Barremian) China The first non-avian dinosaur found with direct evidence of feathers. Analysis of melanosomes suggest it had orange-brown and white countershading with a striped tail and a "bandit mask" around its eyes[125] Sinosaurus 1940 Lufeng Formation (Early Jurassic, Hettangian to Sinemurian) China Had a pair of midline crests similar to Dilophosaurus Sinotyrannus 2009 Jiufotang Formation (Early Cretaceous, Aptian) China One of the earliest known large tyrannosauroids. Closely related to smaller forms such as Proceratosaurus and Guanlong Sinovenator 2002 Yixian Formation (Early Cretaceous, Barremian) China Some specimens are preserved three-dimensionally Sinraptor 1993 Shishugou Formation (Late Jurassic, Oxfordian) China May have used its teeth like blades to inflict deep wounds in prey Sinusonasus 2004 Yixian Formation (Early Cretaceous, Hauterivian) China Had distinctive sinusoid nasal bones Sirindhorna 2015 Khok Kruat Formation (Early Cretaceous, Aptian) Thailand Its fossils were discovered by corn farmers while digging a reservoir Sonidosaurus 2006 Iren Dabasu Formation (Late Cretaceous, Cenomanian to Campanian) China One of the smallest known titanosaurs Stegosaurides 1953 Xinminbao Group (Early Cretaceous, Hauterivian to Albian) China A thyreophoran of uncertain phylogenetic position Suzhousaurus 2007 Xinminbao Group (Early Cretaceous, Barremian to Aptian) China One of the largest Early Cretaceous therizinosaurs Szechuanosaurus 1942 Kuangyuan Series (Late Jurassic, Oxfordian to Tithonian) China Only known from teeth and possibly a very fragmentary skeleton Talarurus 1952 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia Its tail club has been compared to a wicker basket Tambatitanis 2014 Sasayama Group (Early Cretaceous, Albian) Japan Possessed disproportionately large chevrons Tangvayosaurus 1999 Grès supérieurs Formation (Early Cretaceous, Aptian to Albian) Laos Closely related to Phuwiangosaurus Tanius 1929 Wangshi Group (Late Cretaceous, Campanian to Maastrichtian) China Today known from only a few bones; more fossils were once present but were not collected Taohelong 2013 Hekou Group (Early Cretaceous, Albian) China Possessed a sacral shield similar to that of Polacanthus Tarbosaurus 1955 Nemegt Formation, Subashi Formation (Late Cretaceous, Maastrichtian) China
Mongolia An apex predator that hunted large prey. Very similar to Tyrannosaurus Tarchia 1977 Barun Goyot Formation, Nemegt Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia One specimen preserves injuries to its ribs and tail, possibly from a fight with a member of its own kind[126] Tatisaurus 1965 Lufeng Formation (Early Jurassic, Sinemurian) China Potentially a basal thyreophoran Tengrisaurus 2017 Murtoi Formation (Early Cretaceous, Barremian to Aptian) Russia Closely related to South American titanosaurs Therizinosaurus 1954 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Possessed extremely elongated and stiffened hand claws Tianchisaurus 1993 Toutunhe Formation (Late Jurassic, Oxfordian to Kimmeridgian) China Its description uses the spellings Tianchisaurus and Tianchiasaurus interchangeably, but the former is correct[127] Tianyulong 2009 Tiaojishan Formation (Late Jurassic, Oxfordian) China Preserves impressions of long bristles down its back, tail and neck Tianyuraptor 2009 Yixian Formation (Early Cretaceous, Barremian to Aptian) China Combines features of both northern and southern dromaeosaurids. Had unusual proportions Tianzhenosaurus 1998 Huiquanpu Formation (Late Cretaceous, Cenomanian to Campanian) China May be synonymous with Saichania[36] Tienshanosaurus 1937 Shishugou Formation (Late Jurassic, Oxfordian) China Large but basal for a mamenchisaurid[74] Timurlengia 2016 Bissekty Formation (Late Cretaceous, Turonian) Uzbekistan Its inner ear was specialized for detecting low-frequency sounds[128] Tochisaurus 1991 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Known from only a single metatarsus Tonganosaurus 2010 Yimen Formation (Early Jurassic, Pliensbachian) China Potentially the oldest known mamenchisaurid Tongtianlong 2016 Nanxiong Formation (Late Cretaceous, Maastrichtian) China The pose of the holotype suggests it died while trying to free itself from mud Tsaagan 2006 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Very similar to Velociraptor but differs in some features of the skull[129] Tsagantegia 1993 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia Had a long, shovel-shaped snout which may indicate a browsing lifestyle[130] Tsintaosaurus 1958 Wangshi Group (Late Cretaceous, Campanian) China Originally mistakenly believed to have possessed a unicorn horn-like crest Tugulusaurus 1973 Lianmuqin Formation (Early Cretaceous, Barremian to Albian) China Potentially an early, Xiyunykus-grade alvarezsaurian[131] Tuojiangosaurus 1977 Shaximiao Formation (Late Jurassic, Oxfordian to Kimmeridgian) China Possessed two rows of tall, pointed plates, thickened in the center as if they were modified spikes Turanoceratops 1989 Bissekty Formation (Late Cretaceous, Turonian) Uzbekistan Had a pair of brow horns like ceratopsids but was likely not a member of that family Tylocephale 1974 Barun Goyot Formation (Late Cretaceous, Campanian) Mongolia Only known from a partial skull but it is enough to tell that it had a remarkably tall dome Tyrannomimus 2023 Kitadani Formation (Early Cretaceous, Aptian) Japan Its ilium is remarkably similar to that of the supposed tyrannosauroid Aviatyrannis Udanoceratops 1992 Djadochta Formation (Late Cretaceous, Campanian) Mongolia The largest known leptoceratopsid Ultrasaurus 1983 Gugyedong Formation (Early Cretaceous, Aptian to Albian) South Korea Described as very large but this may be due to misidentification of a bone Ulughbegsaurus 2021 Bissekty Formation (Late Cretaceous, Turonian) Uzbekistan Known from only a maxilla. Originally described as a late-surviving carnosaur but may in fact be a large-bodied dromaeosaurid[132] Urbacodon 2007 Bissekty Formation, Dzharakuduk Formation (Late Cretaceous, Cenomanian to Turonian) Uzbekistan The holotype preserves a gap separating the eight rear teeth from the rest of its teeth Vayuraptor 2019 Sao Khua Formation (Early Cretaceous, Barremian) Thailand Potentially ancestral to megaraptorans[133] or an early member of the group[134] Velociraptor 1924 Bayan Mandahu Formation, Djadochta Formation (Late Cretaceous, Campanian) China
Mongolia One potential specimen preserves quill knobs[135] Wakinosaurus 1992 Sengoku Formation (Early Cretaceous, Valanginian to Barremian) Japan May be a close relative of Acrocanthosaurus[109] Wannanosaurus 1977 Xiaoyan Formation (Late Cretaceous, Maastrichtian) China Basal for a pachycephalosaur as indicated by its flat skull with large openings Wuerhosaurus 1973 Ejinhoro Formation, Tugulu Group (Early Cretaceous, Hauterivian) China One of the last and largest known stegosaurs. Preserved with low rectangular plates but these may be broken Wulagasaurus 2008 Yuliangze Formation (Late Cretaceous, Maastrichtian) China A rare hadrosaurid known from far less remains than the contemporary Sahaliyania Wulatelong 2013 Bayan Mandahu Formation (Late Cretaceous, Campanian) China Known from a partial skeleton including some parts of the skull Wulong 2020 Jiufotang Formation (Early Cretaceous, Aptian) China Analysis of preserved melanosomes suggests it was mostly gray with iridescent wings[136] Xianshanosaurus 2009 Haoling Formation (Early Cretaceous, Aptian to Albian) China May have been closely related to Daxiatitan[90] Xiaosaurus 1983 Shaximiao Formation (Middle Jurassic, Bajocian to Callovian) China An ornithischian of uncertain affinities Xiaotingia 2011 Tiaojishan Formation (Middle Jurassic to Late Jurassic, Bathonian to Oxfordian) China Well-preserved but inconsistent in phylogenetic placement. Some studies suggest a position as an early avialan[137] Xingtianosaurus 2019 Yixian Formation (Early Cretaceous, Barremian) China Retained the large third finger that was lost in other caudipterids Xingxiulong 2017 Lufeng Formation (Early Jurassic, Hettangian) China Possessed a robust scapula which increased forelimb mobility for feeding Xinjiangovenator 2005 Lianmuqin Formation (Early Cretaceous, Valanginian to Albian) China Remains originally identified as Phaedrolosaurus Xinjiangtitan 2013 Qiketai Formation (Middle Jurassic, Callovian) China Had an extremely long neck Xiongguanlong 2009 Xinminbao Group, (Early Cretaceous, Aptian) China More robust than other early tyrannosauroids, possibly to support its elongated skull Xixianykus 2010 Majiacun Formation (Late Cretaceous, Santonian to Coniacian) China One of the smallest known non-avian dinosaurs Xixiasaurus 2010 Majiacun Formation (Late Cretaceous, Coniacian to Campanian) China Distinguished from other troodontids by its possession of exactly twenty-two teeth in each maxilla Xixiposaurus 2010 Lufeng Formation (Early Jurassic, Hettangian to Toarcian) China Poorly known Xiyunykus 2018 Tugulu Group (Early Cretaceous, Barremian to Aptian) China Had an unspecialized hand morphology for an alvarezsaur, having three fingers of roughly equal length and construction Xuanhanosaurus 1984 Shaximiao Formation (Middle Jurassic to Late Jurassic, Bathonian) China Originally mistakenly believed to have been capable of quadrupedal locomotion Xuanhuaceratops 2006 Houcheng Formation (Late Jurassic, Tithonian) China Possessed a large premaxillary tooth right behind its beak Xunmenglong 2019 Huajiying Formation (Early Cretaceous, Hauterivian) China The holotype was originally presented as part of a chimera involving three different animals[138] Xuwulong 2011 Xinminbao Group (Early Cretaceous, Aptian to Albian) China The tip of its dentary was V-shaped when viewed from the side Yamaceratops 2006 Javkhlant Formation (Late Cretaceous, Santonian) Mongolia Possessed a short, stubby frill Yamatosaurus 2021 Kita-Ama Formation (Late Cretaceous, Maastrichtian) Japan Basal yet survived late enough to be contemporaneous with more advanced hadrosaurids Yandusaurus 1979 Shaximiao Formation (Middle Jurassic, Bathonian) China Some fossils were destroyed by a composter before they could be studied[139] Yangchuanosaurus 1978 Shaximiao Formation (Middle Jurassic to Late Jurassic, Bathonian to Oxfordian) China The largest theropod known from the Shaximiao Yi 2015 Tiaojishan Formation (Middle Jurassic to Late Jurassic, Callovian to Oxfordian) China Possessed a "styliform element" jutting out from its wrist that supported a bat-like membranous wing Yimenosaurus 1990 Fengjiahe Formation (Early Jurassic, Pliensbachian) China Much of its skeleton is known, including the entirety of the skull Yingshanosaurus 1994 Shaximiao Formation (Late Jurassic, Kimmeridgian) China Possessed greatly enlarged shoulder spines Yinlong 2006 Shishugou Formation (Late Jurassic, Oxfordian) China Its skull displays features of ceratopsians, pachycephalosaurs, and heterodontosaurids Yixianosaurus 2003 Yixian Formation (Early Cretaceous, Aptian) China Inconsistent in phylogenetic placement. Had extremely elongated manual elements Yizhousaurus 2018 Lufeng Formation (Early Jurassic, Sinemurian) China Its skull was very similar to those of sauropods, despite being more primitive Yongjinglong 2014 Hekou Group (Early Cretaceous, Albian) China Possessed an extremely long, broad scapula Yuanmousaurus 2006 Zhanghe Formation (Middle Jurassic, Aalenian to Callovian) China Shares features of its vertebrae with Patagosaurus Yueosaurus 2012 Liangtoutang Formation (Early Cretaceous to Late Cretaceous, Albian to Cenomanian) China Probably closely related to Jeholosaurus[140] Yulong 2013 Qiupa Formation (Late Cretaceous, Maastrichtian) China Known from multiple specimens, most of which are juveniles Yunganglong 2013 Zhumapu Formation (Late Cretaceous, Cenomanian) China Discovered 50 kilometres (31 mi) away from a World Heritage Site Yunmenglong 2013 Haoling Formation (Early Cretaceous, Barremian to Albian) China May have been exceptionally large Yunnanosaurus 1942 Fengjiahe Formation, Lufeng Formation (Early Jurassic, Sinemurian to Pliensbachian) China Its teeth were self-sharpening similar to those of sauropods, likely through convergent evolution[141] Yunyangosaurus 2020 Xintiangou Formation (Middle Jurassic to Late Jurassic, Aalenian to Oxfordian) China Potentially an early megalosauroid Yutyrannus 2012 Yixian Formation (Early Cretaceous, Aptian) China The largest known dinosaur that preserves direct evidence of feathers Yuxisaurus 2022 Fengjiahe Formation (Early Jurassic, Sinemurian to Toarcian) China Had more than one hundred osteoderms Yuzhoulong 2022 Shaximiao Formation (Middle Jurassic, Bathonian) China One of the oldest known macronarians Zanabazar 2009 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Originally named as a species of Saurornithoides. A large troodontid Zaraapelta 2014 Barun Goyot Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia Had an intricate pattern of osteoderms on its skull Zhanghenglong 2014 Majiacun Formation (Late Cretaceous, Santonian) China Reconstructed by its describers with a straight, rectangular back, although no complete neural spines are known[142] Zhejiangosaurus 2007 Chaochuan Formation (Late Cretaceous, Cenomanian) China Has no diagnostic features[36] Zhenyuanlong 2015 Yixian Formation (Early Cretaceous, Aptian) China Possessed large wings with long feathers, but was most likely flightless Zhongjianosaurus 2017 Yixian Formation (Early Cretaceous, Barremian to Aptian) China Distinguishable by its characteristically elongated legs. Described as a microraptorian[143] but it has been noted that some features of its skeleton are similar to avialans[39] Zhuchengceratops 2010 Wangshi Group (Late Cretaceous, Maastrichtian) China Had a particularly deep mandible Zhuchengtitan 2017 Wangshi Group (Late Cretaceous, Campanian) China The proportions of its humerus suggest a close relationship with Opisthocoelicaudia[144] Zhuchengtyrannus 2011 Wangshi Group (Late Cretaceous, Campanian) China Closely related to Tarbosaurus and Tyrannosaurus Zigongosaurus 1976 Shaximiao Formation (Middle Jurassic to Late Jurassic, Bathonian to Tithonian) China May be a species of Mamenchisaurus[145] Zizhongosaurus 1983 Ziliujing Formation (Early Jurassic, Toarcian) China Poorly known but was most likely basal for a sauropod Zuolong 2010 Shishugou Formation (Late Jurassic, Oxfordian China Known from both cranial and postcranial remains
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Struthiosaurinae
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Struthiosaurinae is a subfamily of ankylosaurian dinosaurs from the Cretaceous of Europe. It is defined as "the most inclusive clade containing Europelta but not Cedarpelta, Peloroplites, Sauropelta or Edmontonia" while being reinstated for a newly recognized clade of basal nodosaurids. Struthiosaurinae appeared at about exactly the same time as the North American subfamily Nodosaurinae. Struthiosaurines range all across the Cretaceous, the oldest genus being Europelta at an age of 112 Ma and the youngest being Struthiosaurus at about 85â66 Ma.
It was originally mentioned by Franz Nopcsa in 1923 as a subfamily of Acanthopholidae, along with the previously defined Acanthopholinae. The family has gone through many taxonomic revisions since it was defined by Nopcsa in 1902. It is now recognized as a junior synonym of the family Nodosauridae. The subfamily now includes the genera Anoplosaurus, Europelta, Hungarosaurus, and Struthiosaurus, designated as the type genus. Because of the instability of Acanthopholis, the generic namesake of Acanthopholinae, and its current identification as a nomen dubium, Struthiosaurinae, the next named group, was decidedly used over the older one.
A review of ankylosaur osteoderms was published in 2000, and reviewed the armour of Struthiosaurinae. The group was represented by the single genus Struthiosaurus, known from head, cervical, dorsal, sacral, and caudal scutes. Only a few head osteoderms were identified, so it is unknown how much of the skull was armoured. Many cervical and dorsal scutes have been preserved alongside species of Struthiosaurus. They include cervical bands, which are groups of osteoderms fused together and attached to the vertebrae, and large spines found on the shoulders of nodosaurids like Sauropelta and Edmontonia, although it is not known if the spines were fused like the later of separate like the former. It is quite possible that small ovoid scutes found on Struthiosaurus could have formed a pelvic shield like polacanthids. The caudal scutes of struthiosaurines are small and rough. Even though osteoderms are well-known, it is not certain where they were positioned on the body.
Classification
Struthiosaurinae is a group named by Franz Nopcsa, that was reinstated by James Kirkland for a group of just-european nodosaurids.[1]
History
Baron Franz Nopcsa, in 1902, proposed a new grouping of dinosaurs, Acanthopholididae, a clade of lightly-built thyreophorans. He included in it the genera Acanthopholis, Polacanthus, Syngonosaurus, Struthiosaurus, and possibly Nodosaurus. Soon after, he also added Priodontognathus and Palaeoscincus. He considered Acanthopholididae to be the sister clade to Stegosauridae, and also stated that it might include Hylaeosaurus, which he found had most of the characteristics of the family.[2] Later, in 1915, he rearranged the species included in it. The genera that were later included were Acanthopholis (=Anoplosaurus), Polacanthus, Stegopelta, Stegoceras, and Struthiosaurus. Along with Ankylosaurus, Acanthopholididae was defined to form a subfamily of Nodosauridae.[3] In 1923 he divided up the family into two subfamilies without comment. These two subfamilies were named Acanthopholinae and Struthiosaurinae.[1][4]
In the same year, he corrected the subgroups of Thyreophora. He placed Acanthopholididae, Stegosauridae and Ceratopsidae together inside the group.[5] Five years later, he corrected the name of the family to Acanthopholidae, which is now the correct spelling. In the same publication, he also changed the genera included. He assigned Hylaeosaurus, Stegoceras, Struthiosaurus, and Troodon[6] inside it but moved Stegopelta and Ankylosaurus into the family Ankylosauridae.[7] Nopcsa then downgraded the phylogenetic rank of Acanthopholidae to make it a subfamily inside Nodosauridae. The clade was defined to include Ankylosaurus and Acanthopholinae. Inside Acanthopholinae he placed Acanthopholis, Hylaeosaurus, Rhodanosaurus, Struthiosaurus, and Troodon.[8] Now considered as an artificial grouping, it was defined to include dinosaur taxa now considered to be polacanthids, a pachycephalosaur and Acanthopholis, a genus that is widely considered to be dubious. Acanthopholidae and Acanthopholinae are now dubious groups since the validity of Acanthopholis has changed.[1]
Phylogeny
Historically, Struthiosaurinae has been considered a junior synonym of the family Nodosauridae by Walter P. Coombs in 1978 and Tatyana Tumanova in 1987,[9][10] or a sister clade to Edmontoniinae, Panoplosaurinae, Stegopeltinae, and Sauropeltinae by Tracy L. Ford in 2000.[11] Kirkland and his colleagues followed Ford in using Struthiosaurinae as a clade of basal nodosaurids, but concluded that the only other subfamily of Nodosauridae was Nodosaurinae. According to Kirkland et al., Acanthopholinae was not an acceptable classification for the new clade of previously unrecognized nodosaurids because of the instability of Acanthopholis. Struthiosaurinae was decided on as the name of the clade, as it was the next published term after Acanthopholinae. To ensure the group was rendered valid, Kirkland et al. redefined Struthiosaurinae as the most inclusive clade containing Europelta but not Cedarpelta, Peloroplites, Sauropelta, or Edmontonia. This definition includes the genera Anoplosaurus, Europelta, Hungarosaurus and Struthiosaurus inside the newly defined group.[1] Below is a cladogram from before the recognition of the clade Struthiosaurinae.[12]
Nodosauridae
Struthiosaurus
Hungarosaurus
Silvisaurus
Sauropelta
Pawpawsaurus
Edmontonia
Panoplosaurus
Description
Struthiosaurines are well-known, and include one of the best-preserved species of ankylosaur, Europelta carbonensis.[1]
Osteoderms
In 2000, Ford published a complete description of ankylosaurian osteoderms, in which he recognized the group Struthiosaurinae. Ford's description of Struthiosaurinae was based on the genus Struthiosaurus. Ford found that Struthiosaurus transylvanicus lacked any remains of the jugal, which makes a jugal scute unknown. The skull roof of T. transylvanicus is large and bulbous, preserving a large, flat scute on top, and no osteoderms behind the orbits. Another species, S. austriacus, is known from two incomplete skulls, which preserved irregular scutes parallel to the orbits along the cranium.[11]
Scutes from the postcranial region of the skeleton are also known from struthiosaurines. Cervical bands have been found on S. austriacus, as well as S. sp.. The cervical bands are preserved as a groups of two or three osteoderms that are fused with a large neural spine on the medial edge, and attached to each other through small ovular scutes with short rounded peaks. The scute attaching to the neural spine has a round ridge with a shallow depression ovular in shape. One band was preserved with a primary osteoderm that was angled across the neck from side-to-side and was as long as the whole band itself. The base of the scute is rounded and ends with a tapered point, and the upper side of the scute has a smooth, lightly arching shape. Other bands are preserved with a triangular osteoderms with flat tops and rounded bottoms. The exact placement of cervical bands is not known. In 1995, Pereda-Suberbiola et al. suggested that in a more traditional placement, the bands would have been horizontal along the body, with the neural spine in the middle of the back. That positioning would mean that the medial scute would be next to another osteoderm of equal size, and together they would either fuse, like in Edmontonia, or touch, as in Sauropelta. Another possibility, suggested by Ford, was that the bands were along the side of the neck, pointing dorsally. If oriented along the side, the primary scutes would have pointed up and down, like in polacanthids, and the medial scutes would, by definition, become secondary osteoderms. The set of medial scutes (or secondary) would be possibly oval in shape.[11]
Thoratic scutes on struthiosaurines are oval to teardrop shaped, and possess sharp ridges that rise distally. Some scutes were long and had small domes on them. The first primary osteoderms on the pelvis are large, compressed from the sides, and have a sharp, short point. S. sp. was preserved with five fragmentary scutes from the pelvis. One fragment includes two small scutes with a ridge down the middle, joining the two together, and a compressed osteoderm with a small spike. It is thought that the fragment was from the edge of the pelvic region. Another fragment includes two oval osteoderms with small ossicles fused between them. Pelvic shields were probably formed on struthiosaurines by these scutes.[11]
Caudal scutes have been preserved on struthiosaurines. The osteoderms are compressed inwards from the side, have a slope positioned anteriorly, and a square-shaped posterior edge.[11]
Distinguishing anatomical features
All ankylosaurs that possess these characteristics - a narrow predentary; a nearly horizontal quadrate that is not fused and is oriented 30 degrees from the skull roof; the presence of mandibular condyles that are three times wider than long; premaxillary and dentary teeth that are near a symphysis with the front of the lower jaw (the predentary); a sacrum arched on top; an acromion process above the midpoint of the scapula to coracoid attachment; a straight ischium with a straight dorsal margin; relatively long, slender limbs; sacral shield armour; and the presence of erect pelvic osteoderms with flat bases - form a clade of basal nodosaurids, the Struthiosaurinae. That set of cranial and postcranial features are only present on genera considered to be in the clade. The features above distinguish Struthiosaurinae from other clades and genera found by other analysis'.[1]
Biogeography
The near simultaneous appearance of nodosaurids in both North America and Europe is worthy of consideration, because at the time, they were separated from each other by a huge body of water. Europelta is the oldest nodosaurid from Europe; it is derived from the lower Albian Escucha Formation. The oldest western North American nodosaurid is Sauropelta, from the lower Albian Little Sheep Mudstone Member of the Cloverly Formation, at an age of 108.5±0.2 million years. Eastern North American fossils seem older. Teeth of Priconodon crassus have been derived from the Arundel Clay of the Potomac Group of Maryland, which dates near the AptianâAlbian boundary. A Propanoplosaurus hatchling was uncovered from the base of the underlying Patuxent Formation, dated to the upper Aptian, making Propanoplosaurus the oldest nodosaurid.[1]
Polacanthids are known from pre-Aptian fauna from both Europe and North America. The timing of the appearance of nodosaurids on both continents indicates the origins of the clade preceded the isolation of North America and Europe, thereby pushing the group's date of evolution back to at least the middle Aptian. The separation of Nodosauridae into European Struthiosaurinae and North American Nodosaurinae by the end of the Aptian provides a revised date for the isolation of the continents from each other with rising sealevel.[1]
Struthiosaurinae is one of the longest-lasting groups of ankylosaurians. They range from Europelta at 112 Ma to Struthiosaurus, which lived until the uppermost Cretaceous, or 66 Ma.[1] In between those two early and late struthiosaurines are the genera Anoplosaurus and Hungarosaurus. Hungarosaurus is younger, at about 85 Ma, from the late Santonian of the Csehbánya Formation. Anoplosaurus is a fair amount older, at about 100 Ma, from the late Albian Cambridge Greensand.[13]
See also
Timeline of ankylosaur research
References
1 2 3 4 5 6 7 8 9 10 Kirkland, J. I.; Alcalá, L.; Loewen, M. A.; EspÃlez, E.; Mampel, L.; Wiersma, J. P. (2013). Butler, Richard J, ed. "The Basal Nodosaurid Ankylosaur Europelta carbonensis n. gen., n. sp. From the Lower Cretaceous (Lower Albian) Escucha Formation of Northeastern Spain". PLoS ONE. 8 (12): e80405. doi:10.1371/journal.pone.0080405.
â Nopcsa, B. F. (1902). "Notizen über cretacische Dinosaurier [Notes on Cretaceous dinosaurs]". Sitzungsberichte der Mathematisch-Naturwissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschaften. 111 (1): 93â114.
â Nopcsa, B.F (1915). "Die dinosaurier der Siebenbürgischen landesteile Ungarns". Mitteilungen aus dem Jahrbuche der Königlich-Ungarischen Geologischen Reichsanstalt. 23: 1â26.
â Nopcsa, B. F. (1923). "Die Familien der Reptilien. Fortschritte der Geologie und Paleontologie". Science. 58 (1512): 517â526. doi:10.1126/science.58.1512.517-a.
â Nopcsa, F. (1923). "On the Geological Importance of the Primitive Reptilian Fauna in the Uppermost Cretaceous of Hungary; with a Description of a New Tortoise (Kallokibotion)". Quarterly Journal of the Geological Society. 79: 100â116. doi:10.1144/GSL.JGS.1923.079.01-04.08.
â Nopcsa, B.F. (1928). "The genera of reptiles". Palaeobiologica. 1: 163â188.
â Nopcsa, B.F. (1928). "Palaeontological notes on reptiles. V. On the skull of the Upper Cretaceous dinosaur Euoplocephalus". Geologica Hungarica, Series Palaeontologica. 1 (1): 1â84.
â Nopcsa, B.F. (1929). "Dinosaurierreste aus Seibenburgen V.". Geologica Hungarica. Series Palaeontologica (4): 1â76.
â Coombs, W.P. Jr. (1978). "The Families of the Ornithischia Dinosaur Order Ankylosauria" (PDF). Paleontology. 21 (1): 143â170.
â Tumanova, T.A. (1987). "The Armored Dinosaurs of Mongolia" (PDF). The Joint Soviet-Mongolian Paleontological Expedition Transaction. 32: 9.
1 2 3 4 5 Ford, T.L. (2000). Lucas, S.G.; Heckert, A.B., eds. "Dinosaurs of New Mexico". New Mexico Museum of Natural History and Science Bulletin. 17: 157â167.
â Stein, M.; Hayashi, S.; Sander, P. M. (2013). "Long Bone Histology and Growth Patterns in Ankylosaurs: Implications for Life History and Evolution". PLoS ONE. 8 (7): e68590. doi:10.1371/journal.pone.0068590. PMC3722194. PMID23894321.
â Weishampel, D.B.; Dodson, P.; and Osmólska, H. (2004). The Dinosauria, 2nd. Berkeley: University of California Press. pp. 588â593. ISBN 0-520-24209-2.
"Struthiosaurinae". Fossilworks - Gateway to the Paleobiology Database.
"Struthiosaurus". Fossilworks - Gateway to the Paleobiology Database.
"Struthiosaurinae". Paleobiology Database: Classic.
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UPDATE FOR 1 January 2015: Apologies to all fans and readers of Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages for the three year delay (YIKES) in updates. I hope the new additions are worth the wait.
1 January 2015 Genus List available soon: includes all Mesozoic dinosaurs named up through the end of 2014! Next revision expected in late December 2015
Greetings,
This website is provided especially for readers of my recent book Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages (illustrations by Luis Rey; Random House, published 2007). In the Introduction I begin by saying "The world of dinosaurs is changing", and indeed it has been! In fact, to quote from p. 5:
"I've tried to make this book as up-to-date as I can while I've been writing it. But new dinosaur discoveries are being made all the time. Some of these will just add a species or two to the list of known dinosaurs, but some may be as amazing as finding the first dinosaur fossil with feathers, or the first dinosaur nest, or the first-ever fossilized dinosaur bone!."
The purpose of this website is provide you with an update of information on dinosaurs. I'll try to hit the highlights of recent discoveries in the field, chapter by chapter. And I will update these periodically (with dates of new additions listed before the appropriate section). (The initial list of updates date to October 2007, when the book was released).
Additionally, I will provide a link at the bottom to an updated version of the Appendix: Dinosaur Genus List, containing newly named dinosaurs and revised classifications. Furthermore, I will provide the complete introduction to that Appendix, explaining what I meant by the various size and weight classes.
By the way, the book was featured in an advertisement for the American Museum of Natural History, New York City! (However, the author and publisher make no claims that getting and reading a copy of the book will guarantee that you win the American Awesome Association's Awesome Prize!):
Also, illustrations by Luis Rey--including several from this book--are featured in the closing credits for the theatrical movie Walking with Dinosaurs 3D (2013).
SUPPLEMENTARY INFORMATION TABLE OF CONTENTS:
I. Chapter Updates (Current as of 3 January 2011) II. Updated Dinosaur Genus List (Current as of 29 December 2011) III. Reviews of the book IV. Frequently Asked Questions (coming soon!) V. Website Links
CHAPTER UPDATES
So, let's see what's new with the dinosaurs:
Chapter 3: Fossils and Fossilization
Extraordinarily well preserved "dinosaur mummies" have been in the news. These include "Dakota", an Edmontosaurus (or Anatosaurus), and "Leonardo", a Brachylophosaurus. These two hadrosaurines provide a great amount of information about the "soft tissues" (muscles, skin, etc.) that don't normally preserve. But no, these aren't like Egyptian mummies or the freeze-dried mammoths of the polar regions: the muscles and skin are mineralized, not just dried.
NEW December 2010: You know how in the book I write that dinosaur colors do not preserve? Well, color me astonished! It turns out that there are ways that--in rare circumstances--we can figure out some of the colors of dinosaurs. There are many different factors that go into making scales or feathers different colors, one of which are melanosomes: little microscopic structures that hold different color pigments. In turns out that different color melanosomes actually have different sizes and shapes. For example, some are associated with grey or black, others with reddish-brown, and the lack of melanosomes would mean that the feather was white.
Using this information, different teams of paleontologists have shown that the compsognathid Sinosauropteryx had reddish-brown feathers, and a tail with bands of reddish-brown and white, while the primitive troodontid Anchiornis had a reddish-crest and a body that was mostly dark but with white bands. (In fact, Anchiornis' colors and patterns reminds me a lot of Hylatomus (formerly "Dryocopus") pileatus, the modern pileated woodpecker.)
However, it is important to note a few things. For one, it takes a very special fossil to preserve the melanosome shapes and sizes. Only a few locations with exceptional preservation allow for the survival of the kind of details on the feathers or protofeathers which are visible under a scanning electron microscope. In most dinosaur fossils there simply isn't that kind of microscopic detail present. Also, there is more to color than just melanosomes: they are literally just part of the picture. So these new images of Sinosauropteryx and Anchiornis--amazing as they are--are almost certainly still not 100% correct.
NEW January 2015: There is a debate going on in paleontology as to whether the "melanosomes" are really melanosomes (and thus indication of color) or if they are just preserved bacteria. I am not a microbiologist or a histologist, so I am really not qualified to judge the particulars. Additional lines of evidence for dinosaur (and other fossil animal) colors come from preserved trace elements in the body covering, and structural differences in the same that vary the color.
Chapter 4: Geologic Time
NEW January 2008: This doesn't affect the main portion of the book, but the International Commission on Stratigraphy (the organization in charge of the names of geologic time units) has announced that they will be restoring the old "Quaternary Period" (although somewhat longer than the old version: 2.588 million years ago to the present). So the Cenozoic Era will have three periods: Paleogene, Neogene, and Quaternary.
NEW December 2011: New geological dating of the various boundaries of the Triassic Period means that the dates in the book are slightly off. The Permian-Triassic boundary is about 252.3 million years ago; the Early Triassic/Middle Triassic boundary is 247.2 million years ago; the Middle Triassic/Late Triassic boundary is 235.0 million years ago; and the Triassic/Jurassic boundary is 201.5 million years ago.
NEW December 2013: New estimations and calibrations of the precise half-life of the main type of radiometric dating system means that many dates referred to in the text and the appendix were slightly off the present understanding. For example, while I said the the end of the Cretaceous Period was 65.5 million years ago in the text, the new calculation is 66.0 million years ago. I will try to make the changes in the text. The time chart below shows the current (2013) calculations of the age boundaries. ("Ma" is the geological shorthand for "millions of years ago):
Also, refinement of the precise geological ages within some of the better-studied Late Cretaceous geological formations of the North American West has allowed paleontologists to more accurately narrow down the age ranges of many species of dinosaurs from those rock units. I have tried to show those new refined ages in the appendix. These age ranges are MUCH shorter than seen in the earlier versions of the appendix; in fact, in many cases they are just a million year or so. This doesn't mean that these dinosaur genera only lasted for a million years but the ones elsewhere lasted much longer. Instead, it is very likely that most dinosaur genera lasted only a million years or so: the longer age ranges for these other ones are simply an indication that we really haven't narrowed down the times of these rocks as accurately as we have those in the Late Cretaceous of the North American West.
Chapter 11: The Origin of Dinosaurs
A considerable amount of new discoveries have been made concerning the closest relatives of Dinosauria. One of the more primitive relatives of dinosaurs is little Dromomeron from the Late Triassic of New Mexico, close kin to the earlier Lagerpeton of Middle Triassic Argentina. Together these primitive dinosaur relatives form the group Lagerpetonidae.
More significantly, though, are several finds related to the reptile Silesaurus shown in Chapter 11. One of these is Sacisaurus of the Late Triassic of Brazil, about 5 feet (1.5 meters) long. The first description of this near-dinosaur was published in October 2006. These two represent a distinct group of herbivores very closely related to the dinosaurs. Like true Dinosauria their pubis and ischium bones are very long; unlike true dinosaurs, their hands do not seem to have been large and grasping, nor do their hips have an open socket. Sacisaurus seems to have a pair of small bones in front of the dentary, similar to the larger single predentary bone of ornithischian dinosaurs.
Based on comparisons with the more complete skeletons of Silesaurus and Sacisaurus, it turns out that a number of previously-known creatures from the Triassic belong to this group of "silesaur" near-dinosaurs. These include Lewisuchus and Pseudolagosuchus of the Middle Triassic of Argentina (possibly the same species!) and Technosaurus and Eucoelophysis of the American Southwest. Additionally, some species known only isolated teeth of the Late Triassic Southwest (Crosbysaurus, Galtonia, Krzyzanowskisaurus, Lucianosaurus, Pekinosaurus, and Protecovasaurus) might be from silesaurs, or they might be from a newly-recognized group of herbivorous crocodile relatives the revueltosaurs. In fact, the near-dinosaur silesaurs and the near-crocodilian revueltosaurs represent two of the most important groups of herbivorous reptiles of the Middle and Late Triassic, and they weren't even known five years ago!
NEW January 2010: Silesauridae--the group described in the previous two paragraphs--has been formally named. Many analyses now show this group to be the closest known relatives to the first dinosaurs.
NEW December 2010: Yet another silesaurid (little Asilisaurus of the Middle Jurassic of Tanzania) has been discovered, increasing our knowledge of these close kin to the first dinosaurs. However, Dinosauria has "lost" another member: that is to say, a creature once considered an early dinosaur turns out to be something else. The reptile in question is Azendohsaurus
A sequence of trackway sites in Poland have given us a lot of new information on the rise of dinosaurs and their ancestors. The oldest dinosauromorph tracks are only a few million years after the great Permo-Triassic Extinction, and so it turns out that dinosauromorphs had split from their closest relatives (pterosaurs, and somewhat more distantly the crocodile-line archosaurs) extremely early in the Triassic. Furthermore, there are tracks in the Middle Triassic (older than any dinosaur bone fossil) that are almost certainly from herrerasaur theropods. So it looks like there is strong evidence now for Triassic dinosaurs.
NEW December 2013:: Silesaurids continue to show up in rocks around the world from the Middle and Late Triassic. Even closer to dinosaurs, however, is Nyasasaurus, from the same rocks that yielded the fossils of Asilisaurus. Nyasasaurus is known from very few bones, but distinctive features of the upper arm and elsewhere on the body show that it is either the first Middle Triassic dinosaur ever discovered or an animal which was closer to the common ancestor of all dinosaurs than are silesaurids. Let us hope that more specimens of this fascinating little reptile are discovered soon!
Chapter 12: Saurischians (Lizard-Hipped Dinosaurs)
Debate continues on whether Eoraptor and Herrerasauria are actually theropods or simply primitive saurischians. (Also, note that the new proper term for the larger group containing Herrerasaurus and the other herrerasaurids is "Herrerasauria").
NEW January 2010: In December 2009 this debate may have come to an end. Specifically, discovery of a new primitive theropod dinosaur named Tawa from the Late Triassic of New Mexico has helped to answer some of the old questions. Tawa shows features shared by more advanced theropods and those of primitive saurischians like Herrerasauria and Eoraptor. With all this new information, it now appears that Herrerasauria is a group of very primitive theropod; that Eoraptor is a slightly more advanced form; that Tawa is more advanced still, and is in fact the closest known relative to Coelophysis plus later theropods. (This last group--Coelophysis plus later theropods--is technically known as Neotheropoda.)
NEW January 2011: Yet more changes! Brand new discoveries from Argentina by Ricardo Martinez of Instituto y Museo de Ciencias Naturales, Universidad Nacional de San Juan and Paul Sereno of the University of Chicago and their teams reveals Eodromaeus, a very primitive theropod, more closely related to Tawa and the neotheropods than are herrerasaurs. However, as part of the study of this dinosaur it was discovered the Eoraptor--long considered either a primitive saurischian or a primitive theropod--is more likely a guaibasaurid sauropodomorph, and thus closer to Diplodocus and Brachiosaurus than to Allosaurus and Velociraptor. Still, Eoraptor and Eodromaeus are still very similar to each other in terms of their anatomy, and they show that the earliest saurischians (and the earliest dinosaurs in general) were all about 1-2 m (3.3-6.6 ft) long, bipedal, with grasping hands and teeth suited equally to plants and meat. In other words, early dinosaurs were sort of "reptilian raccoons".
NEW December 2011: Another primitive theropod--closer to neotheropods than are herrerasaurs, but not as close as Tawa is little, buck-toothed Daemonosaurus. Unlike most early theropods, it had a relatively short, blunt snout.
NEW December 2013: As additional specimens have been described, the case that Eoraptor is actually a sauropodomorph rather than a theropod continues to improve. On the other hand, whether herrerasaurs are primitive theropods or simply primitive saurischians is still not secure one way or the other.
NEW January 2014: Guaibasaurus continues to sometimes fall out as a primitive theropod, and sometimes as a primitive sauropodomorph. Eoraptor, however, does seem to be very securely a primitive sauropodomorph. For now.
Chapter 13: Coelophysoids and Ceratosaurs (Primitive Meat-Eating Dinosaurs)
Late Triassic Guaibasaurus of Brazil may actually be a true theropod: if so, it is the most primitive of all. For example, it lacks the three-toed foot typical of all more advanced theropods. (NEW December 2010: However, see Chapter 22 update below for an even newer interpretation of Guaibasaurus.)
There is still confusion about the relationships between the primitive theropods. Some studies support the breakdown that I use in the book: the coelophysoids as the first major branch, and a second group of ceratosaurs more closely related to the tetanurines than to the coelophysoids. However, there are some researchers who have evidence that the old 1980s-1990s idea that coelophysoids and ceratosaurs are each other's closest relatives in a a grander Ceratosauria. To make it even more confusing, still other evidence points to a different arrangement. In this third scheme, the slender small coelophysoids such as Coelophysis, Megapnosaurus, Procompsognathus, Segisaurus, and the like represent the first branch separate from group of larger forms. These larger dinosaurs, the Early Jurassic Dilophosauridae, containing newly-discovered Dracovenator of South Africa, Dilophosaurus of western North America, and "Dilophosaurus" sinensis of China. In fact, work first published in September and October 2007 by Nathan Smith (graduate student at the Field Museum of Natural History in Chicago and the University of Chicago) and his colleagues shows that Cryolophosaurus of Early Jurassic of Antarctica is also a member of this group. I think that Smith and his colleagues probably have the closest approximation to the truth so far when it comes to primitive theropods.
In this new hypothesis, these 16.5 to 21 foot (5-7 m) long dilophosaurids were more closely related to the ceratosaurs and tetanurines than they were to the "true" coelophysoids. I find this a really interesting idea, and look forward to future research that might resolve this issue. Whether the Late Triassic Zupaysaurus and Gojirasaurus turn out to be early representatives of the dilophosaurids, or big true coelophysoids, or something in between the two remains to be seen.
The oldest true ceratosaur in the restricted sense, Berberosaurus of northern Africa, was first described by Ronan Allain (Muséum national d'Histoire Naturelle, Paris) and colleagues in September 2007. This dinosaur shows that true ceratosaurs (that is, the group including Ceratosaurus, Elaphrosaurus, noasaurids, abelisaurids, but not coelophysoids of any sort) were already around by the Middle Jurassic.
NEW July 2008: A major restudy of the ceratosaurs has been published by Matt Carrano (Smithsonian Institution) and Scott Sampson (University of Utah). Among other discoveries, they found that Deltadromeus, previously thought to be a giant noasaurid, is more likely a primitive ceratosaur.
NEW January 2010: There has been a lot of work on the dinosaurs covered in this chapter. The newly-discovered Tawa of the Late Triassic of New Mexico shows a mixture of primitive theropod and neotheropod features; when it is added into our studies, it shows that even the reduced version of "Coelophysoidea" mentioned in the paragraphs above is not a natural group either. Instead, "coelophysoids" simply represent "the primitive phase of the Neotheropoda."
And while Herrerasauria and Eoraptor seem to be more clearly moving INTO Theropoda, Guaibasaurus is moving out: see notes for Chapter 22.
Among the primitive ceratosaurs, the most important new discovery is Limusaurus. This dinosaur from the earliest Late Jurassic of Xinjiang, China (the same site as the early tyrannosauroid Guanlong) shares many features in common with Elaphrosaurus, but is known from more complete fossils. Most significantly, it has a toothless beak very similar to the later ornithomimid coelurosaurs and extremely short arms with stubby hands with only two fingers. These two features suggest that it was not a carnivore at all, but instead a herbivore (or at best an omnivore that ate mostly plants and small animals.)
Skorpiovenator from the Late Cretaceous of Argentina was first named in 2008, and represents one of the mostly completely known skeletons of an abelisaurid so far discovered.
NEW December 2011: Continued work on Late Triassic/Early Jurassic theropods has failed to consistently find a "dilophosaurid" group. It might exist, or some of these dinosaurs (such as Cryolophosaurus) may be closer to averostrans (ceratosaurs plus tetanurines) than to Dilophosaurus.
NEW December 2013: Wow, where to begin? Okay: Guaibasaurus may be a primitive theropod, or it may be a primitive sauropodomorph. The "dilophosaurids" have failed to come together as a group. In recent studies (one big one by Matt Carrano, Roger Benson & Scott Sampson, and on-going research by Xing Lida), it was found that Dilophosaurus proper and Zupaysaurus are true coelophysoids, but that Cryolophosaurus and Sinosaurus triassicus (formerly called "Dilophosaurus" sinensis) are primitive tetanurines.
A large new analysis of ceratosaur relationships by Thierry Tortosa and colleagues in France reveals the following groups (and is the basis for the current version of the appendix): an "elaphrosaur" clade comprised of Spinostropheus, Elaphrosaurus, and Limusaurus; various primitive medium-to-large carnivorous ceratosaurs (such as Ceratosauridae, Berberosaurus, and the newly-discovered Eoabelisaurus [which was first thought to be an incredibly ancient abelisaurid]); a diverse Noasauridae (including giant Deltadromeus and recently-named Daholokely of Madagascar; and Abelisauridae. Within the abelisaurids this study found a variety of early branches, a group Majungasaurinae (which includes Manjugasaurus of Madagascar, the Indian abelisaurids, and some European forms) and Brachyrostra (the South American group.)
NEW January 2015: A new discovery from Venezuela: Tachiraptor from the very beginning of the Jurassic. Although known only from fragmentary material, it seems to be very close to the common ancestor of Ceratosauria and Tetanurae.
Chapter 14: Spinosauroids (Megalosaurs and the Fin-Backed Fish-Eating Dinosaurs)
As mentioned in the comments for Chapter 13, new research shows that the Early Jurassic dinosaurs "Dilophosaurus" sinensis (which will eventually get its own genus name!) and Cryolophosaurus are not primitive tetanurines, but rather part of the Dilophosauridae.
On p. 92, the name "Calvadosaurus" is a mistake: the dinosaur in question is properly called Dubreuillosaurus.
NEW January 2008: The prosauropod behind Cryolophosaurus on p. 90 now has a name: Glacialsaurus.
NEW January 2010: WOW!! There have been some major discoveries and re-analyses within this part of the theropod family tree in the last couple years, many of them due to graduate student Roger Benson of Cambridge University in England. Most of this work was a result of Benson's detailed study of Megalosaurus itself. He and his co-authors have argued that the dinosaurs in this chapter should more accurately be called "Megalosauroidea" rather than "Spinosauroidea", and if I do a new edition of this book I will be using that name.
Here are some of Benson and colleagues discoveries:
Monolophosaurus, Marshosaurus, and Piatnitzkysaurus (among others) are primitive megalosauroids.
Megalosauridae itself contains Torvosaurus, Dubreuillosaurus, Afrovenator, Eustreptospondylus, and some more poorly known forms.
Spinosauridae and Megalosauridae are sister groups
NEW December 2011: A new giant spinosaurid, Oxalaia, has been discovered from the mid-Cretaceous of Brazil. Though known only from incomplete remains, it seems to rival the big African spinosaurids in size.
NEW December 2013: REAL big changes here, mostly from the work of Carrano, Benson and Sampson. Their new study forms the major structure of the primitive tetanurine part of the new appendix. As mentioned above, they found that Cryolophosaurus DOES belong in this chapter, as the most primitive-known tetanurine. Sinosaurus (formerly "Dilophosaurus" sinensis), Chuandongcoelurus, and Monolophosaurus are similarly primitive tetnaurines. A careful reader will note that the heads of all of these dinosaurs (where known) have crests: apparently, that was the "in" thing for Early and Middle Jurassic primitive tetanurines.
Carrano, Benson, and Sampson gave a name to the group of Megalosauroidea plus Avetheropoda: Orionides, the hunters. Their work found that there was a group of primitive megalosauroids, which they named "Piatnitzkysauridae" (including Marshosaurus and Piatnitzkysaurus) and the more specialized "Megalosauria". The megalosaurians include the lightly built Streptospondylus and Eustreptospondylus, the long-snouted giant Spinosauridae, and the Megalosauridae. Within Megalosauridae they found two groups: the heavily-built Megalosaurinae and the slightly less bulky Afrovenatorinae.
In spinosaurid news, at present spinosaurid fossils (sometimes just teeth) have been found in the Early and mid-Cretaceous of South America, Europe, Africa, Asia, and Australia, but still no sign of them in North America. Where are they?!? Chemical study of their teeth shows they they ate more fish than typical theropods in their environment (but that doesn't mean that they actually ate ONLY fish! After all, there are partially-digested bones of a baby Iguanodon in the belly of Baryonyx).
NEW January 2014: Whoops! Should have mentioned this in the previous update. The most complete ever megalosauroid skeleton (or at least, probable megalosauroid) was described. There are some unusual aspects to it. For one thing, it is only a baby! For another, it is covered in fuzz!! It was given the name Sciurumimus, "squirrel mimic". But keep in mind this is just a hatchling! We have no idea how big this animal grew! Furthermore, similarities between it and the "compsognathid" Juravenator suggest that the latter may also be a megalosauroid, and not a coelurosaur as previously thought.
NEW January 2015: The BIG news in megalosauroid research for the year is the discovery of a new, more complete specimen of Spinosaurus! Combining this information and other skeletons found over the years, a team of scientists have presented a new skeletal reconstruction of Spinosaurus and new interpretation of its lifestyle. Here are some of their discoveries and conclusions:
The fragmentary dinosaurs previously called "Spinosaurus B" and "Sigilmassasaurus" are really just Spinosaurus itself.
The hips and legs are rather small for a dinosaur of its size.
The arms, in contrast are quite large. (However, it must be noted that the humerus used to figure out the size of the arms is probably from the sauropod Rebbachisaurus and NOT from Spinosaurus at all!)
The limb bones are solid in Spinosaurus, unlike all other theropods except some diving birds.
The authors therefore conclude that Spinosaurus was an aquatic animal, spending almost all of its time swimming (more like a crocodile than a typical dinosaur)
Their ideas have met with some resistance from other paleontologists, it must be added. I am looking forward to the detailed description of the new material, currently being written by the researchers.
Chapter 15: Carnosaurs (Giant Meat-Eating Dinosaurs)
NEW January 2010: As with the previous chapter, a whole heckuva lot of new discoveries in this part of the dinosaur family tree. In this case one of the main contributors has been Stephen Brusatte (currently a graduate student at Columbia University and the American Museum of Natural History in New York City).
As mentioned in the update to Chapter 14, Monolophosaurus has been moved out of Carnosauria into Megalosauroidea. Also, several European dinosaurs that were previously considered megalosaurids or other primitive tetanurines (Poekilopleuron, Lourinhanosaurus, Metriacanthosaurus) are now recognized to be part of Sinraptoridae; because of this, sinraptorids are not strictly Asian dinosaurs anymore.
More significantly, however, has been the recognition of a whole new branch of carnosaur: the Neovenatoridae and its subgroup the Megaraptora. Neovenatoridae is a newly-recognized sister group to Carcharodontosauridae (together they form Carcharodontosauria.) Neovenatorids have rather large thumb claws, among other features. Neovenator and gigantic Chilantaisaurus are primitive neovenatorids, while the rest form a group of slender forms called Megaraptora. Megaraptora includes: Fukuiraptor of the Early Cretaceous of Japan; Megaraptor itself (and thus it belongs in Chapter 15, not Chapter 14!); Aerosteon of the Late Cretaceous of Argentina (this is the "unnamed Argentine carcharodontosaurid" mentioned on p. 104); Australovenator of the Early Cretaceous of Australia; and Orkoraptor of the latest Cretaceous of Argentina. Since Orkoraptor belongs in this group, the carnosaurs actually did make it to the end of the Cretaceous.
In other news, new studies show that Shaochilong (previously called "Chilantaisaurus" maortuensis) is a carcharodontosaurid, and thus this group is now known to have lived in Asia.
NEW December 2010: A new carnosaur (a primitive carcharodontosaurid) from the Early Cretaceous of Europe has been described: Concavenator. It is known from a nearly complete skeleton with scale impressions. Among the more remarkable things about this dinosaur is a short-but-tall sail right in front of the hips. Possibly more remarkable is some evidence that-- perhaps--it had feathers on its arms! There are a series of knobs along its forearm that are interpreted by its discovered as the connections to large arm quills or feathers (such structures are known on some modern birds and on Velociraptor, for instance). However, an alternative explanation is that these may instead be a line of connection for tissue within the arm muscles.
NEW December 2011: The oldest carcharodontosaurid yet described, Veterupristisaurus is known from a few tail vertebrae from the Late Jurassic Tendaguru Formation of Tanzania.
NEW December 2013: Continuing Carrano, Benson & Sampson's overhaul of primitive tetanurine relationships, some of the big changes. The lightly-built Lourinhanosaurus of Late Jurassic Portugal remains uncertain where it belongs: they find (like I did, back in 2004) that it is a primitive coelurosaur. Within Carnosauria they find the most primitive branch to be Metriacanthosauridae (what I called "Sinraptoridae" in the book) and the more specialized Allosauria. Within Allosauria are Allosauridae (just Allosaurus and Saurophaganax for now) and Carcharodontosauria. As mentioned above, within the carcharodontosaurs are the gigantic Carcharodontosauridae and the more slender Neovenatoridae, the latter containing some primitive forms and the very slender Megaraptora.
In megaraptoran news, the first megaraptoran from North America--Siats of Utah--has been discovered. Orkoraptor of Argentina has been reedited: I mention above that it lived near the end of the Cretaceous, but redating of the rocks from which its fossils came show that it was from around 90 million years ago. So the book was correct in saying that Carnosauria was gone before the end of the Cretaceous.
Are megaraptorans really carnosaurs? In the book I point out that they are hard to place: maybe they are megalosauroids ("spinosauroids" at the time), maybe they are carnosaurs, maybe they are primitive coelurosaurs? A new possibility was suggested by Fernando Novas and his colleagues: that they are actually tyrannosauroids! I do see some similarities there, but am not yet convinced that the evidence fits this hypothesis better than the megaraptorans-as-neovenatorid-carcharodontosaurs hypothesis. We'll see what new discoveries are made.
NEW January 2015: The discovery of a juvenile Megaraptor specimen does show they had long slender snouts, similar to some primitive tyrannosauroids.
Chapter 17: Tyrannosauroids (Tyrant Dinosaurs)
NEW January 2008: Last year there were many news reports about the finding that Tyrannosaurus rex "had three fingers". Unfortunately, those reports were not correct. In fact, the real find was a well-preserved third metacarpal (long bone of the palm of the hand) for T. rex: nota surprise as these were already known in other two-fingered tyrannosaurids. In fact, you can see this little bone on the hand skeleton of Tyrannosaurus on the bottom of page 120.
NEW January 2010: 2009 was a spectacular year for tyrant dinosaur discoveries. New examination of the skull of Middle Jurassic Proceratosaurus of England shows that it is a very primitive tyrannosauroid, and possibly a close relative of Guanlong. From the later Early Cretaceous of China comes long-snouted Xiongguanlong and tiny Raptorex which help us fill in the spaces of the tyrant family tree. Among other things, Raptorex shows that the dinky arms and pinched foot (arctometatarsus) of Tyrannosauridae evolved in smaller bodied ancestors.
Among true Tyrannosauridae comes the most-complete skeleton of Alioramus yet discovered. Isn't it pretty? (Maybe just to me...)
NEW December 2010: A new phylogenetic analysis by Stephen Brusatte and Thomas Carr reveals that Proceratosaurus, Guanlong, and a few other newly discovered genera form a clade called "Proceratosauridae", and represent the oldest and most primitive branches of Tyrannosauroidea.
NEW December 2011: New tyrannosauroids continue to show up. Among these are Teratophoneus of Utah and Zhuchengtyrannus of China.
NEW December 2013: Lots of new tyrannosauroid discoveries. Here are three really important ones: "Raptorrex" is probably not as old as originally thought, and may just be a juvenile of Tarbosaurus; Lythronax the "gore king", the oldest true tyrannosaurid and closer to Tyrannosaurus and Tarbosaurus than to other tyrannosaurids, was named; and most astonishing of all, Yutyrannus of the Early Cretaceous of China was discovered. This is the first giant tyrannosauroid known with fuzz, and it has fuzz all over the body! Previously it was argued that giant theropods may have lost their fuzz, but now we know that even 1.4 ton giants could have it.
Oh, and the great old question "which is the biggest theropod?": dinosaur paleontologist and artist Scott Hartman found that the biggest Tyrannosaurus was BARELY larger than the biggest known carnosaur, but that it is still uncertain if Spinosaurus is more massive then either.
NEW January 2015: Among the new tyrannosaur discoveries last year was Qianzhousaurus, nicknamed "Pinnochio rex" because of its long snout. The discoverers consider it to form a group with Alioramus, and they named the group the "Alioramini". I am not altogether convinced that this isn't simply an older individual of Alioramus itself, but if/when new fossils are found, we can sort this out.
And if you have an hour to kill, hear about the Life and Times of Tyrannosaurus rex by yours truly.
Chapter 18: Ornithomimosaurs and Alvarezsaurs (Ostrich and Thumb-Clawed Dinosaurs)
NEW January 2010: New studies by Lindsay Zanno of the Field Museum in Chicago and her colleagues, based on a new species of the therizinosaur Nothronychus, suggest a new relationship among the dinosaurs of this chapter and the next, and a new understanding of their diet and history. In particular, Zanno and colleague's work shows that ornithomimosaurs, therizinosaurs, alvarezsaurs, oviraptorosaurs, deinonychosaurs, and avialians branched off in that order. So if I ever do a new edition of the book, I would probably include therizinosaurs in this chapter (and possibly move alvarezsaurs into the next).
More significantly, though, these paleontologists observe that with the exception of some deinonychosaurs, all of the dinosaurs in this part of the family tree show signs of eating at least some non-meat (mostly plants, but also insects and/or small bodied vertebrates). The simplest explanation would be that their common ancestor (which would have split off from the common ancestor of the meat-eating compsognathids and tyrannosauroids some time before the Middle Jurassic) began to eat things other than dinosaurs. (It also means that meat-eating deinonychosaurs like Deinonychus and Velociraptor evolved from omnivorous ancestors!)
A flock of the ostrich dinosaur Sinornithomimus was found in China by Dave Varricchio and colleagues. These poor dinosaurs got stuck in some very sticky mud and died, which was bad for them but great for paleontologists! None of them were fully grown, but none were babies either. This suggests that perhaps during the brooding season, when parents were nesting with the new generation of babies the "kids" and "teenagers" hung out together without a parent around. For a modern example of this, visit almost any shopping mall...
NEW December 2010: Jonah Choiniere--then a graduate student at George Washington University in Washington, DC--and colleagues have described one of the most important theropod fossils found in recent years: Haplocheirus. Haplocheirus isn't very big, it isn't very scary, it doesn't have massive teeth or huge crests or anything like that. What makes this dinosaur so important is that it is a very, very primitive alvarezsaur! All the other alvarezsaurs are from the Late Cretaceous, and are very much changed from their ancestral condition, making it hard to clarify their evolutionary relationships. Haplocheirus, however, is from the earliest Late Jurassic (in fact, from the same formation and sites as Guanlong, Limusaurus, and Yinlong) and is not much different from other primitive coelurosaurs. Its hand may not have the giant thumb of later alvarezsaurs, but its thumb is already more powerfully developed than the other fingers. Fossils like these, ones that are near the base of the branch of a particular part of the family tree, are really helpful to paleontologists in figuring out the evolutionary origins and early life styles of different groups.
NEW December 2011: It was a good year for alvarezsaurids! Three new genera were named: Albinykus of Mongolia, Linhenykus of China, and Bonapartenykus of Argentina. Also, Bonapartenykus was found with its own eggs, so we finally have definite alvarezsaurid eggs. But keep your eyes peeled: there are big changes ahead for our current understanding of alvarezsaurids (and parvicursorines).
NEW December 2013: In a bit of a surprise (because these rocks had not yet preserved feathers or fuzz), fossils from the Late Cretaceous of Canada showed that adult Ornithomimus had broad feathers on their arms, but juveniles were simply fuzzy. And the BIGGEST thing in ornithomimosaur studies: more of the skeleton of Deinocheirus has been discovered! The paper hasn't been published yet, but at the 2013 meeting of the Society of Vertebrate Paleontology the first relatively complete material of the giant ostrich dinosaur was revealed. Unfortunately no skull was presented with this material, but it must be out there! Expect more than arms of Deinocheirus in future versions of the book!
NEW January 2015: The BIGGEST news (literally) in ornithomimosaur studies is the description of the new Deinocheirus material. And it is a weird dinosaur! It isn't just a Gallimimus-type dinosaur grown huge, but a weird sort of tiny-brained duckbilled ostrich camel dinosaurs. It has a broad bill like a hadrosaur; a sail on its back, something like Spinosaurus; a forward-tilted pelvis like therizinosaurs; and relatively modestly proportioned legs without an arctometatarsus. The describers consider it, Beishanlong, and Garudimimus to form a group (Deinocheridae) which is the sister taxon to Ornithomimidae.
Chapter 19: Oviraptorosaurs and Therizinosauroids (Egg-Thief and Sloth Dinosaurs)
One of the biggest discoveries (literally) among these dinosaurs is the new giant oviraptorosaur Gigantoraptor of the Late Cretaceous of Asia. Xu Xing (Institute of Vertebrate Paleontology and Paleoanthropology, Beijing) and his colleagues first published on this giant in June 2007. As big as tyrannosaurids like Albertosaurus and Gorgosaurus, Gigantoraptor was one of the largest dinosaurs in its environment. And yet it had long slender legs: longer and more slender than those of tyrannosaurids, in fact, so it may have been the swiftest of big dinosaurs.
What did Gigantoraptor eat? Well, other oviraptorosaurs seem to have eaten both plants and small animals, and the same may be true of Gigantoraptor. However, to this big theropod, "small animals" may have included sheep-sized ceratopsians and jackal-sized dromaeosaurids!
With the discovery of Gigantoraptor, we now recognize that practically all the groups of coelurosaurian theropods produced giants bigger than 1 ton: the tyrannosaurids (of course); Deinocheirus for the ornithomimosaurs; Gigantoraptor for the oviraptorosaurs; and Therizinosaurus for the therizinosaurs.
Some taxonomic notes: Following a new scheme, the name "Therizinosauroidea" would be restricted to the more stump-footed sloth dinosaurs, so that the larger category (including slender-footed Falcarius) would be "Therizinosauria". Also, the larger group of Oviraptorosauria plus Therizinosauria (if they do form a group together) would be Oviraptoriformes.
NEW January 2010: As I mentioned in the notes to Chapter 18, new studies suggest that Therizinosauria and Oviraptorosauria are not each others closest relatives. Instead, Oviraptorosauria seems to be more closely related to Deinonychosauria and Avialae.
NEW December 2011: To add to the discussion of WAIR (wing-assisted incline running), Ken Dial and his students and colleagues have identified another behavior of modern birds that may have been present in broad-feathered dinosaurs: Controlled Flapping Descent (CFD). CFD is a behavior used by young birds, who cannot yet fly, to flap when jumping out of trees to slow and control their landing. It turns out that few birds actually parachute as such; instead, they have an active stroke on the way down.
NEW December 2013: The primitive Early Cretaceous theriziosaur Jianchangosaurus shows that it--like ornithopods and ceratopsians--had teeth where the sides rather than the tips came together.
Many new oviraptorosaurs have been named. Too darn many, probably! I will not be surprised if many wind up collapsing down into a smaller number of genera, and that what we are calling separate genera are really just males, females, and/or juveniles.
NEW January 2015: Important new studies of the function of the claws and beaks of therizinosaurs (both studies led by Stephan Lautenschlager of the University of Bristol) were published.
The big latest Cretaceous North American crested oviraptorosaur (the one whose head is the upper left corner picture on p. 144) has been named: Anzu. On a personal note, I am involved with the excavation and description of a specimen of Anzu that has been nicknamed "Pearl".
The grouping of Oviraptorsauria plus Dromaeosauridae plus Troodontidae plus Avialae now has a name: Pennaraptora. A new detailed analysis of the evolution of the wrists of dinosaurs shows that the origin of the half-moon shaped wrist bone (semilunate carpal) is a pennaraptoran trait.
A new big coelurosaurian phylogenetic study by Steve Brusatte (now at the University of Edinburgh) and colleagues found (among other things), that Pedopenna and the Scansoriopterygidae were actually early members of the Oviraptorosauria. If so, this would mean we have Jurassic oviraptorosaurs (predicted by the presence of troodontids and avialians in the Jurassic) and that the oldest oviraptorosaurs were very small tree-dwelling forms (like the oldest troodontids, dromaeosaurids, and avialians). It will be interesting to see if this is confirmed in later studies.
Chapter 20: Deinonychosaurs (Raptor Dinosaurs)
2007 has seen the discovery of several new small Asian dromaeosaurs, including Shanag and Mahakala. The first of these is the oldest known (and first Asian example) of the unenlagiines; the second is a very small primitive dromaeosaurid no bigger than Archaeopteryx.
While no one has found the actual feathers of Velociraptor yet (the rocks in which it is found do not preserve feather impressions), a very recent discovery (September 2007) shows that it probably had very big arm feathers. Like little Rahonavis and Microraptor, Velociraptor had "quill nodes": bumps on its ulna (forearm bone) where big feathers attached. While Velociraptor would not be able to fly, it may have used its arm feathers as a help in turning quickly; while brooding its nests; for display; and (while a young one) for Wing-Assisted Incline Running.
NEW January 2008: Footprints of a large (Achillobator-sized) dromaeosaurid have been found from the Early Cretaceous of China, showing that they did indeed walk with their sickle-claws raised (as Luis Rey illustrated them in the book.)
NEW January 2010: Despite what I said on p. 154, it now appears that Pedopenna and Epidendrosaurus (and newly discovered Epidexipteryx
) are indeed very primitive avialians. But, to be fair, they are very close to the common ancestry of avialians and deinonychosaurs.
The "giant unenlagiine" discovered by Fernando Novas mentioned on pp. 155-156 has been named: it is Austroraptor (seen in the linked image with Novas himself!) At the other size extreme, little Hesperonychus of the Late Cretaceous of Canada is the smallest known theropod (other than birds) from North America, and the first North American microraptorine. Studies included these new dromaeosaurid have caused some shifting among the various subgroups of dromaeosaurids, which will be included in the 2010 genus list update.
On the troodontid side of things, little Anchiornis shows that troodontids a) were present in the Middle Jurassic; b) had long leg feathers like primitive dromaeosaurids and primitive avialians; and c) were cute.
Also, with regards to the discussion of brains on p. 161: new studies show that Troodon's brain was exceptionally big even for a troodontid, so it (rather than Troodontidae as group) may indeed have been the brainiest Mesozoic dinosaur.
NEW December 2010: The weirdest dromaeosaurid to be discovered in a long time is Balaur bondoc, or as I call it, the "double-barreled dromaeosaurid from Transylvania". Pretty much everything about it is bizarre. It has short arms ending in essentially two-fingered hands (trying to be a tyrannosaurid, I guess); its pubis isn't merely backwards-pointing, it is WAAAYYY backwards-pointing; and it has TWO sickle claws on each foot! (We don't know what the skull looks like yet: who knows what surprises are there?)
NEW December 2011: A big year for deinonychosaur studies! In terms of behavior, Denver Fowler (of Montana State University's Museum of the Rockies) and colleagues [including a former student of mine, Robert Kambic!] have presented a study which suggests that larger dromaeosaurids like Deinonychus and Velociraptor may have used their sickle claws more to pin prey down while their claws and jaws did the "dirty work", rather than as a "belly-ripper" itself.
In terms of classification, big news (and big disagreements!). On the one hand, the discovery of a new little theropod from the early Late Jurassic of China named Xiaotingia shows features of its anatomy that unite it, Anchiornis, and (most important of all) Archaeopteryx into a single group: Archaeopterygidae. What is more, the archaeopterygids were found to be primitive deinonychosaurs, and NOT avialians at all! If this is true, it explains why the pointy faces of archaeopterygids are similar to those of troodontids and dromaeosaurids, but different from the more blunt faces of primitive avialians, oviraptorosaurs, and the like: perhaps the pointy-face condition evolved just once, in the ancestors of all deinonychosaurs. If archaeopterygids are primitive deinonychosaurs, it is quite likely that all eumaniraptorans (deinonychosaurs plus avialians) could fly at first, and that non-flying deinonychosaurs like Troodon, Deinonychus, and Velociraptor all had flying ancestors.
However, other analyses find a very different set of relationships. In these, microraptorines and unenlagiines (or rather Microraptoria and Unenlagiidae, using the terminology of these studies), as well as Anchiornis, Archaeopteryx, and Rahonavis, are all primitive members of Avialae and not deinonychosaurs. Unfortunately these studies don't include the information from Xiaotingia, and the "archaeopterygids are primitive deinonychosaurs" study does not include the new unlagiine information. Hopefully a future study will include all this data.
The main point of all this isn't that dinosaur paleontologists don't know what they are doing! Instead, the important conclusion is that we now have many species which are pretty close in form to the divergence between deinonychosaurs and avialians; so many that it is difficult to sort out exactly what features go on which side. But since all these animals (Archaeopteryx, Xiaotingia, Anchiornis, Rahonavis, Microraptor, etc.) are all anatomically very similar, we have a pretty good idea of the general appearance of the common ancestor of both groups.
2011 was a good year for large troodontids. For many years all the new troodontid discoveries have been of tiny early members of the group. Finally we are beginning to get new types of large, advanced troodontids from the Late Cretaceous: in other words, Troodon-like troodontids. These include Linhevenator of China and Talos of Utah. These genera help us better understand the larger troodontid form. (By the way, almost certainly the name "Troodon" is being applied to too many species. When the skeletons (rather than just teeth or a few isolated bones) of the different troodontids of western North America over the last 15 million years of the Cretaceous are discovered, we'll find that there are several different genera among them; some closer to Talos, some to Saurornithoides, etc. There are a few genus names out there which may return: Stenonychosaurus, Pectinodon, Polydontosaurus. Or, if necessary, new names will be coined.)
NEW December 2013: New Archaeopteryx-like dinosaurs from the late Middle/early Late Jurassic of China have been discovered: Aurornis and Eosinopteryx. These are known from good skeletons. However, the new information doesn't actually solve the problem of where they fit! It is still uncertain if these form their own clade Archaeopterygidae or not, and if so if archaeopterygids (or the individual genera) are primitive deinonychosaurs or primitive avialians (or both!). Similarly, the scansoriopterygids continue to fail to stay in any particular place on the family tree for very long!
Work continues on whether, and how well, the primitive deinonychosaurs and primitive avialians could fly. It was suggested by Michael Habib and Justin Hall that the long leg feathers of Microraptor were useful in steering and controlling the direction of flight rather than as wings as such. If so, this suggests that MANY of the early deinonychosaurs and avialians may have used them in a similar fashion, as long leg feathers were really quite common.
NEW January 2015: New complications in the world of deinonychosaurs and avialians. Based on the new analysis by Brusatte and colleagues, several important new conclusions:
It is not certain if troodontids and dromaeosaurids form their own clade, or if troodontids are closer to avialians, or if dromaeosaurids are closer to avialians. A three-way split seems the safest bet for the moment.
In this newest go-round, Archaeopteryx does fall out as the oldest and most primitive avialian, and Anchiornis, Xiaotingia, Eosinopteryx, Aurornis, and the like are a group of primitive troodontids.
On the other hand, Balaur is coming out as a primitive avialian in more and more analyses.
In additional news, Archeroraptor (the dromaeosaurid from the latest Cretaceous of North America) has been named, and Pectinodon is back as the name for the troodontid from the same rocks (presently only known from teeth and a few isolated bones.)
Chapter 21: Avialians (Birds)
New Mesozoic birds continue to be found and described. I have included a more up-to-date classification of early bird groupings in the revised version of the genus list.
One aspect of the new discovery of early birds, and of bird-like primitive deinonychosaurs, is that it is getting less and less certain that Archaeopteryx really WAS a bird: that is, that it was more closely related to today's birds than were deinonychosaurs.
Work by American paleontologist Julia Clarke and her Chinese colleagues Zonghe Zhou and Fucheng Zhang in 2006 has shown that the typical modern bird tail (with a fan of feathers coming off of a pygostyle) actually shows up later in bird evolution than we used to think. Confuciusornis and the enantiornithines seem to lack this adaptation, which modern birds use to help them steer in flight and in landing.
NEW January 2008: The flying ability of early avialians (like Confuciusornis) and possible avialians (like Archaeopteryx) may not have been very good at all. A recent study by Phil Senter (Fayetteville State University) shows that these early feathered dinosaurs could not effectively flap their arms in the right way to generate lift, so that they couldn't have had any sort of sustained powered flight. True powered flight would have shown up later, among the common ancestors of enatiornithines and more specialized birds.
NEW December 2011: As discussed for Chapt. 20, there is now stronger evidence that Archaeopteryx is not an avialian at all, but is instead a primitive deinonychosaur. If the archaeopterygids are removed from Avialae, than most of the primitive members of this group would seem to be omnivores at most, and more often strict herbivores (like oviraptorosaurs, therizinosaurs, and other non-deinonychosaur maniraptorans, in fact).
Jingmai O'Connor (of the Institute of Vertebrate Paleontology and Paleoanthropology, Beijing), her former advisor Luis Chiappe (of the Los Angeles County Museum), and her former classmate Alyssa Bell, have presented the most extensive-ever analysis of Mesozoic birds. In general the groupings they discovered resemble those in my book and previous versions of the appendix (indeed, earlier smaller versions of their study was the source of a lot of that!) Some of their new conclusions:
At present it cannot be established which are more closely related to advanced birds: the omnivoropterygiforms (Sapeornis and Didactylornis) or the confuciusornithids.
There is still a lot of confusion about the interrelationships among the enantiornithines
The hesperornithiforms are more closely related to living birds than is Ichthyornis: the reverse of most recent studies over the last couple of decades.
Then there was the bird that (probably) wasn't. In the middle of the year, a new genus of giant Mesozoic bird (known only from its jaw) was described: Samrukia of the Late Cretaceous of Kazakhstan. However, later the same year French paleontologist Eric Buffetaut challenged the idea that this is a bird, and instead proposed it to be a pterosaur jaw. Curiously, neither paper has actually been printed on paper yet: both represent online "early" versions of papers later to be printed in physical, hardcopy journals. Because of the analysis of Buffetaut, I am leaving Samrukia out of my appendix list until new data shows it is indeed a bird (or other dinosaur).
NEW January 2015: Yes, Samrukia does seem to be a pterosaur. On the other hand, Archaeopteryx does indeed seem to be a primitive bird (and a new specimen was unveiled.) As mentioned above, Balaur of the latest Cretaceous of Romania appears to have been a weird large flightless long-tailed bird rather than a weird dromaeosaurid.
Chapter 22: Prosauropods (Primitive Long-Necked Plant-Eating Dinosaurs)
Many important new studies of these dinosaurs were published in 2006 and 2007. Here are some of the highlights:
Most recent phylogenetic analyses agree that there are three basic types of "prosauropod". There are the early small-bodied primitive forms like Saturnalia, Efraasia, Thecodontosaurus, and Pantydraco (the last one was still considered a species of Thecodontosaurus when the book went to the printers!). There are the "core prosauropods": larger dinosaurs including Plateosaurus, Massospondylus, Riojasaurus, Lufengosaurus, Coloradisaurus, and their closest relatives. And there are the "near-sauropods": Yunnanosaurus, Anchisaurus, Melanorosaurus and other dinosaurs more similar to sauropods than to "core prosauropods". The different analyses disagree mainly on whether the core prosauropods from their own group (with all members more closely related to each other than to other types of dinosaurs), or if some are closer to true sauropods and some are more distantly related.
Great new skeletons of all three types of prosauropod have been found, and have given us a better view of their anatomy. For example, the primitive sauropodomorphs may have lacked a cheek, but the core prosauropods and the near-sauropods seem to have had a smaller version of the skin or muscular cheek that ornithischians independantly evolved. This would help them keep food in their mouths as they ate. Also, analyses of the forelimbs of these dinosaurs by Matt Bonnan (Western Illinois University), Phil Senter (Fayetteville State University), and Adam Yates (University of Witwatersrand) show that the primitive sauropodomorphs and the classic prosauropods were probably strictly bipedal, and not the "sometimes bipedal, sometimes quadrupedal" dinosaurs that I wrote about and Luis Rey illustrated. On the other hand (so to speak), the near-sauropods seem to have been capable of walking on all fours, and so were more like they were shown in the book.
While the vast majority of the prosauropods died out by the end of the Early Jurassic, the first Middle Jurassic prosauropod (a new species of the near-sauropod Yunnanosaurus of China) has been described.
NEW January 2008: A newly-discovered core prosauropod or near-sauropod, Lamplughsaura of the Early Jurassic of India, is one of the most completely-known early sauropodomorphs. Study of it will help us understand the evolution of the long-necked plant eaters.
NEW January 2010: Yet more discoveries in this part of the tree. Panphagia from the Late Triassic of Argentina (in fact, the same rocks that Eoraptor, Herrerasaurus, and Pisanosaurus come from) is now the most completely known primitive sauropodomorph. Ongoing studies by Martín D. Ezcurra (Museo Argentino de Ciencias Naturales "Bernardino Rivadavia", Buenos Aires, Argentina) and his colleagues suggest that Panphagia forms a group of successful very primitive sauropodomorphs with other genera previously considered theropods (like Guaibasaurus) or sauropodomorphs (like Saturnalia) or even non-dinosaurs (like Agnosphitys).
At the other end of the "prosauropod" range is 7 m long Aardonyx of the Early Jurassic of South Africa. This is the most advanced of the bipedal sauropodomorphs: the next branch known were capable of some quadrupedal walking. It had lost the cheeks that seem to have been present on most other primitive sauropodomorphs (including primitive sauropods), and thus could "bulk browse" like more advanced sauropods: that is, open its jaws wide and chomp down on a lot of food quickly.
NEW December 2010: The Ezcurra and colleagues analysis mentioned above has now been published, and this group of early sauropodomorphs are now called "Guaibasauridae".
NEW December 2011: As noted earlier, at least some studies now place Eoraptor (previously either a primitive theropod or a non-theropod, non-sauropodomorph primitive saurischian) as a guaibasaurid prosauropod. Add to that Pampadromaeus, a primitive prosauropod that looks something like a stretched-out Eoraptor.
NEW December 2013: Guaibasauridae, we hardly knew thee… Most recent studies have failed to find a single grouping for "Guaibasauridae", and in fact Guaibasaurus itself may be back to being a primitive theropod! The old "guaibasaurids" turn out to be a series of primitive prosauropod groups, with Eoraptor among them.
Chapter 23: Primitive Sauropods (Early Giant Long-Necked Dinosaurs)
Depending on how you classify them the "near-sauropods" mentioned in the previous section are considered early sauropods by some paleontologists.
While it is true that most sauropods had the tooth-to-tooth bite I wrote about, newly described jaws from the most primitive sauropods show that they had a bite more similar to prosauropods (with a wraparound overbite and a small cheek). The later sauropods (called the Eusauropoda, or "true sauropods") evolved the tooth-to-tooth contact and lost their cheeks in favor of a jaw that could open much wider. This makes sense when one considers that the heads of eusauropods were about the same size as the heads of the more primitive sauropods and near-sauropods, but the bodies of the eusauropods were MUCH bigger than the other two! They had to chomp and gulp down food as quickly as they could, and not worry about chopping it up in their jaws.
A whole new group of primitive sauropod (in fact, of primitive eusauropod) was announced while the book was at the presses! First described in December 2006, this group is the Turiasauria, a Jurassic-to-Early Cretaceous clade of dinosaurs from Europe containing gigantic Turiasaurus (at 48 tons the largest known European dinosaur and one of the largest sauropods) as well as Galveosaurus and Losillasaurus. Also, new phylogenetic analyses suggest that at least some of the Middle-to-Late Jurassic extremely long necked Chinese sauropods that I wrote about (such as Omeisaurus and Mamenchisaurus) belong to a clade that also includes Middle Jurassic English Cetiosaurus, Middle Jurassic Argentine Patagosaurus, and Early Jurassic Indian Barapasaurus. (Long-necked Euhelopus may be related to these, but more analyses still place it as close kin to the titanosaurs within Macronaria).
NEW January 2010: One of the newest discoveries among the primitive sauropods is the nearly-complete skeleton of Spinophorosaurus of the Middle Jurassic of Niger. Interestingly, like Shunosaurus, Spinophorosaurus has a spiked club on its tail.
On-going work has not resolved yet if Turaiasauria represents a natural group or not. Mamenchisauridae (the group containing Omeisaurus and Mamenchisaurus) does appear to be well-supported, but Euhelopus is not part of it: instead, this other long-necked Chinese form is close to the ancestry of the Titanosauria.
Chapter 24: Diplodocoids (Whip-Tailed Giant Long-Necked Dinosaurs
NEW January 2008: On p. 199 I wrote "At the time of this writing, no one has put together a complete rebbachisaurid skull or skeleton, so we aren't sure what they looked like." That has changed, thanks to Paul Sereno (University of Chicago), Jeff Wilson (University of Michigan), and their team. They have finally put together the skeleton and utterly-bizarre skull of Nigersaurus. And it is even stranger than Luis drew and I wrote about in our book! Its teeth were apparently replaced at a rate of about once a month: twice as fast as duckbills, which were previously thought to have the fastest rate of tooth replacement in dinosaurs by far. The snout of Nigersaurus was normally head facing straight downwards. It was little (for a sauropod: only the size of an Indian elephant).
NEW July 2008: David Lovelace (University of Wyoming), Scott Hartman (Wyoming Dinosaur Center), and William Wahl (Bighorn Basin Foundation) have recently described Jimbo the Supersaurus, a specimen that demonstrates that Supersaurus is its own distinct genus and that was an apatosaurine (that is, it was a diplodocid that was more closely related to Apatosaurus than to Diplodocus). Additionally, their study shows that "Seismosaurus" is not a distinct genus or species, but is just a very very old, very very large individual of Diplodocus longus.
NEW January 2015: A newly named diplodocid Leinkupal from the Early Cretaceous of Argentina made the news, but not in a good way. It is the first definite diplodocid from the Early Cretaceous, but this got misreported as being a dinosaur which survived a mass extinction (and some reporters even implied it survived the K/Pg extinction!!). It is a cool discovery, but not THAT incredible!
Forgot to mention this last year: Tatouinea, a new rebbachisaurid that shares its name (kind of) with the home world of Luke Skywalker in Star Wars. (Both the dinosaur and the planet are named after the province of Tataouine in Tunisia, which is where the dinosaur was discovered and where the scenes in Star Wars on the planet Tatooine were filmed.
A new specimen of Apatosaurus has been found. It is still under study, but based on the length of its femur it is in the same size range as Supersaurus, Alamosaurus, and Argentinosaurus! So yes, the old "Brontosaurus" may regain its claim to fame in the late 19th Century as the "largest known dinosaur"!!
Chapter 25: Macronarians (Big-Nosed Giant Long-Necked Dinosaurs)
The biggest new discovery (September 2007) for this chapter is 106 to 112 feet (32-34 m) and 70 or more ton Futalognkosaurus, a rival with fellow titanosaurs Argentinosaurus and Puertasaurus for the "Largest Dinosaur of Them All" title. Unlike its giant cousins, however, a good percentage of the skeleton is actually known. Its 46 to 50 foot (14-15 m) long neck is the longest of any dinosaur currently known.
Many new titanosaurs have been discovered, and the relationships between the various groups is still being worked out. It is likely that the relatively simplistic version I used in the book (with a bunch of primitive titanosaurs and the advanced saltasaurids) will be replaced by a classification with multiple groups of advanced titanosaurs. I have used such a system (based on the phylogenetic analysis of Kristi Curry Rogers (Science Museum of Minnesota) and using names already available from previous classifications) in the revised appendix. In this scheme, Titanosauria includes primitive forms like Phuwiangosaurus, Janenschia, Andesaurus, and the gigantic Argyrosauridae (containing Argyrosaurus and Paralititan); the brachiosaurid-like Antarctosauridae (including Argentinosaurus, Alamosaurus, and Antarctosaurus, as well as some titanosaurs whose names DON'T begin with "A"...); and the Lithostrotia. The latter batch are the forms known to have armored backs (not definitely known in the previously-listed types). Among the lithostrotians are the relatively small Saltasauridae (like Saltasaurus and Neuquensaurus) and the more diverse broad-snouted Nemegtosauridae (including Rapetosaurus, Nemegtosaurus, and spike-backed Augustinia, among many others). However, with all the new titanosaurs showing up around the world, our understanding of the diversity of the titanosaurs is going to change even more!
NEW July 2008: Many new titanosaurs have been found in South America. Among them, Futalognkosaurus and Mendozasaurus form a group that has been named the Lognkosauria, or "chief reptiles."
NEW January 2010: Jobaria does not seem to be a macronarian, but instead a more primitive form. Also, it and the megalosaurid Afrovenator are NOT from the Early Cretaceous as long thought: instead, the rocks they are found in come have been redated to the Middle Jurassic.
Re-examination by Mike Taylor of the true North American Brachiosaurus altithorax and the African species typically called "Brachiosaurus" brancai confirms that the latter is distinct from Brachiosaurus proper, and thus is more properly called Giraffatitan.
A comment on p. 205 of the book led to this discussion on the Sauropod Vertebra Picture of the Week blog.
NEW December 2010: Nearly all sauropods are known from the limbs and vertebrae, but skulls are rare. An exception is newly discovered Early Cretaceous brachiosaurid Abydosaurus of Utah, for which four extremely good skulls are known.
NEW December 2011: One of the most complete-ever titanosaur skulls has been found: that of Tapuiasaurus of Brazil.
Also, new specimens of the Late Cretaceous North American titanosaur Alamosaurus show that it is about the same size as the largest known dinosaurs (such as Argentinosaurus, Puertasaurus, Futalognkosaurus, and Ruyangosaurus). This means that Tyrannosaurus (the largest known North American theropod) likely preyed on the largest North American dinosaur, at least on occasion and at least in the southern part of its range (Alamosaurus is not found in the northern states nor in Canada, yet).
NEW December 2013: Some major revisions of the macronarians have been published: some by Michael D'Emic, others by Phil Mannion and colleagues. Unfortunately, they don't always agree with each other, so I've sort of picked and chosen from both of them for the organization of the current appendix. Most of the new studies do support a Brachiosauridae group, and several a Euhelopodidae group; I use both of these. Sauroposeidon is consistently closer to titanosaurs than to brachiosaurids. And Titanosauria is now a strictly Cretaceous group.
NEW January 2015: The big news in macronarian studies is a new big macronarian: Dreadnoughtus. Despite some news reports it is definitely NOT the largest known dinosaur, but it is the most completely known really large dinosaur. And the specimen was not an adult, so it might have achieved the size of Argentinosaurus, Alamosaurus, and kin. In related news, Futalognkosaurus was probably overestimated in size, and was slightly less heavy than Dreadnoughtus.
Chapter 26: Ornithischians
Everything you know about early ornithischians is wrong! Okay, that's an overstatement... But recent work in 2006 and 2007 by the team of Randall Irmis (University of California, Berkeley), Sterling Nesbitt (American Museum of Natural History), William Parker (Petrified Forest National Park), and various colleagues have revolutionized our identification of Triassic ornithischian dinosaurs. Basically, a re-examination of the evidence shows that most of the previously described Triassic "ornithischians" turn out to be from either the near-crocodilian revueltosaurs or the near-dinosaurian silesaurs, two non-dinosaurian types of herbivorous archosaur they we didn't even know about a few years ago. The only remaining definite Triassic ornithischian is the oldest and most primitive one, Pisanosaurus, and an unnamed Triassic member of the Heterodontosauridae.
Oh, wait: turns out another Triassic ornithischian showed up in June 2007! This is Eocursor, a Late Triassic South African ornithischian. Like heterodontosaurids and saurischians, but unlike all the other ornithischians, Eocursor still had the relatively large grasping hand of the earliest dinosaurs. In their initial description of this new dinosaur, Richard Butler (Cambridge University and the Natural History Museum, London), Roger Smith (Iziko South African Museum), and David Norman (Cambridge University) also reanalysed the relationships between various ornithischians, and found that heterodontosaurids are indeed very primitive ornithischians and not ornithopods at all.
NEW January 2010: A couple of important new discoveries among the heterodontosaurids. One of these is tiny Fruitadens of the Late Jurassic of western North America. This is one of the smallest ornithischian dinosaurs ever found: only 70 cm (not quite 28 inches) long!
More spectacular--or maybe just more surprising--is Tianyulong. It was first thought to come from the Early Cretaceous, but now is known to date to the Middle Jurassic. That's not the surprising thing, though. What is the surprising thing is that it had protofeathers along its back (at least). Now up until this discovery there was no definite evidence of feathers or protofeathers outside of the coelurosaurian theropods. (It is true that Psittacosaurus had its quills, but these were not clearly derived from the same anatomical features as protofeathers.) Now we've got fuzzy ornithischians. This means that the common ancestor of all dinosaurs may have sported some fuzz, and that Luis Rey's fuzzy Leaellynasaura on p. 248 may turn out to be accurate!
NEW December 2011: To go along with tiny Fruitadens among the smallest ornithischians, we now have Manidens of the Middle Jurassic of Argentina.
NEW January 2015: From the very beginning of the Jurassic of Venezuela comes Laquintasaura, a primitive ornithischian. It was found in a group of individuals, which may suggest that herding went very far back in ornithischian history.
Chapter 28: Stegosaurs (Plated Dinosaurs)
NEW January 2010: A bizarre new discovery is Miragaia of the Late Jurassic of Portugal: a stegosaur with a long flexible neck like a sauropod.
Chapter 29: Ankylosaurs (Tank Dinosaurs)
The first definite Asian members of Nodosauridae have been described: Zhejiangosaurus and Zhongyuansaurus.
NEW January 2010: Biomechanical studies led by graduate student Victoria Arbour of the University of Alberta calculated the force that ankylosaurids could generate with their club tails. The short answer: they could indeed break bone, especially the big ones.
NEW December 2011: Zhongyuansaurus has subsequently been found out to have been an ankylosaurid, and not a nodosaurid after all. This was part of a massive phylogenetic study of the Ankylosauria by Richard Thompson, Susannah Maidment, and Paul Barrett (all of the Natural History Museum, London) and Jolyon Parish (of the University of Oxford). This study found that basically all ankylosaurs could be divided into Nodosauridae and Ankylosauridae (the old model), and that "polacanthids" do not form their own distinct group. These changes are reflected in the new appendix.
Also, one of the weirdest fossils of any dinosaur was named this year: what looks like the impression of a body of a baby nodosaurid from Early Cretaceous rocks pretty close to where I work! This has been given the name Propanoplosaurus. I'm still suspicious if this is really the impression of a dinosaur at all, and not some sort of other object that our minds are tricking us into thinking is a dinosaur. But I don't have any actual data to back up that suspicion, so until someone can demonstrate this is not the fossil of a dinosaur, I'll give it the benefit of the doubt.
NEW December 2013: The Ankylosauridae, and in particular the Late Cretaceous North American ankylosaurids, have been re-examined by Victoria Arbour, who has found that we have been "lumping" many different distinct genera into "Euoplocephalus". So old, early 20th Century names like Anoplosaurus, Scolosaurus, and Anodontosaurus are back! A HUGE new study of ankylosaur interrelationships is on its way, but hasn't come out by the time I write this. A few aspects of this (such as the diversity of Struthiosaurinae) are included in this version of the appendix, and when the analysis is out I will revise the entire appendix accordingly.
NEW January 2015: Two new ankylosaurids, and two new dinosaurs for "Z" in your "Dinosaurs A-to-Z"-type encyclopediae: Ziapelta of New Mexico and Zaarapelta of Mongolia.
Chapter 30: Primitive Ornithopods (Primitive Beaked Dinosaurs)
The analysis of ornithischian dinosaurs by Butler, Smith and Norman (see the comments on Chapter 26 above) indicates that "Othnielia" (recently renamed Othnieliosaurus by Peter Galton), Agilisaurus, and Hexinlusaurus are all non-ornithopods, but are simply Jurassic relatives of the Ornithopoda and Marginocephalia. Jeholosaurus, on the other hand, may be a true ornithopod.
Bob Bakker (currently at the Houston Museum of Natural History) once speculated that the primitive ornithopod (or ornithopod-like dinosaur) Drinker was a burrower. More recently, a newly discovered (first described in March 2007) Early Cretaceous ornithopod Oryctodromeus was found actually buried inside its burrow.
NEW December 2011: South Korea finally has some dinosaur genera first names from that nation. One of these is the small Late Cretaceous ornithopod Koreanosaurus, a close relative of the burrowing "zephyrosaurs" of North America.
Another small Asian ornithopod named this year, Haya of the Late Cretaceous of China, seems to form a group with the older Jeholosaurus and Changchunsaurus (also of China).
NEW December 2013: Caleb Brown and colleagues have restudied the material of the small ornithopods, and find that the burrowing Orodrominae (what I called "zephyrosaurs") and the Thescelosaurinae (which includes Thescelosaurus and the Asian "jeholosaurs") form a single major group, Thescelosauridae.
NEW January 2015: New shaking around the primitive ornithopod and primitive neornithischian part of the tree. In the latest studies, Orodrominae turns out to be the most primitive ornithopods, followed by Jeholosauridae (a group known from the Cretaceous of Asia). Thescelosaurus is much more closely related to iguanodonts than these dinosaurs.
The biggest news in primitive neornithischian studies, though, is Kulindadromeus of the Jurassic of Siberia. In terms of its skeleton it is nothing special: it is a lot like Hexinlusaurus or Othnieliosaurus. But because it was found in very fine grained lake sediments, its body covering was preserved. And there is a LOT of stuff going on in body covering: some rounded scales, some plate-like scales; some ring-like scales on its tail (sort of like the ring-like scales the feet of some birds); simple fuzz as in Tianyulong and many theropods; and bizarre plates with fuzz coming out from them. This shows that a) even more ornithischians than we knew were fuzzy and b) the variety of body coverings of dinosaurs (even the very same dinosaur!) could be very complex. It also points to the real possibility that the ancestor of ALL dinosaurs was fuzzy, or at least could grow fuzz on some parts of its body. Different groups of dinosaurs did this to different degrees: at present there is no evidence for fuzz in sauropodomorphs, for instance, and big hadrosaurids and ceratopsids were definitely covered mostly in scales. But fuzz is no longer just a theropod trait. (Okay, Tianyulong already proved that, but this reinforces it!).
Oh, and Kulindadromeus is known from many, many individuals, so we will know lots more about its anatomy and growth in the near future.
Chapter 31: Iguanodontians (Advanced Beaked Dinosaurs)
"Iguanodon" atherfieldensis was slender European iguanodontian once considered a species of Iguanodon. However, recent work by Gregory Paul suggests it is actually more closely related to hadrosaurids than to Iguanodon proper, and so in late 2006 he gave it its own genus name: Mantellisaurus. We will have to see if that name stands, or if instead this species turns out to be the same as the previously-named but poorly-known Vectisaurus.
In the past, when you have seen the skull of the Late Jurassic iguanodontian Camptosaurus, it turns out you've seen the wrong dinosaur! Re-examination of the skull by Kathleen Brill and Kenneth Carpenter (Denver Museum of Nature & Science) published in late 2006 shows that the long, squared off skull that everyone (including I and Luis Rey) thought was Camptosaurus was really from a younger, Early Cretaceous dinosaur. Brill and Carpenter have given this dinosaur the name Theiophytalia. Thankfully, however, the actual Camptosaurus skull is now known from a nearly complete specimen, and it is found to be more triangular (something like the head of Dryosaurus, only larger).
Although it is in the genus list in the book, I think it is worth mentioning one of the strangest iguanodontians here as well. That is Lanzhousaurus of Early Cretaceous China. While most iguanodontians evolved more but smaller teeth over time, Lanzhousaurus decided to "buck the trend" and developed fewer but enormous teeth. In fact, these are the largest teeth known of any herbivorous dinosaur.
NEW January 2008: Greg Paul has additionally recognized two new Early Cretaceous dinosaurs that were once considered species of Iguanodon: slender Dollodon bampingi of Belgium and Dakotadon lakotaensis of the United States.
NEW December 2010: The break-up of Iguanodon continues, as more and more species are assigned to their own genera. Darren Naish has written an series of excellent reviews of this topic. Basically, the name "Iguanodon" was used in the past to cover a wide variety of Early Cretaceous iguanodontians: some were more distantly related to hadrosaurids than true Iguanodon and others closer. Among the more notable is tall-spined Hypselospinus.
Brand new (well, newly discovered) iguanodontians that were never part of the classic "Iguanodon" genus have been unearthed from Early Cretaceous-aged rocks of Utah, and named Hippodraco and Iguanacolossus.
NEW December 2011 After the "Iguanodon Explosion" of the last few years, there has been a bit of an implosion. Dave Norman (Cambridge University) has been re-examining much of the material once considered "Iguanodon", and agrees that there are several dinosaur genera represented here, but perhaps not as many as thought in the last few years. He considered "Dollodon" to belong to Mantellisaurus, and "Kukufeldia" and "Sellicoxa" to belong to Barilium. Furthermore, Andrew McDonald (a graduate student at the University of Pennsylvania) suggests that "Proplanicoxa" is simply a specimen of Mantellisaurus. These changes are reflected in the Winter 2011 version of the appendix.
However, 2011 is the year of the "Camptosaurus" explosion. Andrew McDonald (a student at the University of Pennsylvania) has examined the primitive ornithopods once considered to be species of Camptosaurus, and has found instead that some of these are closer to Iguanodon and the hadrosaurs than they are to true Camptosaurus. Thus, the former "Camptosaurus" aphanoecetes is now Uteodon, and "Camptosaurus" depresses is now Osmakasaurus.
NEW January 2015: What is new in the realm of iguanodonts? Dave Norman's clarification of the "Iguanodon Explosion" continues. In his analyses, Iguanodon bernissartensis contains "Dollodon seelyi"; Mantellisaurus atherfieldensis contains Vectisaurus, Sphenospondylus, Proplanicoxa, Dollodon proper, Mantellodon carpenteri, and some of the specimens once called Darwinsaurus; and Hypselospinus fittoni contains Wadhurstia, Huxleysaurus, and most of the specimens called Darwinsaurus. Barilium dawsoni contains Kukufeldia (possibly), Torilion (unquestionably), and Sellacoxa. One issue that isn't resolved is the position of Iguanodon anglicus (the ORIGINAL Iguanodon species): is it actually closer to I. bernissartensis than to these other forms?
In his reanalysis, Iguanodon, Barilium, and Mantellisaurus are all each other's closest relatives, and together with Jinzhousaurus and Bolong of China and Proa of Spain form a group for which the proper name would be "Iguanodontidae". Hypselospinus is not an iguanodontid, and instead is farther from hadrosaurs than are the iguanodonts.
Norman also has introduced several useful new group names within Ornithpoda: Clypeodonta, for all ornithopods closer to Parasaurolophus than to Thescelosaurus; Neoiguanodontia, for the group uniting Hypselospinus and hadrosaurids (and thus containing the iguanodontids as well); and Hadrosauromorpha, for all dinosaurs closer to Parasaurolophus than to Probactrosaurus.
In small ornithopod news, newly named is Eousdryosaurus, a dryosaurid from Portugal.
Chapter 32: Hadrosauroids (Duckbilled Dinosaurs)
NEW January 2010: Some changes in terminology for future editions of this chapter: I will probably call the whole chapter "Hadrosauria", and where I used "Hadrosaurinae" I will use "Saurolophinae" following work by Albert Prieto-Marquez of the American Museum of Natural History. (Hadrosaurus may actually be more distantly related to other "hadrosaurines" than the lambeosaurines are!) These changes, and some changes among the interrelationships of the duckbills, will be reflected in the revised 2010 genus list.
A new discovery is the primitive Italian hadrosaurian Tethyshadros, one of the most complete skeletons of a large Mesozoic dinosaur from Europe.
NEW December 2011: A new study by Nic Campione (University of Toronto) and David Evans (Royal Ontario Museum) shows that Anatotitan is just the adult stage of Edmontosaurus annectens, and not a distinct genus. (Okay, in the appendix I place this species back in its old genus name Anatosaurus. True Edmontosaurus regalis is an older dinosaur.)
NEW December 2013: The lambeosaurines on the cover the book now have a name! When Luis Rey drew them, we went by the idea that the giant lambeosaurine specimens from Baja California were actually Lambeosaurus. However, new studies show that this dinosaur is its own distinct genus, Magnapaulia. The "unicorn" crested lamebosaurine Tsintaosaurus turns out to have a very different kind of crest.
NEW January 2014: Forgot these in the last update! Even well-known duckbills like Edmontosaurus can have surprises: for instance, at least some of them had a small fleshy comb on the tops of their heads. Also, it turns out that hadrosaurid chewing movements were even more complex than I discussed in the book.
NEW January 2015: "Hadrosaurinae" is back, probably. Some recent analyses do find that Hadrosaurus is closer to Saurolophus, Edmontosaurus, and company than to Lambeosaurus, so the flat-head dinosaurs are back to being "hadrosaurines".
A new Mongolian dinosaur Plesiohadros has been described; it is not quite a hadrosaurid, but very closely related to them. It is also the largest dinosaur found so far among the desert deposits of Mongolia.
A new review of the giant Chinese edmontosaurinin hadrosaurine Shantungosaurus confirms what most people suspected: that "Zhuchengosaurus" and "Huaxiaosaurus" were just specimens of Shantungosaurus.
A new study by Scott Persons and Phil Currie of the University of Alberta suggests that hadrosaurs--although slower than tyrannosaurs--were endurance runners ("marathoners"). This may have helped them survive in a world with faster predators: if they managed to escape for long enough, the tyrant dinosaurs would get winded and give up the chase.
Chapter 33: Pachycephalosaurs (Domeheaded Dinosaurs)
Jack Horner (Montana State University) has recently suggested that Pachycephalosaurus, Stygimoloch, and Dracorex are all different growth stages (basically adult, "teenager", and kid) of the same species. I think that this idea has a lot going for it, and look forward to this study being presented in more detail. (As I mention in the book, dinosaurs had a lot of growing up to do in their life, so they went through a lot of different sizes and shapes between hatchling and adulthood. So many different "species" of dinosaurs will likely turn out to be just different growth stages).
NEW January 2010: Related to the above study, new work suggests that Homalocephale is just the juvenile stage of Prenocephale.
NEW December 2011: Functional studies by Eric Snively (Ohio University) and Jessica Theodor (University of Calgary) strongly support the idea that at least some pachycephalosaurs were in fact headbangers.
NEW January 2015: Forgot to get this in last year: a pair of studies by Joseph Peterson (University of Wisconsin) and colleagues supports the hypothesis that head-banging, not erosion after death caused the damage seen on the domes of pachycephalosaurs.
Chapter 34: Primitive Ceratopsians (Parrot and Frilled Dinosaurs)
Plenty of new primitive ceratopsians have been discovered, including the first European ceratopsian fossils (teeth only, but resembling most closely those of leptoceratopsids, announced in July 2007). As more and more of these are discovered, they highlight the fact that ceratopsians spent most of their history as small dinosaurs under the shadows of sauropods, thyreophorans, and iguanodontians. It is only at the very end of the Age of Dinosaurs, and only in western North America, that they became a major group of large-bodied dinosaur.
NEW December 2010: Additional definite primitive ceratopsians from Europe has been discovered. Ajkaceratops from the Late Cretaceous of Hungary is a cousin to Asian Bagaceratops.
NEW January 2015: You want cute? We've got your cute dinosaurs right here: Aquilops of the Early Cretaceous of western North America. Granted, the only known fossil was not fully grown, but this little dinosaur represents the oldest and most primitive ceratopsian from North America. (Hey! Want your own Aquilops skull? Here are directions to making your own version with a fancy 3D printer or a good old-fashioned paper printer!)
Chapter 35: Ceratopsids (Horned Dinosaurs)
NEW January 2008: Eotriceratops xeriinsularis is a newly described ceratopsine very closely related to Triceratops (heck, possibly even ancestral to that later and more famous dinosaur!).
NEW January 2010: With the discovery that primitive centrosaurines (like Albertaceratops) and advanced non-ceratopsid ceratopsians (like Zuniceratops) had long brow horns suggests that poorly-known Ceratops itself may not be closely related to the other dinosaurs that are called "Ceratopsinae" in the book. Therefore, in future versions I will stick to more common use and call Triceratops, Chasmosaurus, and their kin "Chasmosaurinae".
It has been suggested that Turanoceratops from Uzbekistan is a chasmosaurine ceratopsid: if true, it would be the first true ceratopsid from outside of western North America. However, it might be a Zuniceratops-like near-ceratopsid.
On-going research is suggesting that Torosaurus may not be a distinct type of dinosaur, but rather just a fully-adult Triceratops. This is an interesting idea, and I await the full analysis of this idea.
NEW December 2010: The "Torosaurus is just the adult form of Triceratops" paper mentioned above is finally published. As with most things in Science, you can't say that it is 100% proven. However, my personal opinion is that the scientific ball is in the opposition's court now: it is up to those who reject this claim to give positive evidence showing that this hypothesis is incorrect. One amusing (and at times very frustrating) thing was that the news media and blog-commentators often got the implications of this totally backwards! There were lots of articles saying "Triceratops never existed" and that we have to call this dinosaur Torosaurus now. If only those reporters had read my book!! (Okay, or any other book that explains taxonomy...) It isn't whether the name is associated with an adult or not, or a bigger skeleton or not, or a cooler name or not, which is the deciding factor. It is just the date of publication. Since Triceratops was named in 1889, and Torosaurus in 1891, "Triceratops" is the proper name.
(Also, in that same paper, the authors indicate that "Diceratops" aka "Diceratus" aka "Nedoceratops" belongs in Triceratops, too. I definitely concur on that one!)
While we may have "lost" the name "Torosaurus", we gained a dozen brand new genera of Ceratopsidae this year, by far the most ever added to any of the traditional dinosaur "families" in any year! Some, like Mojoceratops, Vagaceratops, and Rubeosaurus were new names for specimens already known (and referred to Chasmosaurus, Chasmosaurus, and Styracosaurus, respectively). Others, though, were brand new: Mexican Coahuilaceratops with horns even more massive than Triceratops; tall-spiked Diabloceratops of Utah; Kosmoceratops and Utahceratops, also of Utah; little Tatankaceratops (which I suspect will wind up being a juvenile Triceratops); and, in the last days of 2010, Titanoceratops, a giant older New Mexican relative of Triceratops (almost as big as that dinosaur), based on a skeleton previously considered to be a giant "Pentaceratops". And for the first time, confirmation of a definite Asian ceratopsid, the Chinese centrosaurine Sinoceratops.
NEW December 2011: New data for and against Nedoceratops being its own distinct genus (rather than a growth stage of Triceratops) were presented this year. The number of new genera has slowed down a bit, with Spinops of the Late Cretaceous of Alberta being the only new entry. (However, several more are on their way for 2012 and/or 2013!)
NEW December 2013: Even more new centrosaurines and chasmosaurines: see the appendix for details! Also, still no resolution on the whole "Is Torosaurus just a grown-up Triceratops?" thing. Both sides have stated their case, but the evidence doesn't decisively fall in one camp. Yet.
NEW January 2015: Got a spare hour? Watch and hear about the "Triceratops vs. Torosaurus" debate.
Chapter 36: Dinosaur Eggs and Babies
In September 2007, an assemblage of Psittacosaurus fossils found together was described. This collection of skeletons of the little ceratopsian were buried together under volcanic ashes. Examination of the fossils by Zhao Qi (Institute of Vertebrate Paleontology and Paleoanthropology, Beijing), Paul Barrett (The Natural History Museum, London), and Dave Eberth (Royal Tyrrell Museum of Palaeontology) indicates two different age groups of youngsters: some about 1 and a half years old and the others about 3 years old. This shows that they were born at two different times, but the fact that they were buried together may indicate that they lived together as a big family.
NEW December 2010: Yet another peril that baby dinosaurs had to face: early snakes!.
NEW December 2013: A new discovery is nesting colonies of therizinosaurs. And check out this cute baby Chasmosaurus!
NEW January 2015: I should have included this before: watch the changes in Tarbosaurus as it grew up.
Chapter 37: Dinosaur Behavior: How Did Dinosaurs Act, and How Do We Know?
NEW December 2010: New news on the locomotion of tyrannosaurs front. A new study by University of Alberta graduate student Scott Persons and his advisor Phil Currie shows that the tail muscles of Tyrannosaurus and its kin were bigger than previously thought. Although it added more weight to the dinosaurs, it also added power, and suggests that they may have been faster than some previously calculated.
Other work by Casey Holliday of the University of Missouri and colleagues shows that people have been underestimating the amount of cartilage between bones in dinosaurs. This effects things like our calculations of their height and length, as well as of the mechanics of their joints.
Chapter 38: Dinosaur Biology: Living, Breathing Dinosaurs
NEW December 2010: As I discuss in Chapter 38, there is a lot of evidence suggesting that dinosaurs had metabolisms more like modern mammals and birds than like cold-blooded lizards and snakes and turtles. Similar evidence exists for dinosaurs weird-looking flying cousins, the pterosaurs. This year, however, comes evidence in the form of the chemical make-up of the bones of marine reptiles that the advanced tuna-shaped ichthyosaurs and the paddle-finned plesiosaurs may also have been warm-blooded, even though the more eel-shaped primitive ichthyosaurs and the fingered-and-toed relatives of plesiosaurs were not. Additionally the mosasaur sea-lizards of the Late Cretaceous may have also been warm-blooded as well, but the evidence for this is less strong. It is worth noting that modern day tuna and billfish (sailfish, swordfish, etc.) are essentially "warm-blooded fish", and evolved a very similar body shape to the advanced ichthyosaurs.
NEW December 2011: New lines of evidence that dinosaurs had warm-blooded metabolisms come from new studies of the body temperatures of sauropods and the presence of a high number of nutrient foramina (blood vessel openings) in dinosaus compared to cold-blooded animals.
In other aspects of their biology, at least some dinosaurs and pterosaurs may have been nocturnal based on aspects of the relative proportions of their eyes.
NEW January 2015: New analyses of the insides of the noses of pachycephalosaurs show that their nasal turbinates would help cool the blood going to the brain. This was actually probably widespread in all dinosaurs, but it is easier to study in pachycephalosaurs due to the very solid skulls which preserve many details.
Were dinosaurs (including Archaeopteryx and other early avialians) intermediate between fully cold and fully warm-blooded? A new study suggests this. This supposed "mesothermy" in dinosaurs (a term originally coined by Dr. Scott Sampson) is an intriguing possibility, but needs further study. One thing to point out is that the authors agree that dinosaurs generated heat internally (and thus were endothermic by definition), but that their control over the body temperature wasn't as strong as in homeothermic animals.
Chapter 40: Life in the Jurassic Period
NEW December 2010: Chris Noto, currently of Grand Valley State University in Michigan, has published an examination of the differences and similarities of the different communities of dinosaurs in the Late Jurassic, and found that brachiosaurs and their kin preferred more arid (dry) habitats, while stegosaurs and diplodcoids preferred wetter ones.
NEW December 2011: A new study of the chemical composition of sauropod teeth shows that at least some of them seem to have migrated to the mountains and back to the lowlands in the course of a year.
Chapter 41: Life in the Cretaceous Period
NEW December 2011: Tyler Lyson and Nick Longrich at Yale University have presented a study of the paleoecology of dinosaurs from the Hell Creek Formation, the very last dinosaur-dominated community of North America. They found that even a relatively small region they could detect habitat preferences for different types of dinosaurs, with hadrosaurids and the primitive ornithopod Thescelosaurus closer to the rivers, and Triceratops in the drier land. (Carnivores like Tyrannosaurus didn't have a strong preference: after all, meat is meat, no matter where you eat it!)
II. Updated Dinosaur Genus List
The genus update is available here.
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Yangchuanosaurus (Dinosaurs of China by Safari Ltd) – Dinosaur Toy Blog
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2009-12-08T16:51:29+00:00
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https://dinotoyblog.com/yangchuanosaurus-dinosaurs-of-china-by-safari-ltd/
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Yangchuanosaurus is sorely underrepresented as a dinosaur toy so I’m glad Safari Ltd decided to make one as part of their Dinosaurs of China line (and moreover, make it good!) Yangchuanosaurus was a large theropod from the Late Jurassic of China – the T. rex of it’s time – and lived alongside other contemporary Chinese dinosaurs such as Sinraptor and the behemoth sauropod Mamenchisaurus.
The sculpt is very nice and its obvious that attention has been given to the texture: this guy scales all over his body. His legs aren’t deathly thin like some of Safari Ltd’s Carnegie replicas and they look beautiful. The head is well done, I love the nostrils and the thick ridges going down the snout. It also has noticeable external ears. Unlike many other Safari dinosaurs, this Yangchuanosaurus isn’t covered in wrinkles; the only wrinkles visible are behind the right side of the head. The pose is creative and makes a welcome change. It is actually in a natural pose, not so much “RAWR look at me I’m a dinosaur!”. It looks to be wiping it’s mouth off after taking a drink from a nice prehistoric stream.
This is one of the handful of Safari theropods that actually has its mouth closed and it looks great. At first glance, the arms seem to be of two different lengths, but when you measure them out, they are really equal. As with the the other dinosaurs of China fugures, this Yangchuanosaurus was distributed in a box together with a special display stand featuring the skeleton of the animal. Although this is rather crudely sculpted, it makes for an eye-catching display.
As with any dinosaur sculpt, this replica has its faults. The left side of the head has more teeth than the right side, and the tail is particularly thin. There is also the absence of a dew claw on each foot.
The paint job on this figure is very nice too, and it fits the sculpt well. It is mostly a (slightly greenish) brown all over, and it is slightly more green on the head than other places on the body. The eyes are a fierce forest green with circular black pupils, and to give the illusion of sunlight hitting the eyes, there is a little white dot above each pupil. Fading black bands adorn this beast’s flanks. The ribcages are highlighted in a bright orange yellow color. The claws, unfortunately, are unpainted, and at first glance they look like part of the digits themselves. Oddly, the line of the mouth is highlighted in black– This is the only Safari theropod figure that I can think of that has this feature. This figure is about 8 inches long.
Overall, the ‘Dinosaurs of China’ Yangchuanosaurus is a worthwhile figure – I rate it 9/10. However, since he has a very non-hunting pose with a closed mouth and docile look, he probably didn’t go over so well with children, which may explain why he was discontinued. If you can find him you better get him soon because I wouldn’t be surprised if he becomes highly sought after.
Edit – this post was updated by ‘Plesiosauria’ on 2/9/2012
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https://kids.kiddle.co/List_of_dinosaur_genera
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List of dinosaur genera facts for kids
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Learn List of dinosaur genera facts for kids
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en
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https://kids.kiddle.co/List_of_dinosaur_genera
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Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago, although the exact origin and timing of the evolution of dinosaurs is the subject of active research. They became the dominant terrestrial vertebrates after the Triassic–Jurassic extinction event 201.3 million years ago; their dominance continued throughout the Jurassic and Cretaceous periods. The fossil record demonstrates that birds are modern feathered dinosaurs, having evolved from earlier theropods during the Late Jurassic epoch. Birds were therefore the only dinosaur lineage to survive the Cretaceous–Paleogene extinction event approximately 66 million years ago. Dinosaurs can be divided into avian dinosaurs (birds) and non-avian dinosaurs, which are all dinosaurs other than birds. Birds are feathered theropod dinosaurs and constitute the only known living dinosaurs.
This list of dinosaurs is a comprehensive listing of all genera that have ever been considered to be non-avian dinosaurs, but also includes some dinosaurs of disputed status (avian? or non-avian?, where "avian" refers to the clade Avialae), as well as purely vernacular terms.
The list includes all commonly accepted genera, but also genera that are now considered invalid, doubtful (nomen dubium), or were not formally published (nomen nudum), as well as junior synonyms and genera that are no longer considered dinosaurs. Many listed names have been reclassified as everything from true birds to crocodilians to petrified wood. The list contains 1764 names, of which approximately 1328 are considered either valid dinosaur genera or nomina dubia.
Scope and terminology
There is no official, canonical list of all non-avian dinosaur genera. The closest is the Dinosaur Genera List, compiled by biological nomenclature expert George Olshevsky, which was first published online in 1995 and was regularly updated until June 2021. The most authoritative general source in the field is the second (2004) edition of The Dinosauria. The vast majority of names listed below are sourced to Olshevsky's list, and all subjective determinations (such as junior synonymy or non-dinosaurian status) are based on The Dinosauria, except where they conflict with primary literature. These exceptions are noted.
Naming conventions and terminology follow the International Code of Zoological Nomenclature. Technical terms used include:
Junior synonym: A name which describes the same taxon as a previously published name. If two or more genera are formally designated and the type specimens are later assigned to the same genus, the first to be published (in chronological order) is the senior synonym, and all other instances are junior synonyms. Senior synonyms are generally used, except by special decision of the ICZN (see Tyrannosaurus), but junior synonyms cannot be used again for a different genus, even if deprecated. Junior synonymy is often subjective, unless the genera described were both based on the same type specimen.
Nomen nudum (Latin for "naked name"): A name that has appeared in print but has not yet been formally published by the standards of the ICZN. Nomina nuda (the plural form) are invalid, and are therefore not italicized as a proper generic name would be. If the name is later formally published, that name is no longer a nomen nudum and will be italicized on this list. Often, the formally published name will differ from any nomina nuda that describe the same specimen.
Nomen oblitum (Latin for "forgotten name"): A name that has not been used in the scientific community for more than fifty years after its original proposal.
Nomen manuscriptum (Latin for "manuscript name"): A name that appears in manuscript of a formal publication that has no scientific backing.
Preoccupied name: A name that is formally published, but which has already been used for another taxon. This second use is invalid (as are all subsequent uses) and the name must be replaced. Preoccupied names are not valid generic names.
Nomen dubium (Latin for "dubious name"): A name describing a fossil with no unique diagnostic features. As this can be an extremely subjective and controversial designation (see Hadrosaurus), no genera should be marked as such on this list.
A
Contents: A B C D E F G H I J K L M N O P Q R S T U V W X Y Z – See also
Aachenosaurus – subsequently found to be a piece of petrified wood
Aardonyx
"Abdallahsaurus" – nomen nudum, synonym of Giraffatitan
Abdarainurus
Abditosaurus
Abelisaurus
Abrictosaurus
Abrosaurus
Abydosaurus
Acantholipan
Acanthopholis
Achelousaurus
Acheroraptor
Achillesaurus
Achillobator
Acristavus
Acrocanthosaurus
Acrotholus
Actiosaurus – subsequently found to be a choristoderan
Adamantisaurus
Adasaurus
Adelolophus
Adeopapposaurus
Adratiklit
Adynomosaurus
Aegyptosaurus
Aeolosaurus
Aepisaurus
Aepyornithomimus
Aerosteon
Aetonyx – junior synonym of Massospondylus
Afromimus
Afrovenator
Agathaumas – possible synonym of Triceratops
Aggiosaurus – subsequently found to be a metriorhynchid crocodilian
Agilisaurus
Agnosphitys – possibly non-dinosaurian
Agrosaurus – probably a junior synonym of Thecodontosaurus
Agujaceratops
Agustinia
Ahshislepelta
"Airakoraptor" – nomen nudum; Kuru
Ajancingenia – synonym of Heyuannia
Ajkaceratops
Ajnabia
Akainacephalus
Alamosaurus
Alaskacephale
Albalophosaurus
Albertaceratops
Albertadromeus
Albertavenator
Albertonykus
Albertosaurus
Albinykus
Albisaurus – subsequently found to be a non-dinosaurian reptile
Alcovasaurus – possible junior synonym of Miragaia
Alectrosaurus
Aletopelta
Algoasaurus
Alioramus
Aliwalia – junior synonym of Eucnemesaurus
Allosaurus
Almas
Alnashetri
Alocodon
Altirhinus
Altispinax
Alvarezsaurus
Alwalkeria
Alxasaurus
Amanasaurus – possibly non-dinosaurian
Amanzia
Amargasaurus
"Amargastegos" – nomen nudum
Amargatitanis
Amazonsaurus
Ambopteryx
Ammosaurus – junior synonym of Anchisaurus
Ampelognathus
Ampelosaurus
Amphicoelias
"Amphicoelicaudia" – nomen nudum; synonym of Huabeisaurus
"Amphisaurus" – preoccupied name, now known as Anchisaurus
Amtocephale
Amtosaurus – possibly a junior synonym of Talarurus
Amurosaurus
Amygdalodon
Anabisetia
Analong
Anasazisaurus
Anatosaurus – junior synonym of Edmontosaurus
Anatotitan – junior synonym of Edmontosaurus
Anchiceratops
Anchiornis
Anchisaurus
Andesaurus
"Andhrasaurus" – nomen nudum
Angaturama – possible junior synonym of Irritator
"Angloposeidon" – nomen nudum
Angolatitan
Angulomastacator
Anhuilong
Aniksosaurus
Animantarx
Ankistrodon – subsequently found to be a proterosuchid archosauriform
Ankylosaurus
Anodontosaurus
Anomalipes
Anoplosaurus
Anserimimus
Antarctopelta
Antarctosaurus
Antetonitrus
Anthodon – subsequently found to be a pareiasaur
Antrodemus – possibly a synonym of Allosaurus
Anzu
Aoniraptor
Aorun
Apatodon – possibly a junior synonym of Allosaurus
Apatoraptor
Apatosaurus
Appalachiosaurus
Aquilarhinus
Aquilops
Arackar
Aragosaurus
Aralosaurus
Aratasaurus
"Araucanoraptor" – nomen nudum; Neuquenraptor
Archaeoceratops
Archaeodontosaurus
Archaeopteryx – possibly a bird
Archaeoraptor – a chimaera of the bird Yanornis and the dromaeosaur Microraptor
Archaeornis – junior synonym of Archaeopteryx
Archaeornithoides
Archaeornithomimus
Arcovenator
Arctosaurus – subsequently found to be a non-dinosaurian reptile
Arcusaurus
Arenysaurus
Argentinosaurus
Argyrosaurus
Aristosaurus – junior synonym of Massospondylus
Aristosuchus
Arizonasaurus – subsequently found to be a rauisuchian
Arkansaurus
Arkharavia
Arrhinoceratops
Arrudatitan
Arstanosaurus
Asfaltovenator
Asiaceratops
Asiamericana – a fish
Asiatosaurus
Asilisaurus – possibly non-dinosaurian
Astrodon
Astrodonius – junior synonym of Astrodon
Astrodontaurus – junior synonym of Astrodon
Astrophocaudia
Asylosaurus
Atacamatitan
Atlantosaurus
Atlasaurus
Atlascopcosaurus
Atrociraptor
Atsinganosaurus
Aublysodon
Aucasaurus
"Augustia" – preoccupied name, now known as Agustinia
Augustynolophus
Auroraceratops
Aurornis
Australodocus
Australotitan
Australovenator
Austrocheirus
Austroposeidon
Austroraptor
Austrosaurus
Avaceratops
"Avalonia" – preoccupied name, now known as Avalonianus
Avalonianus – subsequently found to be a non-dinosaurian archosaur
Aviatyrannis
Avimimus
Avipes – probably a non-dinosaurian dinosauromorph
Avisaurus – subsequently found to be an enantiornithine bird
Azendohsaurus – subsequently found to be a non-dinosaurian archosauromorph
B
Contents: A B C D E F G H I J K L M N O P Q R S T U V W X Y Z – See also
Baalsaurus
Bactrosaurus
Bagaceratops
Bagaraatan
Bagualia
Bagualosaurus
Bahariasaurus
Bainoceratops
Bajadasaurus
"Bakesaurus" – nomen nudum; Bactrosaurus
Balaur – possibly a bird
"Balochisaurus" – nomen nudum
Bambiraptor
Banji
Bannykus
Baotianmansaurus
Barapasaurus
Barilium
Barosaurus
Barrosasaurus
Barsboldia
Baryonyx
Bashanosaurus
Bashunosaurus
Basutodon – subsequently found to be a non-dinosaurian archosaur
Bathygnathus – a pelycosaur, Dimetrodon
Batyrosaurus
Baurutitan
Bayannurosaurus
"Bayosaurus" – nomen nudum
Becklespinax – junior synonym of Altispinax
"Beelemodon" – nomen nudum
Beg
Beibeilong
Beipiaognathus – chimera of several unnamed dinosaurs
Beipiaosaurus
Beishanlong
Bellusaurus
Belodon – subsequently found to be a phytosaur
Berberosaurus
Berthasaura
Betasuchus
Bicentenaria
Bienosaurus
"Bihariosaurus" – nomen nudum
"Bilbeyhallorum" – nomen nudum; Cedarpelta
Bissektipelta
Bistahieversor
Bisticeratops
"Blancocerosaurus" – nomen nudum, synonym of Giraffatitan
Blasisaurus
Blikanasaurus
Bolong
Bonapartenykus
Bonapartesaurus
Bonatitan
Bonitasaura
Borealopelta
Borealosaurus
Boreonykus
Borogovia
Bothriospondylus
Brachiosaurus
Brachyceratops
Brachylophosaurus
Brachypodosaurus
Brachyrophus – junior synonym of Camptosaurus
Brachytaenius – subsequently found to be a metriorhynchid; junior objective synonym of Dakosaurus or Geosaurus
Brachytrachelopan
Bradycneme
Brasileosaurus – subsequently found to be a non-dinosaurian archosaur
Brasilotitan
Bravasaurus
Bravoceratops
Breviceratops
Brighstoneus
Brohisaurus
Brontomerus
"Brontoraptor" – nomen nudum, synonym of Torvosaurus
Brontosaurus
Bruhathkayosaurus
Bugenasaura – junior synonym of Thescelosaurus
Buitreraptor
Burianosaurus
Buriolestes
Bustingorrytitan
"Byranjaffia" – nomen nudum; Byronosaurus
Byronosaurus
C
Contents: A B C D E F G H I J K L M N O P Q R S T U V W X Y Z – See also
Caenagnathasia
Caenagnathus
Caieiria
Caihong
Calamosaurus
"Calamospondylus" – preoccupied name, now known as Calamosaurus
Calamospondylus
Callovosaurus
Calvarius
Camarasaurus
Camarillasaurus
Camelotia
Camposaurus
"Camptonotus" – preoccupied name, now known as Camptosaurus
Camptosaurus
"Campylodon" – preoccupied name, now known as Campylodoniscus
Campylodoniscus
Canardia
"Capitalsaurus" – nomen nudum
Carcharodontosaurus
Cardiodon
Carnotaurus
Caseosaurus
Cathartesaura
Cathetosaurus — possibly a species of Camarasaurus
Caudipteryx
Caudocoelus – junior synonym of Teinurosaurus
Caulodon – junior synonym of Camarasaurus
Cedarosaurus
Cedarpelta
Cedrorestes
Centemodon – subsequently found to be a phytosaur
Centrosaurus
Cerasinops
Ceratonykus
Ceratops
Ceratosaurus
Ceratosuchops
Cetiosauriscus
Cetiosaurus
Chakisaurus
Changchunsaurus
"Changdusaurus" – nomen nudum
Changmiania
Changyuraptor
Chaoyangsaurus
Charonosaurus
Chasmosaurus
Chassternbergia – junior synonym of Edmontonia
Chebsaurus
Chenanisaurus
Cheneosaurus – junior synonym of Hypacrosaurus
Chialingosaurus
Chiayusaurus
Chienkosaurus – possible junior synonym of Szechuanosaurus
"Chihuahuasaurus" – nomen nudum; Sonorasaurus
Chilantaisaurus
Chilesaurus
Chindesaurus
Chingkankousaurus
Chinshakiangosaurus
Chirostenotes
Choconsaurus
Chondrosteosaurus
Choyrodon
Chromogisaurus
Chuandongocoelurus
Chuanjiesaurus
Chuanqilong
Chubutisaurus
Chucarosaurus
Chungkingosaurus
Chuxiongosaurus
"Cinizasaurus" – nomen nudum
Cionodon
Citipati
Citipes
Cladeiodon – subsequently found to be a non-dinosaurian rauisuchian; synonym of Teratosaurus
Claorhynchus – possibly Triceratops
Claosaurus
Clarencea – subsequently found to be a sphenosuchian; synonym of Sphenosuchus
Clasmodosaurus
Clepsysaurus – subsequently found to be a phytosaur, possibly Palaeosaurus
Coahuilaceratops
Coelophysis
"Coelosaurus" – preoccupied genus name, species "Coelosaurus" antiquus
Coeluroides
Coelurosauravus – subsequently found to be a primitive diapsid
Coelurus
Colepiocephale
"Coloradia" – preoccupied name, now known as Coloradisaurus
Coloradisaurus
"Colossosaurus" – nomen nudum; Pelorosaurus
Comahuesaurus
"Comanchesaurus" – nomen nudum
Compsognathus
Compsosuchus
Concavenator
Conchoraptor
Condorraptor
Convolosaurus
Coronosaurus
Corythoraptor
Corythosaurus
Craspedodon
Crataeomus – junior synonym of Struthiosaurus
Craterosaurus
Creosaurus – junior synonym of Allosaurus
Crichtonpelta
Crichtonsaurus
Cristatusaurus
Crittendenceratops
Crosbysaurus – subsequently found to be a non-dinosaurian archosauriform
Cruxicheiros
Cryolophosaurus
Cryptodraco – junior synonym (unneeded replacement name) of Cryptosaurus
"Cryptoraptor" – nomen nudum
Cryptosaurus
Cryptovolans – junior synonym of Microraptor
Cumnoria
D
Contents: A B C D E F G H I J K L M N O P Q R S T U V W X Y Z – See also
Daanosaurus
Dacentrurus
"Dachongosaurus" – nomen nudum
Daemonosaurus
Dahalokely
Dakosaurus – subsequently found to be a metriorhynchid crocodilian
Dakotadon
Dakotaraptor
Daliansaurus
"Damalasaurus" – nomen nudum
Dandakosaurus
Danubiosaurus – junior synonym of Struthiosaurus
"Daptosaurus" – nomen nudum; early manuscript name for Deinonychus
Darwinsaurus – junior synonym of Hypselospinus or Mantellisaurus
Dashanpusaurus
Daspletosaurus
Dasygnathoides – subsequently found to be a non-dinosaurian archosaur; possible junior synonym of Ornithosuchus
"Dasygnathus" – preoccupied name, now known as Dasygnathoides
Datai
Datanglong
Datonglong
Datousaurus
Daurlong
Daurosaurus – synonym of Kulindadromeus
Daxiatitan
Deinocheirus
Deinodon – possibly Gorgosaurus
Deinonychus
Delapparentia – junior synonym of Iguanodon
Deltadromeus
Demandasaurus
Denversaurus
Deuterosaurus – subsequently found to be a therapsid
Diabloceratops
Diamantinasaurus
Dianchungosaurus – subsequently found to be a crocodilian
"Diceratops" – preoccupied name, now known as Nedoceratops
Diceratus – junior synonym of Nedoceratops
Diclonius
Dicraeosaurus
Didanodon – synonym of Lambeosaurus; possibly a nomen nudum
Dilong
Dilophosaurus
Diluvicursor
Dimodosaurus – junior synonym of Plateosaurus
Dineobellator
Dinheirosaurus – possible junior synonym of Supersaurus
Dinodocus
"Dinosaurus" – preoccupied name for a junior synonym of Brithopus; now a junior synonym of Plateosaurus
Dinotyrannus – junior synonym Tyrannosaurus or some other tyrannosaurid
Diodorus – possibly non-dinosaurian
Diplodocus
Diplotomodon
Diracodon – junior synonym of Stegosaurus
Dolichosuchus
Dollodon – junior synonym of Mantellisaurus
"Domeykosaurus" – nomen nudum, synonym of Arackar
Dongbeititan
Dongyangopelta
Dongyangosaurus
Doratodon – subsequently found to be a crocodilian
Dornraptor
Doryphorosaurus – junior synonym (unneeded replacement name) of Kentrosaurus
Draconyx
Dracopelta
Dracoraptor
Dracorex – junior synonym of Pachycephalosaurus
Dracovenator
Dravidosaurus – possibly non-dinosaurian
Dreadnoughtus
Drinker – junior synonym of Nanosaurus
Dromaeosauroides
Dromaeosaurus
Dromiceiomimus
Dromicosaurus – junior synonym of Massospondylus
Drusilasaura
Dryosaurus
Dryptosauroides
Dryptosaurus
Dubreuillosaurus
"Duranteceratops" – nomen nudum
Duriatitan
Duriavenator
Dynamosaurus – junior synonym of Tyrannosaurus
Dynamoterror
Dyoplosaurus
Dysalotosaurus
Dysganus
Dyslocosaurus
Dystrophaeus
Dystylosaurus – junior synonym of Supersaurus
Dzharaonyx
Dzharatitanis
E
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Echinodon
Edmarka – junior synonym of Torvosaurus
Edmontonia
Edmontosaurus
Efraasia
Einiosaurus
Ekrixinatosaurus
Elachistosuchus – subsequently found to be a rhynchocephalian
Elaltitan
Elaphrosaurus
Elemgasem
Elmisaurus
Elopteryx
Elosaurus – junior synonym of Brontosaurus
Elrhazosaurus
"Elvisaurus" – nomen nudum; Cryolophosaurus
Emausaurus
Embasaurus
Enigmosaurus
Eoabelisaurus
Eobrontosaurus – junior synonym of Brontosaurus
Eocarcharia
Eoceratops – junior synonym of Chasmosaurus
Eocursor
Eodromaeus
"Eohadrosaurus" – nomen nudum; Eolambia
Eolambia
Eomamenchisaurus
Eoneophron
"Eoplophysis" – nomen nudum
Eoraptor
Eosinopteryx
Eotrachodon
Eotriceratops
Eotyrannus
Eousdryosaurus
Epachthosaurus
Epanterias – may be Allosaurus
"Ephoenosaurus" – nomen nudum; Machimosaurus (a crocodilian)
Epicampodon – subsequently found to be a proterosuchid archosauriform, Ankistrodon
Epichirostenotes
Epidendrosaurus – synonym of Scansoriopteryx
Epidexipteryx
Equijubus
Erectopus
Erketu
Erliansaurus
Erlikosaurus
Erythrovenator
Eshanosaurus
"Euacanthus" – nomen nudum; junior synonym of Polacanthus
Eucamerotus
Eucentrosaurus – junior synonym (unneeded replacement name) of Centrosaurus
Eucercosaurus
Eucnemesaurus
Eucoelophysis – possibly non-dinosaurian
"Eugongbusaurus" – nomen nudum
Euhelopus
Euoplocephalus
Eupodosaurus – subsequently found to be a nothosaur synonymous with Lariosaurus
"Eureodon" – nomen nudum; Tenontosaurus
Eurolimnornis – subsequently found to be a pterosaur
Euronychodon
Europasaurus
Europatitan
Europelta
Euskelosaurus
Eustreptospondylus
F
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Fabrosaurus – possibly Lesothosaurus
Falcarius
"Fendusaurus" – nomen nudum
"Fenestrosaurus" – nomen nudum; Oviraptor
Ferganasaurus
"Ferganastegos" – nomen nudum
Ferganocephale
Ferrisaurus
Foraminacephale
Fosterovenator
Fostoria
Frenguellisaurus – junior synonym of Herrerasaurus
Fruitadens
Fujianvenator
Fukuiraptor
Fukuisaurus
Fukuititan
Fukuivenator
Fulengia
Fulgurotherium
Furcatoceratops
Fushanosaurus
"Fusinasus" – nomen nudum; Eotyrannus
Fusuisaurus
"Futabasaurus" – nomen nudum; not to be confused with the formally named plesiosaur Futabasaurus
Futalognkosaurus
Fylax
G
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"Gadolosaurus" – nomen nudum
Galeamopus
Galesaurus – subsequently found to be a therapsid
Galleonosaurus
Gallimimus
Galtonia – subsequently found to be a pseudosuchian; possibly a junior synonym of Revueltosaurus
Galveosaurus – synonym of Galvesaurus
Galvesaurus
Gamatavus — possibly non-dinosaurian
Gandititan
Gannansaurus
"Gansutitan" – nomen nudum; Daxiatitan
Ganzhousaurus
Gargoyleosaurus
Garrigatitan
Garudimimus
Garumbatitan
Gasosaurus
Gasparinisaura
Gastonia
"Gavinosaurus" – nomen nudum; Eotyrannus
Geminiraptor
Genusaurus
Genyodectes
Geranosaurus
Gideonmantellia
Giganotosaurus
Gigantoraptor
"Gigantosaurus" – preoccupied name, now known as Tornieria
Gigantosaurus
Gigantoscelus – Probable junior synonym of Euskelosaurus
Gigantspinosaurus
Gilmoreosaurus
"Ginnareemimus" – nomen nudum; Kinnareemimus
Giraffatitan
Glacialisaurus
Glishades
Glyptodontopelta
Gnathovorax
Gobiceratops – possibly a junior synonym of Bagaceratops
Gobihadros
Gobiraptor
Gobisaurus
Gobititan
Gobivenator
"Godzillasaurus" – nomen nudum; Gojirasaurus
Gojirasaurus
Gondwanatitan
Gongbusaurus
Gongpoquansaurus
Gongxianosaurus
Gonkoken
Gorgosaurus
Goyocephale
Graciliceratops
Graciliraptor
Gracilisuchus – subsequently found to be a non-dinosaurian archosaur
Gravitholus
Gremlin
Gresslyosaurus – possible junior synonym of Plateosaurus
Griphornis – junior synonym of Archaeopteryx
Griphosaurus – junior synonym of Archaeopteryx
Gryphoceratops
Gryponyx
Gryposaurus
"Gspsaurus" – nomen nudum
Guaibasaurus
Gualicho
Guanlong
Guemesia
Gwyneddosaurus – subsequently found to be a tanystrophid
Gyposaurus – possibly a junior synonym of Massospondylus
H
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"Hadrosauravus" – nomen nudum; junior synonym of Gryposaurus
Hadrosaurus
Haestasaurus
Hagryphus
Hallopus – subsequently found to be a crocodylomorph
Halszkaraptor
Halticosaurus
Hamititan
Hanssuesia
"Hanwulosaurus" – nomen nudum
Haplocanthosaurus
"Haplocanthus" – preoccupied name, now known as Haplocanthosaurus
Haplocheirus
Harpymimus
Haya
Hecatasaurus – junior synonym of Telmatosaurus
"Heilongjiangosaurus" – nomen nudum
Heishansaurus
Helioceratops
"Helopus" – preoccupied name, now known as Euhelopus
Heptasteornis
Herbstosaurus – subsequently found to be a pterosaur
Herrerasaurus
Hesperonychus
Hesperonyx
Hesperornithoides
Hesperosaurus
Heterodontosaurus
Heterosaurus – possible synonym of Mantellisaurus
Hexing
Hexinlusaurus
Heyuannia
Hierosaurus
Hikanodon – junior synonym of Iguanodon
Hippodraco
"Hironosaurus" – nomen nudum
"Hisanohamasaurus" – nomen nudum
Histriasaurus
Homalocephale
"Honghesaurus" – nomen nudum later described as Yandusaurus; name later used for a genus of marine reptile
Hongshanosaurus – junior synonym of Psittacosaurus
Hoplitosaurus
Hoplosaurus – junior synonym of Struthiosaurus
Horshamosaurus
Hortalotarsus – possible junior synonym of Massospondylus
Huabeisaurus
Hualianceratops
Huallasaurus
Huanansaurus
Huanghetitan
Huangshanlong
Huaxiagnathus
Huaxiaosaurus – junior synonym of Shantungosaurus
"Huaxiasaurus" – nomen nudum; Huaxiagnathus
Huayangosaurus
Hudiesaurus
Huehuecanauhtlus
Huinculsaurus
Hulsanpes
Hungarosaurus
Huxleysaurus – junior synonym of Hypselospinus
Hylaeosaurus
Hylosaurus – junior synonym of Hylaeosaurus
Hypacrosaurus
Hypselorhachis – subsequently found to be a ctenosauriscid
Hypselosaurus
Hypselospinus
Hypsibema
Hypsilophodon
Hypsirhophus
I
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Iani
Iberospinus
Ibirania
"Ichabodcraniosaurus" – nomen nudum; Shri
Ichthyovenator
Igai
Ignavusaurus
Ignotosaurus – possibly non-dinosaurian
Iguanacolossus
Iguanodon
"Iguanoides" – nomen nudum; Iguanodon
"Iguanosaurus" – nomen nudum; Iguanodon
Iliosuchus
Ilokelesia
Imperobator
Inawentu
Incisivosaurus
Indosaurus
Indosuchus
"Ingenia" – preoccupied name, now known as Heyuannia yanshini
Ingentia
Inosaurus
Invictarx
Irisosaurus
Irritator
Isaberrysaura
Isanosaurus
Isasicursor
Ischioceratops
Ischisaurus – junior synonym of Herrerasaurus
"Ischyrosaurus" – preoccupied genus name, species Ischyrosaurus manseli
Isisaurus
"Issasaurus" – nomen nudum; Dicraeosaurus
Issi
Itapeuasaurus
Itemirus
Iuticosaurus
Iyuku
J
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Jaculinykus
Jainosaurus
Jakapil
Jaklapallisaurus
Janenschia
Jaxartosaurus
Jeholosaurus
Jenghizkhan – junior synonym of Tarbosaurus
"Jensenosaurus" – nomen nudum; Supersaurus
Jeyawati
Jianchangosaurus
"Jiangjunmiaosaurus" – nomen nudum; Monolophosaurus
Jiangjunosaurus
Jiangshanosaurus
Jiangxisaurus
Jiangxititan
Jianianhualong
Jinbeisaurus
Jinfengopteryx
"Jingia" – preoccupied name, now known as Jingiella
Jingiella
Jingshanosaurus
Jintasaurus
Jinyunpelta
Jinzhousaurus
Jiutaisaurus
Jobaria
Jubbulpuria – possible junior synonym of Laevisuchus
Judiceratops
Jurapteryx – junior synonym of Archaeopteryx
"Jurassosaurus" – nomen nudum; Tianchisaurus
Juratyrant
Juravenator
K
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Kaatedocus
"Kagasaurus" – nomen nudum
Kaijiangosaurus
Kaijutitan
Kakuru
Kamuysaurus
Kangnasaurus
Kansaignathus
Karongasaurus
Katepensaurus
"Katsuyamasaurus" – nomen nudum
Kayentavenator
Kazaklambia
Kelmayisaurus
Kelumapusaura
Kemkemia – subsequently found to be a crocodyliform
Kentrosaurus
Kentrurosaurus – junior synonym (unneeded replacement name) of Kentrosaurus
Kerberosaurus
Khaan
"Khetranisaurus" – nomen nudum
Kholumolumo
Khulsanurus
Kileskus
Kinnareemimus
"Kitadanisaurus" – nomen nudum; Fukuiraptor
"Kittysaurus" – nomen nudum; Eotyrannus
Klamelisaurus
Kol
Koparion
Koreaceratops
Koreanosaurus
"Koreanosaurus" – nomen nudum; name later used formally for a genus of ornithopod
Koshisaurus
Kosmoceratops
Kotasaurus
Koutalisaurus – possible junior synonym of Pararhabdodon
Kritosaurus
Kryptops
Krzyzanowskisaurus – probably a pseudosuchian (Revueltosaurus?)
Kukufeldia – junior synonym of Barilium
Kulceratops
Kulindadromeus
Kulindapteryx – synonym of Kulindadromeus
Kunbarrasaurus
Kundurosaurus
"Kunmingosaurus" – nomen nudum
Kuru
Kurupi
Kuszholia – subsequently found to be a bird
Kwanasaurus – possibly non-dinosaurian
L
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Labocania
Labrosaurus – junior synonym of Allosaurus
"Laelaps" – preoccupied name, now known as Dryptosaurus
Laevisuchus
Lagerpeton – subsequently found to be a non-dinosaurian pterosauromorph
Lagosuchus – subsequently found to be a non-dinosaurian dinosauromorph
Laiyangosaurus
Lajasvenator
Lamaceratops – possible junior synonym of Bagaceratops
Lambeosaurus
Lametasaurus
Lamplughsaura
Lanasaurus – junior synonym of Lycorhinus
"Lancangosaurus" – variant spelling of "Lancanjiangosaurus"
"Lancanjiangosaurus" – nomen nudum
Lanzhousaurus
Laosaurus
Lapampasaurus
Laplatasaurus
Lapparentosaurus
Laquintasaura
Latenivenatrix – possible junior synonym of Stenonychosaurus
Latirhinus
Lavocatisaurus
Leaellynasaura
Ledumahadi
Leinkupal
Leipsanosaurus – junior synonym of Struthiosaurus
"Lengosaurus" – nomen nudum; Eotyrannus
Leonerasaurus
Lepidocheirosaurus — junior synonym of Kulindadromeus
Lepidus
Leptoceratops
Leptorhynchos
Leptospondylus – junior synonym of Massospondylus
Leshansaurus
Lesothosaurus
Lessemsaurus
Levnesovia
Lewisuchus – possibly non-dinosaurian
Lexovisaurus
Leyesaurus
Liaoceratops
Liaoningosaurus
Liaoningotitan
Liaoningvenator
"Liassaurus" – nomen nudum; possible synonym of Sarcosaurus
Libycosaurus – subsequently found to be an anthracothere mammal
Ligabueino
Ligabuesaurus
"Ligomasaurus" – nomen nudum, synonym of Giraffatitan
"Likhoelesaurus" – nomen nudum; possibly non-dinosaurian
Liliensternus
Limaysaurus
"Limnornis" – preoccupied name, now known as Palaeocursornis ( a pterosaur)
"Limnosaurus" – preoccupied name, now known as Telmatosaurus
Limusaurus
Lingwulong
Lingyuanosaurus
Linhenykus
Linheraptor
Linhevenator
Lirainosaurus
Lisboasaurus – subsequently found to be a crocodilian
Liubangosaurus
Llukalkan
Lohuecotitan
Loncosaurus
Longisquama – subsequently found to be a non-dinosaurian reptile
Longosaurus – junior synonym of Coelophysis
Lophorhothon
Lophostropheus
Loricatosaurus
Loricosaurus
Losillasaurus
Lourinhanosaurus
Lourinhasaurus
Luanchuanraptor
"Luanpingosaurus" – nomen nudum; Psittacosaurus
Lucianosaurus – subsequently found to be a non-dinosaurian archosauriform
Lucianovenator
Lufengosaurus
Lukousaurus – possibly a crurotarsan
Luoyanggia
Lurdusaurus
Lusitanosaurus
Lusotitan
Lusovenator
Lutungutali – possibly non-dinosaurian
Lycorhinus
Lythronax
M
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Macelognathus – subsequently found to be a sphenosuchian crocodilian
Machairasaurus
Machairoceratops
Macrocollum
Macrodontophion – subsequently found to be a member of Lophotrochozoa
Macrogryphosaurus
Macrophalangia – junior synonym of Chirostenotes
"Macroscelosaurus" – nomen nudum; junior synonym of Tanystropheus
Macrurosaurus
"Madsenius" – nomen nudum, Allosaurus
Magnamanus
Magnapaulia
Magnirostris – possible junior synonym of Bagaceratops
Magnosaurus
"Magulodon" – nomen nudum
Magyarosaurus
Mahakala
Mahuidacursor
Maiasaura
Maip
Majungasaurus
Majungatholus – junior synonym of Majungasaurus
Malarguesaurus
Malawisaurus
Maleevosaurus – junior synonym of Tarbosaurus
Maleevus
Malefica
Mamenchisaurus
Mandschurosaurus
Manidens
Manospondylus – synonym of Tyrannosaurus
Mansourasaurus
Mantellisaurus
Mantellodon – junior synonym of Mantellisaurus
"Maojandino" – nomen nudum
Mapusaurus
Maraapunisaurus
Marasuchus – subsequently found to be a non-dinosaurian dinosauromorph
"Marisaurus" – nomen nudum
Marmarospondylus
Marshosaurus
Martharaptor
Masiakasaurus
Massospondylus
Matheronodon
Maxakalisaurus
Mbiresaurus
Medusaceratops
"Megacervixosaurus" – nomen nudum
"Megadactylus" – preoccupied name, now known as Anchisaurus
"Megadontosaurus" – nomen nudum; Microvenator
Megalosaurus
Megapnosaurus – possible junior synonym of Coelophysis
Megaraptor
Mei
Melanorosaurus
Mendozasaurus
Menefeeceratops
Menucocelsior
Meraxes
Mercuriceratops
Meroktenos
"Merosaurus" – nomen nudum; Dornraptor
Metriacanthosaurus
"Microcephale" – nomen nudum
"Microceratops" – preoccupied name, now known as Microceratus
Microceratus
Microcoelus
"Microdontosaurus" – nomen nudum
Microhadrosaurus
Micropachycephalosaurus
Microraptor
Microvenator
Mierasaurus
"Mifunesaurus" – nomen nudum
Migmanychion
Minimocursor
Minmi
Minotaurasaurus
Minqaria
Miragaia
Mirischia
Mnyamawamtuka
Moabosaurus
Mochlodon
"Mohammadisaurus" – nomen nudum; Tornieria
Mojoceratops – junior synonym of Chasmosaurus
Mongolosaurus
Mongolostegus
Monkonosaurus
Monoclonius
Monolophosaurus
"Mononychus" – preoccupied name, now known as Mononykus
Mononykus
Montanoceratops
Morelladon
Morinosaurus
Moros
Morosaurus – junior synonym of Camarasaurus
Morrosaurus
Mosaiceratops
"Moshisaurus" – nomen nudum; possibly Mamenchisaurus
"Mtapaiasaurus" – nomen nudum, synonym of Giraffatitan
"Mtotosaurus" – nomen nudum; Dicraeosaurus
Murusraptor
Mussaurus
Muttaburrasaurus
Muyelensaurus
Mymoorapelta
N
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Naashoibitosaurus
Nambalia
Nankangia
Nanningosaurus
Nanosaurus
Nanotyrannus – possible junior synonym of Tyrannosaurus
Nanshiungosaurus
Nanuqsaurus
Nanyangosaurus
Napaisaurus
Narambuenatitan
Narindasaurus
Nasutoceratops
Natovenator
"Natronasaurus" – invalid name, either Alcovasaurus or Miragaia
Navajoceratops
Nebulasaurus
"Nectosaurus" – preoccupied name, now known as Kritosaurus
Nedcolbertia
Nedoceratops – possible junior synonym of Triceratops
Neimongosaurus
"Nemegtia" – preoccupied name, now known as Nemegtomaia
Nemegtomaia
Nemegtonykus
Nemegtosaurus
"Neosaurus" – preoccupied name; renamed Parrosaurus, which is now Hypsibema
Neosodon
Neovenator
Neuquenraptor
Neuquensaurus
Nevadadromeus
"Newtonsaurus" – nomen nudum, possibly Zanclodon
"Ngexisaurus" – nomen nudum
Ngwevu
Nhandumirim
Nicksaurus – nomen manuscriptum
Niebla
Nigersaurus
Ningyuansaurus
Ninjatitan
Niobrarasaurus
Nipponosaurus
Noasaurus
Nodocephalosaurus
Nodosaurus
Nomingia – possible junior synonym of Elmisaurus
Nopcsaspondylus
Normanniasaurus
Notatesseraeraptor
Nothronychus
Notoceratops
Notocolossus
Notohypsilophodon
Nqwebasaurus
"Nteregosaurus" – nomen nudum; Janenschia
Nullotitan
"Nurosaurus" – nomen nudum
Nuthetes
Nyasasaurus — possibly non-dinosaurian
"Nyororosaurus" – nomen nudum; Dicraeosaurus
O
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Oblitosaurus
Oceanotitan
Ohmdenosaurus
Ojoceratops – possible synonym of Eotriceratops
Ojoraptorsaurus
Oksoko
Oligosaurus – possible synonym of Mochlodon
Olorotitan
Omeisaurus
"Omosaurus" – preoccupied name, now known as Dacentrurus
Ondogurvel
Onychosaurus – junior synonym of Zalmoxes or Rhabdodon, or an ankylosaurian
Oohkotokia
Opisthocoelicaudia
Oplosaurus
"Orcomimus" – nomen nudum
Orinosaurus – junior synonym (unneeded replacement name) of Orosaurus
Orkoraptor
Ornatops
Ornatotholus – junior synonym of Stegoceras
Ornithodesmus
"Ornithoides" – nomen nudum; Saurornithoides
Ornitholestes
Ornithomerus – possible synonym of Mochlodon
Ornithomimoides
Ornithomimus
Ornithopsis
Ornithosuchus – subsequently found to be a non-dinosaurian archosaur
Ornithotarsus – junior synonym of Hadrosaurus
Orodromeus
Orosaurus
Orthogoniosaurus
Orthomerus
Oryctodromeus
"Oshanosaurus" – nomen nudum
Osmakasaurus
Ostafrikasaurus
Ostromia
Othnielia – junior synonym of Nanosaurus
Othnielosaurus – junior synonym of Nanosaurus
Otogosaurus — possibly a nomen nudum
Ouranosaurus
Overoraptor
Overosaurus
Oviraptor
"Ovoraptor" – nomen nudum; Velociraptor
Owenodon
Oxalaia – possible junior synonym of Spinosaurus
Ozraptor
P
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Pachycephalosaurus
Pachyrhinosaurus
Pachysauriscus – junior synonym of Plateosaurus
Pachysaurops – junior synonym of Plateosaurus
"Pachysaurus" – preoccupied name, now known as Pachysauriscus; junior synonym of Plateosaurus
Pachyspondylus – junior synonym of Massospondylus
Pachysuchus
Padillasaurus
"Pakisaurus" – nomen nudum
Palaeoctonus – subsequently found to be a phytosaur
Palaeocursornis – subsequently found to be an azhdarchoid pterosaur
"Palaeolimnornis" – nomen nudum; Palaeocursornis, pterodactyloid pterosaur belonging to Azhdarchoidea
Palaeopteryx – possibly a bird
Palaeosauriscus – junior synonym of Palaeosaurus
Palaeosaurus – subsequently found to be a non-dinosaurian reptile
"Palaeosaurus" – preoccupied name, now known as Sphenosaurus and considered to be a non-dinosaurian procolophonid
Palaeoscincus
Paleosaurus – subsequently found to be a non-dinosaurian archosaur; junior synonym (unneeded replacement name) of Palaeosaurus
Paludititan
Paluxysaurus – junior synonym of Sauroposeidon
Pampadromaeus
Pamparaptor
Panamericansaurus
Pandoravenator
Panguraptor
Panoplosaurus
Panphagia
Pantydraco – possible synonym of Thecodontosaurus
Papiliovenator
"Paraiguanodon" – nomen nudum; Bactrosaurus
Paralitherizinosaurus
Paralititan
Paranthodon
Pararhabdodon
Parasaurolophus
Paraxenisaurus
Pareiasaurus – subsequently found to be a pareiasaur
Pareisactus
Parksosaurus
Paronychodon
Parrosaurus – now known as Hypsibema missouriensis
Parvicursor
Patagonykus
Patagopelta
Patagosaurus
Patagotitan
Pawpawsaurus
Pectinodon
Pedopenna
Pegomastax
Peishansaurus
Pekinosaurus – subsequently found to be a pseudosuchian; junior synonym of Revueltosaurus
Pelecanimimus
Pellegrinisaurus
Peloroplites
Pelorosaurus
"Peltosaurus" – preoccupied name, now known as Sauropelta
Pendraig
Penelopognathus
Pentaceratops
Perijasaurus
Petrobrasaurus
Phaedrolosaurus
Philovenator
Phuwiangosaurus
Phuwiangvenator
Phyllodon
Piatnitzkysaurus
Picrodon – possibly non-dinosaurian
Pilmatueia
Pinacosaurus
Pisanosaurus — possibly non-dinosaurian
Pitekunsaurus
Piveteausaurus
Planicoxa
Plateosauravus
Plateosaurus
Platyceratops – possible junior synonym of Bagaceratops
Platypelta
Platytholus
Plesiohadros
Pleurocoelus – possible junior synonym of Astrodon
Pleuropeltus – junior synonym of Struthiosaurus
Pneumatoarthrus – subsequently found to be a turtle
Pneumatoraptor
Podokesaurus
Poekilopleuron
Polacanthoides – chimera of Hylaeosaurus and Polacanthus
Polacanthus
Polyodontosaurus
Polyonax
Ponerosteus – subsequently found to be a non-dinosaurian archosaur
Poposaurus – subsequently found to be a non-dinosaurian archosaur
Portellsaurus
Postosuchus – subsequently found to be a rauisuchian
Powellvenator
Pradhania
Prenocephale
Prenoceratops
Priconodon
Priodontognathus
Proa
Probactrosaurus
Probrachylophosaurus
Proceratops – junior synonym (unneeded replacement name) of Ceratops
Proceratosaurus
Procerosaurus – subsequently found to be a tanystropheid protorosaur, Tanystropheus
"Procerosaurus" – preoccupied name, now known as Ponerosteus
Procheneosaurus – junior synonym of Lambeosaurus
Procompsognathus
Prodeinodon
"Proiguanodon" – nomen nudum; Iguanodon
Propanoplosaurus
Proplanicoxa – junior synonym of Mantellisaurus
Prosaurolophus
Protarchaeopteryx
Protathlitis
Protecovasaurus – subsequently found to be a non-dinosaurian archosauriform
Protiguanodon – junior synonym of Psittacosaurus
Protoavis – described as a bird, probably a chimera including theropod dinosaur bones
Protoceratops
Protognathosaurus
"Protognathus" – preoccupied name, now known as Protognathosaurus
Protohadros
"Protorosaurus" – preoccupied name, now known as Chasmosaurus
Protorosaurus – subsequently found to be a non-dinosaurian reptile
"Proyandusaurus" – nomen nudum; Hexinlusaurus.
Pseudolagosuchus – possibly non-dinosaurian; a junior synonym of Lewisuchus
Psittacosaurus
Pteropelyx
Pterospondylus
Puertasaurus
Pukyongosaurus
Pulanesaura
Punatitan
Pycnonemosaurus
Pyroraptor
Q
Contents: A B C D E F G H I J K L M N O P Q R S T U V W X Y Z – See also
Qantassaurus
Qianlong
Qianzhousaurus
Qiaowanlong
Qijianglong
Qingxiusaurus
Qinlingosaurus
Qiupalong
Qiupanykus
Quaesitosaurus
Quetecsaurus
Quilmesaurus
R
Contents: A B C D E F G H I J K L M N O P Q R S T U V W X Y Z – See also
Rachitrema – subsequently found to be a chimera primarily based on ichthyosaur fossils
Rahiolisaurus
"Rahona" – preoccupied name, now known as Rahonavis
Rahonavis – possibly a bird
Rajasaurus
Rapator
Rapetosaurus
Raptorex – possible junior synonym of Tarbosaurus
Ratchasimasaurus
Rativates
Rayososaurus
Razanandrongobe – subsequently found to be a crocodylomorph
Rebbachisaurus
Regaliceratops
Regnosaurus
Revueltosaurus – subsequently found to be a pseudosuchian
Rhabdodon
Rhadinosaurus – may be non-dinosaurian, possibly crocodilian
Rhinorex – possible synonym of Gryposaurus
Rhodanosaurus – junior synonym of Struthiosaurus
Rhoetosaurus
Rhomaleopakhus
Rhopalodon – subsequently found to be a synapsid
Riabininohadros
Richardoestesia
"Rileya" – preoccupied name, now known as Rileyasuchus
Rileyasuchus – subsequently found to be a phytosaur
Rinchenia
Rinconsaurus
Rioarribasaurus – junior synonym of Coelophysis
"Riodevasaurus" – nomen nudum; Turiasaurus
Riojasaurus
Riojasuchus – subsequently found to be a non-dinosaurian archosaur
Riojavenatrix
Riparovenator
Rocasaurus
"Roccosaurus" – nomen nudum; Melanorosaurus
"Ronaldoraptor" – nomen nudum
Rubeosaurus – junior synonym of Styracosaurus
Ruehleia
Rugocaudia
Rugops
Ruixinia
Rukwatitan
Ruyangosaurus
S
Contents: A B C D E F G H I J K L M N O P Q R S T U V W X Y Z – See also
Sacisaurus – possibly non-dinosaurian
Sahaliyania
Saichania
"Saldamosaurus" – nomen nudum
"Salimosaurus" – nomen nudum, synonym of Giraffatitan
Saltasaurus
Saltopus – possibly non-dinosaurian
"Saltriosaurus" – nomen nudum
Saltriovenator
"Sanchusaurus" – nomen nudum, possible Gallimimus
"Sangonghesaurus" – nomen nudum, Tianchisaurus
Sanjuansaurus
Sanpasaurus
Santanaraptor
Sanxiasaurus
Sarahsaurus
Saraikimasoom – nomen manuscriptum
Sarcolestes
Sarcosaurus
Sarmientosaurus
Saturnalia
"Sauraechinodon" – nomen nudum; Echinodon
"Sauraechmodon" – nomen nudum; Echinodon
"Saurechinodon" – nomen nudum; Echinodon
Saurolophus
Sauroniops
Sauropelta
Saurophaganax
"Saurophagus" – preoccupied name, now known as Saurophaganax
Sauroplites
Sauroposeidon
Saurornithoides
Saurornitholestes
Savannasaurus
Scansoriopteryx
Scaphonyx – subsequently found to be a rhynchosaur, Hyperodapedon
Scelidosaurus
Schleitheimia
Scipionyx
Sciurumimus
Scleromochlus – subsequently found to be a non-dinosaurian avemetatarsalian
Scolosaurus
Scutellosaurus
Secernosaurus
Sefapanosaurus
Segisaurus
Segnosaurus
Seismosaurus – junior synonym of Diplodocus
Seitaad
Sektensaurus
"Selimanosaurus" – nomen nudum; Dicraeosaurus
Sellacoxa – junior synonym of Barilium
Sellosaurus – junior synonym of Plateosaurus
Serendipaceratops
Serikornis
Shamosaurus
Shanag
Shanshanosaurus – junior synonym of Tarbosaurus
Shantungosaurus
Shanxia
Shanyangosaurus
Shaochilong
Shenzhousaurus
Shidaisaurus
Shingopana
Shishugounykus
Shixinggia
Shri
Shuangbaisaurus – possible synonym of Sinosaurus
Shuangmiaosaurus
Shunosaurus
Shuvosaurus – subsequently found to be a rauisuchian
Shuvuuia
Siamodon
"Siamodracon" – nomen nudum
Siamosaurus
Siamotyrannus
Siamraptor
Siats
"Sibirosaurus" – nomen nudum, now known as Sibirotitan
Sibirotitan
Sidersaura
"Sidormimus" – nomen nudum
Sierraceratops
Sigilmassasaurus – possible junior synonym of Spinosaurus
Silesaurus – possibly non-dinosaurian
Siluosaurus
Silutitan
Silvisaurus
Similicaudipteryx
Sinankylosaurus
Sinocalliopteryx
Sinocephale
Sinoceratops
Sinocoelurus
"Sinopelta" – nomen nudum; synonym of Sinopeltosaurus
"Sinopeltosaurus" – nomen nudum
Sinopliosaurus – a pliosaur; one species, "S." fusuiensis, is actually a dinosaur that may be synonymous with Siamosaurus
Sinornithoides
Sinornithomimus
Sinornithosaurus
Sinosauropteryx
Sinosaurus
Sinotyrannus
Sinovenator
Sinraptor
Sinusonasus
Sirindhorna
Skorpiovenator
"Smilodon" – preoccupied name, now known as Zanclodon
Smitanosaurus
Smok — possibly non-dinosaurian
Sonidosaurus
Sonorasaurus
Soriatitan
Soumyasaurus – possibly non-dinosaurian
Spectrovenator
Sphaerotholus
Sphenosaurus – subsequently found to be a non-dinosaurian reptile
Sphenospondylus – junior synonym of Mantellisaurus
Spiclypeus
Spicomellus
Spinophorosaurus
Spinops
Spinosaurus
Spinostropheus
Spinosuchus – subsequently found to be a non-dinosaurian reptile
Spondylosoma – subsequently found to be an aphanosaur
Squalodon – subsequently found to be a cetacean
Staurikosaurus
Stegoceras
Stegopelta
Stegosaurides
Stegosaurus
Stegouros
Stellasaurus
Stenonychosaurus
Stenopelix
Stenotholus – junior synonym of Stygimoloch, which is a possible junior synonym of Pachycephalosaurus
Stephanosaurus
"Stereocephalus" – preoccupied name, now known as Euoplocephalus
Sterrholophus – junior synonym of Triceratops
Stokesosaurus
Stormbergia – junior synonym of Lesothosaurus
Strenusaurus – junior synonym of Riojasaurus
Streptospondylus
Struthiomimus
Struthiosaurus
Stygimoloch – junior synonym of Pachycephalosaurus
Stygivenator – junior synonym of Tyrannosaurus
Styracosaurus
Succinodon – subsequently found to be fossilized mollusc borings
Suchomimus
Suchoprion – subsequently found to be a phytosaur
Suchosaurus – possible synonym of Baryonyx
"Sugiyamasaurus" – nomen nudum
"Sulaimanisaurus" – nomen nudum
Supersaurus
Suskityrannus
Suuwassea
Suzhousaurus
Symphyrophus – junior synonym of Camptosaurus
Syngonosaurus
"Syntarsus" – preoccupied name, sometimes assigned to Coelophysis or Megapnosaurus
Syrmosaurus – junior synonym of Pinacosaurus
Szechuanosaurus
T
Contents: A B C D E F G H I J K L M N O P Q R S T U V W X Y Z – See also
Tachiraptor
Talarurus
Talenkauen
Talos
Tamarro
Tambatitanis
Tangvayosaurus
Tanius
Tanycolagreus
Tanystropheus – subsequently found to be a protorosaur
Tanystrosuchus
Taohelong
Tapinocephalus – subsequently found to be a therapsid
Tapuiasaurus
Tarascosaurus
Tarbosaurus
Tarchia
Tastavinsaurus
Tatankacephalus
Tatankaceratops – probable junior synonym of Triceratops
Tataouinea
Tatisaurus
Taurovenator
Taveirosaurus
Tawa
Tawasaurus – junior synonym of Lufengosaurus
Tazoudasaurus
Technosaurus – possibly non-dinosaurian
Tecovasaurus – subsequently found to be a non-dinosaurian archosauriform
Tehuelchesaurus
"Teihivenator" – nomen nudum
Teinurosaurus
Teleocrater – subsequently found to be a basal avemetatarsalian
Telmatosaurus
"Tenantosaurus" – nomen nudum; Tenontosaurus
"Tenchisaurus" – nomen nudum; an unpublished museum name for Tianchisaurus
Tendaguria
Tengrisaurus
Tenontosaurus
Teratophoneus
Teratosaurus – subsequently found to be a non-dinosaurian archosaur
Termatosaurus – subsequently found to be a phytosaur
Terminocavus
Tethyshadros
Tetragonosaurus – junior synonym of Lambeosaurus
Texacephale
Texasetes
Teyuwasu – possibly junior synonym of Staurikosaurus
Thanatotheristes
Thanos
Tharosaurus
Thecocoelurus
Thecodontosaurus
Thecospondylus
Theiophytalia
Therizinosaurus
Therosaurus – a synonym of Iguanodon
Thescelosaurus
Thespesius
"Thotobolosaurus" – nomen nudum; Kholumolumo
Thyreosaurus
Tianchiasaurus – alternate spelling of Tianchisaurus
Tianchisaurus
"Tianchungosaurus" – nomen nudum; Dianchungosaurus (crocodilian)
Tianyulong
Tianyuraptor
Tianzhenosaurus
Tichosteus
Tienshanosaurus
Tietasaura
Timimus
Timurlengia
Titanoceratops
Titanomachya
Titanosaurus
"Titanosaurus" – preoccupied name, now known as Atlantosaurus
Tlatolophus
Tochisaurus
"Tomodon" – preoccupied name, now known as Diplotomodon
Tonganosaurus
Tongtianlong
"Tonouchisaurus" – nomen nudum
Torilion – junior synonym of Barilium
Tornieria
Torosaurus
Torvosaurus
Tototlmimus
Trachelosaurus – subsequently found to be a basal archosauromorph
Trachodon
Tralkasaurus
Transylvanosaurus
Tratayenia
Traukutitan
Trialestes – subsequently found to be a basal crocodylomorph
"Triassolestes" – preoccupied name, now known as Trialestes
Tribelesodon – junior synonym of Tanystropheus, a protorosaur
Triceratops
Trierarchuncus
Trigonosaurus
Trimucrodon
Trinisaura
Triunfosaurus
Troodon
Tsaagan
Tsagantegia
Tsintaosaurus
Tuebingosaurus
Tugulusaurus
Tuojiangosaurus
Turanoceratops
Turiasaurus
Tylocephale
Tylosteus – synonym of Pachycephalosaurus
Tyrannomimus
Tyrannosaurus
Tyrannotitan
U
Contents: A B C D E F G H I J K L M N O P Q R S T U V W X Y Z – See also
Uberabatitan
"Ubirajara" – nomen nudum
Udanoceratops
Udelartitan
Ugrosaurus – junior synonym of Triceratops
Ugrunaaluk – junior synonym of Edmontosaurus
Uintasaurus – junior synonym of Camarasaurus
Ultrasauros – junior synonym of Supersaurus
"Ultrasaurus" – preoccupied name, renamed Ultrasauros which is now a junior synonym of Supersaurus
Ultrasaurus
Ulughbegsaurus
"Umarsaurus" – nomen nudum; Barsboldia
Unaysaurus
Unenlagia
Unescoceratops
"Unicerosaurus" – nomen nudum, subsequently found to be a fish
Unquillosaurus
Urbacodon
Utahceratops
Utahraptor
Uteodon
V
Contents: A B C D E F G H I J K L M N O P Q R S T U V W X Y Z – See also
Vagaceratops
Vahiny
Valdoraptor – possible synonym of Thecocoelurus
Valdosaurus
Vallibonavenatrix
Variraptor
Vayuraptor
Vectaerovenator
Vectensia – junior synonym of Polacanthus or Hylaeosaurus
Vectidromeus
Vectipelta
Vectiraptor
Vectisaurus – junior synonym of Mantellisaurus
Velafrons
Velocipes
Velociraptor
Velocisaurus
Venaticosuchus – subsequently found to be a non-dinosaurian archosaur
Venenosaurus
Vespersaurus
Veterupristisaurus
Viavenator
"Vitakridrinda" – nomen nudum
"Vitakrisaurus" – nomen nudum
Volgatitan
Volkheimeria
Vouivria
Vulcanodon
W
Contents: A B C D E F G H I J K L M N O P Q R S T U V W X Y Z – See also
Wadhurstia – junior synonym of Hypselospinus
Wakinosaurus
Walgettosuchus – possible synonym of Rapator
"Walkeria" – preoccupied name, now known as Alwalkeria
"Walkersaurus" – nomen nudum; Duriavenator
Wamweracaudia
"Wangonisaurus" – nomen nudum, synonym of Giraffatitan
Wannanosaurus
Weewarrasaurus
Wellnhoferia – subsequently found to be a bird, possible junior synonym of Archaeopteryx
Wendiceratops
Wiehenvenator
Willinakaqe
Wintonotitan
Wuerhosaurus
Wulagasaurus
Wulatelong
Wulong
Wyleyia – subsequently found to be a bird
"Wyomingraptor" – nomen nudum, synonym of Allosaurus
X
Contents: A B C D E F G H I J K L M N O P Q R S T U V W X Y Z – See also
Xenoceratops
Xenoposeidon
Xenotarsosaurus
Xianshanosaurus
Xiaosaurus
Xiaotingia
Xingtianosaurus
Xingxiulong
Xinjiangovenator
Xinjiangtitan
Xiongguanlong
Xixianykus
Xixiasaurus
Xixiposaurus
Xiyunykus
Xuanhanosaurus
Xuanhuaceratops
"Xuanhuasaurus" – nomen nudum; Xuanhuaceratops
Xunmenglong
Xuwulong
Y
Contents: A B C D E F G H I J K L M N O P Q R S T U V W X Y Z – See also
Yaleosaurus – junior synonym of Anchisaurus
Yamaceratops
Yamanasaurus
Yamatosaurus
Yanbeilong
Yandusaurus
Yangchuanosaurus
Yaverlandia
Yehuecauhceratops
"Yezosaurus" – nomen nudum; subsequently found to be a junior synonym of the mosasaur Taniwhasaurus
Yi
"Yibinosaurus" – nomen nudum
Yimenosaurus
Yingshanosaurus
Yinlong
Yixianosaurus
Yizhousaurus
Yongjinglong
Ypupiara
"Yuanmouraptor" – nomen nudum
Yuanmousaurus
Yueosaurus
Yulong
Yunganglong
Yunmenglong
Yunnanosaurus
"Yunxianosaurus" – nomen nudum
Yunyangosaurus
Yurgovuchia
Yutyrannus
Yuxisaurus
Yuzhoulong
Z
Contents: A B C D E F G H I J K L M N O P Q R S T U V W X Y Z – See also
Zalmoxes
Zanabazar
Zanclodon – subsequently found to be non-dinosaurian
Zapalasaurus
Zapsalis
Zaraapelta
Zatomus – subsequently found to be a non-dinosaurian archosaur
Zby
Zephyrosaurus
Zhanghenglong
Zhejiangosaurus
Zhenyuanlong
Zhongjianosaurus
Zhongornis – subsequently found to be a bird
Zhongyuansaurus – possible junior synonym of Gobisaurus
Zhuchengceratops
Zhuchengosaurus – junior synonym of Shantungosaurus
Zhuchengtitan
Zhuchengtyrannus
Ziapelta
Zigongosaurus
Zizhongosaurus
Zuniceratops
"Zunityrannus" – nomen nudum, Suskityrannus
Zuolong
Zuoyunlong
Zupaysaurus
Zuul
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The Basal Nodosaurid Ankylosaur Europelta carbonensis n. gen., n. sp. from the Lower Cretaceous (Lower Albian) Escucha Formation of Northeastern Spain
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[
"James I. Kirkland",
"Luis Alcalá",
"Mark A. Loewen",
"Eduardo Espílez",
"Luis Mampel",
"Jelle P. Wiersma"
] |
2013-08-10T00:00:00
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Nodosaurids are poorly known from the Lower Cretaceous of Europe. Two associated ankylosaur skeletons excavated from the lower Albian carbonaceous member of the Escucha Formation near Ariño in northeastern Teruel, Spain reveal nearly all the diagnostic ...
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https://www.ncbi.nlm.nih.gov/coreutils/nwds/img/favicons/favicon.ico
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PubMed Central (PMC)
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https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3847141/
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PLoS One. 2013; 8(12): e80405.
PMCID: PMC3847141
PMID: 24312471
The Basal Nodosaurid Ankylosaur Europelta carbonensis n. gen., n. sp. from the Lower Cretaceous (Lower Albian) Escucha Formation of Northeastern Spain
, 1 , * , 2 , 3 , 4 , 2 , 2 and 3 , 4
James I. Kirkland
1 Utah Geological Survey, Salt Lake City, Utah, United States of America
Find articles by James I. Kirkland
Luis Alcalá
2 Fundación Conjunto Paleontológico de Teruel-Dinópolis, Museo Aragonés de Paleontología, Teruel, Spain
Find articles by Luis Alcalá
Mark A. Loewen
3 Natural History Museum of Utah, Salt Lake City, Utah, United States of America
4 Department of Geology & Geophysics, University of Utah, Salt Lake City, Utah, United States of America
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Eduardo Espílez
2 Fundación Conjunto Paleontológico de Teruel-Dinópolis, Museo Aragonés de Paleontología, Teruel, Spain
Find articles by Eduardo Espílez
Luis Mampel
2 Fundación Conjunto Paleontológico de Teruel-Dinópolis, Museo Aragonés de Paleontología, Teruel, Spain
Find articles by Luis Mampel
Jelle P. Wiersma
3 Natural History Museum of Utah, Salt Lake City, Utah, United States of America
4 Department of Geology & Geophysics, University of Utah, Salt Lake City, Utah, United States of America
Find articles by Jelle P. Wiersma
Richard J. Butler, Editor
1 Utah Geological Survey, Salt Lake City, Utah, United States of America
2 Fundación Conjunto Paleontológico de Teruel-Dinópolis, Museo Aragonés de Paleontología, Teruel, Spain
3 Natural History Museum of Utah, Salt Lake City, Utah, United States of America
4 Department of Geology & Geophysics, University of Utah, Salt Lake City, Utah, United States of America
University of Birmingham, United Kingdom
Competing Interests: The authors have declared that no competing interests exist.
Conceived and designed the experiments: JIK LA MAL. Performed the experiments: JIK MAL. Analyzed the data: JIK LA MAL. Wrote the paper: JIK LA MAL. Oversaw the entire Ariño project: LA. Co-directed the excavation: EE LM. Oversaw the preparation of all the fossil materials: EE. Assisted with character evaluations and in constructing many of the figures: JPW. Coordinated quarry mapping and photography: LM EE.
Copyright © 2013 Kirkland et al
This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are properly credited.
Abstract
Nodosaurids are poorly known from the Lower Cretaceous of Europe. Two associated ankylosaur skeletons excavated from the lower Albian carbonaceous member of the Escucha Formation near Ariño in northeastern Teruel, Spain reveal nearly all the diagnostic recognized character that define nodosaurid ankylosaurs. These new specimens comprise a new genus and species of nodosaurid ankylosaur and represent the single most complete taxon of ankylosaur from the Cretaceous of Europe. These two specimens were examined and compared to all other known ankylosaurs. Comparisons of these specimens document that Europelta carbonensis n. gen., n. sp. is a nodosaur and is the sister taxon to the Late Cretaceous nodosaurids Anoplosaurus, Hungarosaurus, and Struthiosaurus, defining a monophyletic clade of European nodosaurids– the Struthiosaurinae.
Introduction
Ankylosaurs were first described from the Lower Cretaceous of England with Hylaeosaurus armatus (Valanginian) described in 1833 [1]–[3]. Hylaeosaurus is one of the three dinosaurs on which the Dinosauria were defined [4] and one of the first dinosaurs for which a full-sized life reconstruction was attempted at the Crystal Palace Park in London in 1854 [5]. Although first mentioned in an anonymous article in the September 16th 1865 issue of the “The Illustrated London News” by Sir Richard Owen [6], the Early Cretaceous (Barremian) Polacanthus was not described formally as Polacanthus foxii by Hulke until 1882 [7]–[10]. The abundant plates and spines of these ankylosaurs are characteristic of the Lower Cretaceous up into the lower part of the Aptian stage [11], [12]. In 1867, Huxley described the fragmentary Acanthopholis from the base of the Upper Cretaceous (Cenomanian) [13]–[15]. Additionally, in 1879, Seeley [16] described the juvenile nodosaurid Anoplosaurus curtonotus [17] from the uppermost Lower Cretaceous (upper Albian) Cambridge Greensand. Subsequent descriptions of the fragmentary remains of ankylosaurs from the Early Cretaceous of Europe have been tentatively assigned to the genus Polacanthus [18].
Only nodosaurids have been described from the Upper Cretaceous of Europe with Struthiosaurus austriacus described from the Campanian of Austria in 1871 [19]–[24] followed by Struthiosaurus transylvanicus [25], [26], [27] from the uppermost Cretaceous (upper Maastrichtian) strata of Romania. Until recently, all Late Cretaceous ankylosaur fossils in Europe have been assigned to Struthiosaurus [28]–[30] including Struthiosaurus languedocensis from the Campanian of southern France [31]. The primitive nodosaurid Hungarosaurus tormai [32], [33] from the mid-Late Cretaceous (Santonian) is now known from multiple specimens and has become the best documented ankylosaur in Europe.
Fragmentary ankylosaur remains are also known from a number of localities from the Middle to Upper Jurassic strata of Europe, but have been relatively uninformative as specimens are based largely on isolated skeletal elements [34].
Northeastern Spain has contributed many dinosaur discoveries from both Lower and Upper Cretaceous strata in recent years [35]. The Early Cretaceous dinosaurs discovered to date include numerous sauropods, iguanodonts, and ankylosaurs from the Barremian-lower Aptian, with all the fragmentary ankylosaur material assigned tentatively to the genus Polacanthus [25], [28], [36]–[40]. All the Late Cretaceous ankylosaurs from Spain have in turn been assigned to Struthiosaurus [28]–[30].
The earliest reported dinosaur remains from Spain were found in the Escucha Formation, few significant vertebrate fossils had been recovered from these rocks in the 140 intervening years [41], [42]. Current research on vertebrate sites in the Escucha Formation in the northern Teruel Province in the Community of Aragón, Spain, by the Fundación Conjunto Paleontológico of Teruel-Dinópolis has resulted in the discovery of an extensive new dinosaur locality in the open-pit Santa María coal mine near Ariño ( ) operated by Sociedad Anónima Minera Catalano-Aragonesa (SAMCA Group) [42]. The most abundant dinosaur identified is a distinctive iguanodontian ornithopod recently described as Proa valdearinnoensis [43]. Among the many other significant fossils excavated are two associated partial skeletons of a new species of ankylosaur, described herein as Europelta carbonensis n. gen., n. sp. This new taxon is the most completely known ankylosaur in Europe and adds considerable new information about Early Cretaceous ankylosaurian phylogeny and biogeography.
Geological Setting
Counterclockwise rotation of the Iberian Plate toward the end of the Early Cretaceous resulted in the development of a series of syndepositional sub-basins bounded by active faults within Ebro Basin south of the Pyrenean ranges, northeast of the Iberian Range, and northwest of the Catalan/Coastal Range [44], [45]. The new dinosaur locality is within the Oliete sub-basin on the northwest margin of the Escucha outcrop belt [42], [44]. The Formación Lignitos de Escucha and overlying Formación Arenas de Utrillas were initially described in 1971 [46]. These largely Albian-aged strata were deposited along the northwestern margin of the Tethys Sea during the fragmentation of this terrain, and overlie Aptian strata in the center of each sub-basin and unconformably overlie progressively older strata toward their margins. Initially, the Escucha Formation was divided into three members [47] and interpreted to be an unconformity-bounded lower to middle Albian depositional sequence, representing a progradational, tidally-dominated delta sequence [44], [48]–[52]. Recently, the upper “fluvial” member has been reinterpreted as an eolian depositional sequence separated from the underlying portions of the Escucha Formation by a regional unconformity [53]. We recognize this bipartite division of the Escucha Formation ( ).
The geologic age of the Escucha Formation has been considered to be early to middle Albian. It overlies Aptian strata in central basinal settings and is, in turn, overlain by the upper Albian Utrillas Formation [44]. However, both calcareous plankton (foraminifera and nanoplankton) [54] and palynomorphs [55], [56] indicate that the lower Escucha Formation is late Aptian in age. Both fresh and brackish coal-bearing strata are recognized below the regional unconformity within the Escucha [43]. However, reports on the microplankton restrict marine and marginal marine facies to the late Aptian in the lower Escucha Formation [54]–[56]. Marine ostracods have been reported from the upper Escucha Formation northeast of Teruel that confirm an Albian age for the upper portion of these strata in this area [57].
A sample of the matrix from the bonebed was processed for both palynomorphs and calcareous microfossils. The palynomorphs were exclusively of terrestrial origin and indicated an Albian age (Gerry Waanders, 2012, personal communication). The microfossils consisted exclusively of freshwater ostracods and charophytes. The ostracods represent new species and the charophytes are also reported from the Albian of Tunisia [58]. No arenaceous foraminifera were identified, which, along with the absence of dinoflagelates, indicates that the bonebed formed well inland of marine and brackish water influences ( ).
The bonebed is located immediately below the lowest mineable coal seam in the Santa María coal mine ( ), in a dark olive-gray to olive-black mudstone that preserves a high percentage of fossil plant debris. In overall appearance, the rock is much like the plant debris beds in the Wessex Formation on the Isle of Wight [59], [60] and, as in those beds, there is a great amount of pyrite (iron sulfide) disseminated through the matrix and in the fossils. Significant amounts of iron sulfide in the coals were found to decrease up section, away from marine and brackish-water environments. In addition to this depositional relationship, it has been speculated that detrital evaporites from exposed Triassic strata on the north and northwest sides of the basin have secondarily contributed significant amounts of sulfur to these coals [43], [61]. Additionally, the abundance of pyrite in the bones indicates that the long-term stability of the fossils is in question as pyrite breaks down in an expansive oxidation reaction that liberates corrosive sulfuric acid compounds that cannot be reversed [62]. The degradation by this pyrite is apparent on most of the bones soon after exposure to the surface. This is indicated by the rapid appearance of fine, powdery to crystalline gypsum coating bones and teeth, and by the expansion and shattering of some bones and teeth with internal gypsum formation ( ). Protocols are being developed to ensure the preservation of the primary data represented by these important fossils [42], [62].
The bonebed was located many tens of meters underground prior to strip mining operations in the Santa María coal mine. As mining operations proceed, more of the plant debris stratum containing the bonebed is exposed as simultaneous reclamation covers the previously exposed surface. Thus, with the help of mine managers, efficient methodologies for the documentation and extraction of significant fossils have been established [42]. By the end of 2012, an area of approximately 25 ha had been investigated and the areal distributions of 101 vertebrate concentrations were documented; 33 of these consisted of associated dinosaur skeletons (mostly iguanodonts) and 68 consisted of other vertebrate remains (mostly turtles and crocodilians). During this stage of the project, numerous dinosaurs (ornithischian elements and associated skeletons, and saurischian teeth), two types of turtle, crocodilians, fish (both ostheicthyians and selachiens), coprolites, molluscs (freshwater bivalves and gastropods), arthropods (ostracods), and abundant plant remains (logs, plant fragments, palynomorphs, and amber) have been excavated.
The bonebed designated AR-1 contains more than 5000 identifiable vertebrate specimens recovered from isolated skeletal remains and associated individual animals. All fossils receive a consecutive number from the site, each association is numbered as well. Thus:
AR-1-#fossil identifies each fossil found at the Ariño site (the ID written on each fossil);
AR-1/#concentration identifies a collection of bones belonging to a single skeleton;
AR-1-#fossil/#concentration identifies a fossil from a bone concentration # belonging or not belonging to a single skeleton.
The two associated ankylosaur skeletons described herein were separated by 200 meters. The location of the holotype AR-1/10 ( ) was still available for examination and sampling for microfossils in December of 2011 [58], while that of the paratype AR-1/31 ( ) was already inaccessible.
Materials and Methods
Paleontological Ethics Statement
All of the specimens described in this paper (AR-1/10 and AR-1/31) are reposited in the collections of the Fundación Conjunto Paleontológico de Teruel-Dinópolis/Museo Aragonés de Paleontología (FCPTD/MAP). Locality information is available from the registrar of the museum as per museum policy. All necessary permits were obtained for the described study, which complied with all relevant regulations. All of these specimens were collected under permits obtained from the Sociedad Anónima Minera Catalano-Aragonesa.
Nomenclatural Acts
The electronic edition of this article conforms to the requirements of the amended International Code of Zoological Nomenclature, and hence the new names contained herein are available under that Code from the electronic edition of this article. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information viewed through any standard web browser by appending the LSID to the prefix “http://zoobank.org/”. The LSID for this publication is: urn:lsid:zoobank.org:pub:9246FFA7-6271-4734-8E01-5590BE4A80C2. The LSID for Europelta carbonensis is: urn:lsid:zoobank.org:act:089040A3-1BCF-42D1-B99F-94840E2BB96D. The electronic edition of this work was published in a journal with an ISSN (1932-6203), and has been archived and is available from the following digital repositories: LOCKSS (http://www.lockss.org); PubMed Central (http://www.ncbi.nlm.nih.gov/pmc).
Terminology
We do not refer to the “armor” on the skull roof as caputegulae, as we consider these patterns in the Nodosauridae to reflect impressions of scale boundaries on the skull roof as opposed to thickened remodeled cranial bone. We use the term caudal rib instead of caudal transverse process. We employ the monophyletic clade Polacanthidae of Carpenter [63] to facilitate comparison with and discussion of a number of similar taxa (Gargoyleosaurus, Mymoorapelta, Hylaeosaurus, Polacanthus, Hoplitosaurus, and Gastonia). The most recent analysis of polacanthids as a monophylogenetic subfamily of nodosaurids was by Yang and others [64], who similarly defined them as the most inclusive clade containing Polacanthus foxii but not Ankylosaurus magniventris or Panoplosaurus mirus.
Institutional Abbreviations
AMNH, American Museum of Natural History, New York, New York, NHMUK, Natural History Museum, London, England, CEUM, Prehistoric Museum, Utah State University, Price, Utah, DMNH, Denver Museum of Nature and Science, Denver, Colorado, MPC, Geological Institute, Ulaan Bataar, Mongolia, FCPTD/MAP, Fundación Conjunto Paleontológico de Teruel-Dinópolis/Museo Aragonés de Paleontología, Teruel, Spain, FMNH, Field Museum of Natural History, Chicago, MPC, Institute of Geology, Mongolian Academy of Sciences, Ulaan Baatar, Mongolia; INBR, Victor Valley Museum, Apple Valley, California, IVPP, Institute of Vertebrate Paleontology and Paleoanthropology, Beijing, China, KUVP, Kansas Museum of Natural History, Lawrence, Kansas, MPC, Mongolian Paleontological Center, Ulaan Baatar, Mongolia; MNA, Museum of Northern Arizona, Flagstaff, Arizona, NMC, National Museum of Canada, Ottawa, Canada, NMW, National Museum of Wales, Cardiff, England, PIN, National Institute of Paleontology, Moscow, Russia, QM, Queensland Museum, Queensland, Australia, ROM, Royal Ontario Museum, Toronto, Canada, SDNHM, San Diego Natural History Museum, San Diego, California, SGDS, Saint George Dinosaur Discovery Site at Johnson Farm, St. George, Utah, SMP, State Museum of Pennsylvania, Harrisburg, Pennsylvania, SMU, Schuler Museum, Southern Methodist University, Dallas, Texas, USNM, National Museum of Natural History, Smithsonian Institution, Washington D.C.
Comparative Material
In addition to accessing the ever-expanding ankylosaur literature, the senior and third authors have had the opportunity to study firsthand much of the important ankylosaur material collected globally. From the basal thyreophorans: the type material of Scutellosaurus lawleri (MNA P1.175), the type material of Scelidosaurus harrisoni (NHMUK R 1111), and a large, exceptionally well-preserved, articulated Scelidosaurus specimen with intact armor, collected and owned by David Sole and currently exhibited at the University of Bristol. Also, a full cast of the left side of the skeleton (SGDS 1311) exhibited in southwestern Utah was examined.
In regards to Jurassic ankylosaurs: the extensive type and paratype material of Mymoorapelta maysi housed at the Museum of Western Colorado, Gargoyleosaurus parkpinorum (DMNH 27726), and the dentary of Sarcolestes leedsi (NHMUK R 2682) were studied.
Early Cretaceous polacanthine ankylosaur material examined includes Polacanthus foxii (NHMUK R 175, 9293), Hylaeosaurus armatus (NHMUK R 3775), Hoplitosaurus marshi (USNM 4752), and the extensive material of Gastonia burgei material housed at the Prehistoric Museum (including holotype CEUM 1307 and paratype material), and cranial material from a minimum of six individuals at Brigham Young University's Earth Science Museum, together with the postcranial skeleton of an unnamed new species of polacanthine (BYU 245).
Among basal shamosaurine-grade ankylosaurids, Cedarpelta bilbyhallorum (including CEUM 12360 and paratype material), Shamosaurus scutatus (PIN 3779/2), and a cast of the skull of Gobisaurus domoculus (IVPP 12563) housed at the Royal Tyrell Museum were studied.
Among derived North American ankylosaurs, Nodocephalosaurus kirtlandensis (SMP-VP-900), Ankylosaurus magniventris (AMNH 5214, 5859; NMC 8880), Anadontosaurus lambei (NMC 8530), Dyoplosaurus acutosquameus (ROM 784), Scolosaurus cutleri (NHMUK, R 5161), and several important examples of Euoplocephalus tutus, (AMNH 5404, 5409; RTMP 91.127.1) were examined.
Asian ankylosaur material researched include an adult skull of Tsagantegia longicranialis (MPC 100/1306), China, Pinacosaurus grangeri (AMNH 6523) and three undescribed skulls personally excavated by JIK from the Djadokhta Formation, Shabarakh Usu (Flaming Cliffs, Mongolia) and housed at MAS, Talarurus plicatospineus (composite skeleton made up of parts of many individuals assigned to PIN 557), cast skull of Saichania chulsanensis (PIN 3141/251), a relatively complete specimen referred to Saichania with in situ armor but lacking its skull (MPC 100/1305), Tarchia gigantea (PIN 3142/250), a cast skull of Minotaurasaurus ramachandrani (INBR 21004), and a cast skeleton of Crichtonsaurus benxiensis housed in the Museum at the Chaoyang Bird National Geopark, Liaoning.
Numerous nodososaurids were examined, including the Early Cretaceous nodosaurids Sauropelta edwardsi (AMNH, 3016, 3032, 3035, 3036; YPM 5502, 5529, 5499, 5178), Peloroplites cedrimontanus (CEUM 26331 and the extensive paratype material), and Pawpawsaurus campbelli (SMU 73203; = “Texasestes” pleurohalio USNM 337987). The early Late Cretaceous nodosaurids reviewed include Animantarx ramaljonesi (CEUM 6228), Silvisaurus condrayi (KUVP 10296), Nodosaurus textilis (YPM 1815), and Stegopelta landerensis(FMNH UR88) and the Late Cretaceous nodosaurids Panoplosaurus mirus (NMC 2759), Edmontonia rugosidens (USNM 11868; AMNH 5665), Edmontonia longiceps (NMC 8531), Denversaurus schlessmani (DMNH 468), casts of Struthiosaurus austriacus at the Carnegie Museum (PIUW 2349) and Struthiosaurus transylvanicus (NHMUK R 4966).
Enigmatic taxa such as the skull of Minmi paravertebrata (QM {"type":"entrez-nucleotide","attrs":{"text":"F18101","term_id":"4824805","term_text":"F18101"}}F18101), the skeleton of Liaoningosaurus paradoxus (IVPP V12560), and Aletopelta coombsi (SDNHM 33909) were also examined.
Results
Systematic Paleontology
Dinosauria Owen, 1842 [65]
Ornithischia Seeley, 1887 [66]
Thyreophora Nopcsa, 1915 [25]
Ankylosauria Osborn, 1908 [67]
Nodosauridae Marsh, 1890 [68]
Struthiosaurinae Nopcsa, 1923 [69]
Diagnosis
Nodosaurid ankylosaurs that share a combination of characters including: narrow predentaries; a nearly horizontal, unfused quadrates that are oriented less than 30° from the skull roof, and condyles that are 3 times transversely wider than long; premaxillary teeth and dentary teeth that are near the predentary symphysis; dorsally arched sacra; an acromion process dorsal to midpoint of the scapula-coracoid suture; straight ischia, with a straight dorsal margin; relatively long slender limbs; a sacral shield of armor; and erect sacral armor with flat bases. Struthiosaurinae is defined as the most inclusive clade containing Europelta but not Cedarpelta , Peloroplites, Sauropelta or Edmontonia .
Europelta Kirkland, Alcalá, Loewen, Espílez, Mampel, and Wiersma 2013 gen. nov.
urn:lsid:zoobank.org:act:62808E3D-85BE-4AE3-B771-9CFF2C6AC054
Etymology
“Euro” as a contraction for Europe in regard to its origin and “pelta” Greek for shield, a common root for ankylosaurian genera; “Europe's shield”.
Diagnosis
Same as for the only known species below.
Europelta carbonensis Kirkland, Alcalá, Loewen, Espílez, Mampel, and Wiersma 2013 gen. et sp. nov.
urn:lsid:zoobank.org:act:089040A3-1BCF-42D1-B99F-94840E2BB96D
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Etymology
The specific name “carbonensis” from the coal, is in honor of access to the fossil locality in the Santa María coal mine provided by Sociedad Anónima Minera Catalano-Aragonesa (SAMCA Group), which has been extracting coal in Ariño (Teruel) since 1919.
Holotype
AR-1/10, a disarticulated partial skeleton reposited at Fundación Conjunto Paleontológico de Teruel-Dinópolis/Museo Aragonés de Paleontología (FCPTD/MAP). The holotype consists of: a mostly complete skull (AR-1-544), isolated left and right nasals (AR-1-133, and AR-1-639), a dentary fragment (AR-1-362), 15 isolated teeth (AR-1-323 to AR-1-325, AR-1-343, AR-1-358, AR-1-417, AR-1-418, AR-1-423, AR-1-424, AR-1-428, AR-1-454, AR-1-482, AR-1-563, AR-1-564 and AR-1-567), an atlas (AR-1-649), five cervical vertebrae (AR-1-431, AR-1-449, AR-1-533, AR-1-637, AR-1-650), two cervical ribs (AR-1-450, AR-1-4452), AR-1-638 (possibly the first dorsal vertebrae), seven more posterior dorsal vertebrae (AR-1-154, AR-1-155, AR-1-322, AR-1-430, AR-1-448, AR-1-478, AR-1-535, AR-1-556), a section of synsacrum (AR-1-154), three isolated dorsal ribs (AR-1-331, AR-1-333, AR-1-476), seven dorsal rib fragments (AR-1-339, AR-1-341, AR-1-427, AR-1-534, AR-1-641, AR-1-642, AR-1-676), three caudal vertebrae (AR-1-562, AR-1-635, AR-1-636), four chevrons (AR-1-560, AR-1-561, AR-1-569, AR-1-4451), a coracoid with a small portion of scapula (AR-1-657), a scapular blade fragment (AR-1-429), two xiphosternal plates (AR-1-252, AR-1-4675), two partial humeri (AR-1-327, AR-1-655), a right ilium-ischium-pubis (AR-1-479), a left ischium-pubis (AR-1-129), and 70 osteoderms (AR-1-126 to AR-1-128, AR-1-192, AR-1-234, AR-1-241, AR-1-246, AR-1-247, AR-1-272, AR-1-276, AR-1-438, AR-1-444, AR-1-447, AR-1-461, AR-1-462, AR-1-464, AR-1-467, AR-1-472, AR-1-496 to AR-1-530, AR-1-553, AR-1-651 to AR-1-653, AR-1-659, AR-1-675, AR-1-4450, AR-1-4454 to AR-1-4463).
Paratype
AR-1/31, a partial skeleton deposited at Fundación Conjunto Paleontológico de Teruel-Dinópolis/Museo Aragonés de Paleontología (FCPTD/MAP). The paratype consists of a partial left jaw with dentary and surangular (AR-1-3698) and isolated angular (AR-1-2945), 10 teeth (AR-1-3432, AR-1-3495, AR-1-3524, AR-1-3650, AR-1-3699 to AR-1-3701, AR-1-3705, AR-1-3706, AR-1-3961), five cervical vertebrae (AR-1-3586, AR-1-3632, AR-1-3657, AR-1-3671, AR-1-3676), nine dorsal vertebrae (AR-1-3489, AR-1-3586, AR-1-3633, AR-1-3662, AR-1-3672 to 3675, AR-1-3677, AR-1-3704), three to four? dorsosacral vertebrae (AR-1-3450, AR-1-3451), a sacrum (AR-1-3446), a caudosacral vertebra (AR-1-3512), two sacral rib fragments (AR-1-3452, AR-1-3460), 14 caudal vertebrae (AR-1-2950, AR-1-3204, AR-1-3206, AR-1-3243, AR-1-3265, AR-1-3348, AR-1-3398, AR-1-3478, AR-1-3615, AR-1-3616, AR-1-3714 to 3717), a right ilium (AR-1-3490), two left ilium fragments (AR-1-3521, AR-1-3571), two ischia with fused pubes (AR-1-3648, AR-1-3649), a right femur (AR-1-3244), a right tibia (AR-1-3237), a right fibula (AR-1-3238), a calcaneum (AR-1-3239), four metatarsals (AR-1-3100, AR-1-3173, AR-1-3233, AR-1-3234), eight phalanges (AR-1-3032, AR-1-3066, AR-1-3174, AR-1-3179, AR-1-3324, AR-1-3234, AR-1-3292, AR-1-3356), nine unguals (AR-1-2952, AR-1-2986, AR-1-3172, AR-1-3181, AR-1-3182, AR-1-3288, AR-1-3291, AR-1-3386, AR-1-3711), and 90 osteoderms (AR-1-3024, AR-1-3030, AR-1-3074 to AR-1-3076, AR-1-3080, AR-1-3145, AR-1-3159, AR-1-3180, AR-1-3207 to AR-1-3209, AR-1-3216, AR-1-3223, AR-1-3226 to AR-1-3229, AR-1-3292, AR-1-3236, AR-1-3242, AR-1-3338 to AR-1-3340, AR-1-3390, AR-1-3438, AR-1-3447 to AR-1-3449, AR-1-3491, AR-1-3492, AR-1-3494, AR-1-3506, AR-1-3540, AR-1-3572 to AR-1-3576, AR-1-3587, AR-1-3588, AR-1-3590, AR-1-3597, AR-1-3598, AR-1-3608 to AR-1-3613, AR-1-3638, AR-1-3658, AR-1-3680 to AR-1-3684, AR-1-3687, AR-1-3708, AR-1-3720, AR-1-3721, AR-1-3932 to AR-1-3960).
Locality and Horizon
The type locality, Fundación Conjunto Paleontológico of Teruel-Dinópolis locality AR-1, is located east of Ariño, Teruel Province, Spain. The fossil horizon is below the lowest mineable coal seam at Sociedad Anónima Minera Catalano-Aragonesa Group's Ariño coal mine in a plant debris bed in the lower Escucha Formation [42]. The paratype AR-1/31 was located 200 m laterally from the holotype AR-1/10 in the same bed. Pyrite is common within the bone and the surrounding sediment of the bonebed, common also in plant debris beds in the older Wessex Formation on the Isle of Wight [58].
Age
Elsewhere, the Escucha Formation has been interpreted as late Aptian to early Albian in age based on nanofossils, planktonic foraminifera, dinoflagellates and palynomorphs [50], [52]. An analysis of the palynomorphs, ostracods, and charophytes from AR-1 indicates that the site is completely of early Albian age [57].
Diagnosis
The quadrate is shorter and mediolaterally wider than in any other ankylosaur. The posterior margin of the skull is concave in dorsal view. The sacrum is arched dorsally about 55° in lateral view. The pubis is fully and uniquely fused to the ischium with a slot-shaped foramen between the post-pubic process and the position of the pubic peduncle forming an ischiopubis. The tibia is longer relative to the length of the femur (90%) than in other ankylosaurs for which these proportions are known. Laterally compresed, flanged osteoderm with a flat plate-like base is present anteriorly on the pelvic shield.
Description and Comparisons
Skull
The skull (AR-1-544/10) was lying on its dorsal surface and is moderately well preserved although distorted through compaction ( ). The palate is crushed in toward the skull roof, resulting in the medial rotation of both maxillae with the posterior teeth displaced into the posterior palate. The sheet-like palatal bones are highly fragmented. The braincase is crushed along the plane of the cranial nerve openings and the fenestra ovalis completely obscures them. Unexpectedly, the right quadrate ( ) and associated portion of the palate was dislodged from the skull and subsequently crushed across the ventral side of the basicranium. This gives the impression that these bones had been expelled from inside the skull prior to compaction. Both the left and right nasals were separated from the skull and the premaxillae (whereas possibly present upon discovery) have not been identified.
The skull has a minimum length of 370.3 mm from the anterior end of the maxillae to the rear margin of the squamosals. The skull has a maximum width of 299.1 mm at the orbits and narrows to 203.7 mm at the posterior end of the skull at the squamosals, giving the skull the “pear-shaped” dorsal profile characteristic of derived nodosaurids [70], [71]. Although tapering posteriorly, there is no distinct post-temporal notch as in polacanthids and other nodosaurids [63].
The maxillae ( ) are irregularly sculptured externally with a flattened, horizontally oriented buccal recesses that are inset approximately 2 cm. The anterior margin of the maxilla appears to form the posterior margin of a relatively simple naris relative to derived nodosaurids and ankylosaurids. Medially, there is no evidence that the maxilla formed a portion of a secondary palate. The tooth row was arched ventrally with an estimated 22–25 alveoli increasing in size posteriorly as in Edmontonia [72]. In ventral orientation, the tooth rows are only moderately deflected medially, such that the palate would not have had a pronounced hourglass appearance typical of derived nodosaurs such as Pawpawsaurus, Edmontonia, and Panoplosaurus [73]–[75]. However, it is not dissimilar from that of the primitive nodosaurid Silvisaurus [76], [77].
The nasals (AR-1-133/10, AR-1-639/10) are relatively large and subrectangular, tapering somewhat anteriorly ( ). Both nasals extend laterally from their relatively straight, unfused midline suture before flexing down to a sutural contact with the maxillae that extends for most of their length. When rearticulated onto the skull, they appear to fit well, despite the skull's distortion. Most ankylosaurs have fused nasals except the nodosaurids Silvisaurus [76], [77] and Niobrarasaurus [78], although the nasals are unknown in European nodosaurids [24], [32], [33]. A distinct tongue-like process projects from the nasal's posterior margin and would have overlapped the frontals. The external surface is lightly textured and the internal surface is relatively smooth, suggesting the narial passage was large and simple, rather than convolute as in derived nodosaurids and ankylosaurids [79], [80].
The orbits are somewhat crushed and the sutures of the bones surrounding them are obscured by fusion. The orbits are subrectangular in shape, are slightly more elongate anteoposteriorly and are directed anterolaterally. The prominent and evenly rounded suborbital horn is formed mostly from the quadratojugal posterior to the ventral margin of the orbit, as in most derived ankylosaurs [81], [82] and unlike that in polacanthids such as Mymoorapelta, Gargoyleosaurus, and Gastonia where the suborbital horn is below the orbit and is formed exclusively by the jugal [83]–[85]. The suborbital horn appears to be unornamented and hides the head of the quadrate in lateral view.
The lateral wall of the skull extends posteriorly behind orbit with a dorsoventally wide posterior notch, such that the lower temporal opening is just visible in lateral view. There is no lateral wall of skull behind the orbits in polacanthids [70], [81] and most nodosaurids other than Peloroplites [86], Silvisaurus [76], Struthiosaurus transylvanicus [22], [23] and one specimen from the Dinosaur Park Formation assigned to Edmontonia (ROM 1215) [88], although in these taxa the lower temporal opening is still visible in lateral view as in Europelta. The lower temporal opening is completely obscured in lateral view in Cedarpelta [84], [86], Shamosaurus [89]–[91], Gobisaurus, [92] Zhongyuansaurus [93] and all derived ankylosaurids.
Although the palate is fragmented and crushed along the internal surface of the skull roof, the fragments of the vomer suggest it did not extend ventrally to the level of the tooth row. Additionally, the broad sheet-like pterygoids appear to have been flexed nearly dorsally against the anterior portion of the basicranium as in nodosaurids and not like the open transversely oriented pterygoids characteristic of ankylosaurids or polacanthids [94].
The posterolateral margin of the pterygoid is fully fused to the quadrate. There is a sutural contact between the straight, nearly vertical quadrates and the quadratojugal laterally. The quadrates are wide transversely and thin rostrocaudally as compared to the mediolaterally narrower quadrates of other ankylosaurs [82]. The contact with the squamosal is also transversely wide, unlike the narrow, rounded contact seen in many ankylosaurs such as Mymoorapelta (Kirkland, pers. obs.) and Cedarpelta [63], [86]. The mandibular articulation is proportionally wider than in any other ankylosaur examined as a part of this study and the medial condyle larger than the lateral condyle. The ratio of mediolateral quadrate width to dorsoventral quadrate length is 0.77 (94 mm/122 mm). The anteropostior length of the quadrate condyle is 31 mm. There is no fusion between the quadrates and the paroccipital processes.
Vertical compaction has obscured the posterior view of the skull, in particular the foramen magnum and the supraoccipital. However, even with compaction it is apparent that in occipital view the skull was subrectangular and wider than tall as in Gargoyleosaurus, Gastonia, and most other derived anklylosaurs, and unlike the narrow, highly arched occipital region of Struthiosaurus [22]. The paroccipital processes extend horizontally lateral to the foramen magnum and then flare dorsoventrally by approximately 100% of their minimum widths. They angle posteriorly at about 30 degrees when viewed ventrally ( ). In morphology and orientation, they are most similar to those in Gargoyleosaurus [95] although ventral twisting is not present. In most other ankylosaurs, the paroccipital processes extend straight laterally [81], [96] or may be flexed ventrally as in Gastonia [83]. A triangular wedge of bone of unknown identity is fused to the anterior ventrolateral margin of the paroccipital, separating it from the quadrate.
The subspherical occipital condyle ( ) has a width of 59.4 mm and height of 46.5 mm and lacks a distinct neck to separate it from the rest of the basicranium. Although no cranial sutures are visible, the occipital condyle does appear to be composed exclusively of the basioccipital. It is similar in overall morphology to that of the basal ankylosaurid Cedarpelta [88] except that the occipital condyle angles somewhat ventrally, but not as much as in more derived nodosaurids [71], [82]. The ventral surface of the relatively elongate basioccipital is broadly convex. Again, as in Cedarpelta [88], there are no distinct, separate basal tubera between the basioccipital and the short basisphenoid, but instead there is a prominent transverse flange extending across the ventral surface of the basicranium along the line of this suture. The pterygoid processes appear to be short, but are completely obscured by crushed pterygoids bone fragments that wall off the anterior part of the braincase as in most nodosaurids.
The skull roof ( ) is roughened texturally by remodeling of the bone surface as in Cedarpelta, the nodosaurids Sauropelta and Peloroplites, and the shamosaurine-grade ankylosaurids Shamosaurus and Gobisaurus [81], [86], [88]. Europelta differs from these specimens in that some of the margins of the scale impressions on the skull roof are visible, as seen in Edmontonia, Panoplosaurus and Struthiosaurus [22], [77]. These scale margins are represented by shallow grooves that are difficult to see relative to the textured surface of the skull and the cracks in the bone due to compaction. These grooves are particularly evident along the lateral margins of the skull roof above the orbit. An extensive median scale appears to have covered much of the central portion of the skull between and posterior to the orbits on the frontals and parietals as other nodosaurids [63], [82]. There does not appear to be any distinct nuchal ornamentation. The skull is thickened above the orbit, but there is not a distinct supraorbital boss, a condition similar to Peloroplites, Cedarpelta, Shamosaurus, and Gobisaurus [86], [88]–[90], [92]. Narrow grooves along the margin of the skull in this area above the orbits suggest that a particularly robust pair of scales were present in this area as indicated by a deep groove bisecting this ornamented area directly above the orbit. Weak grooves delineate a small scale without underlying ornamentation separating the posterior supraorbital scale from the squamosal horn forming the posteriolateral margin of the skull roof. The squamosal horn is ornamented by narrow grooves radiating from its apex onto the skull roof. Grooves on the anterolateral sides of the fronto-parietal scale appear to delineate two scales between the anterior supraorbital scales. Unfortunately, no distinctive scale boundaries are recognizable on the nasals, although the dorsal surfaces of the nasals are textured. Several elongate scales rimmed the lateral raised margin around the orbit.
In dorsal view, the posterior margin of the skull is concave, whereas it is nearly straight or convex in all other nodosaurids. This reflects the posterior angulation of the paraoccipital processes and the squamosal horns. Interestingly, the occipital condyle is barely visible, though not completely obscured in dorsal view. There is no evidence of any distinct nuchal sculpturing. The skull roof is relatively flat but a slight dome may have been present prior to crushing. However, it is clear that the skull roof is not as highly domed as in many other nodosaurids, such as Struthiosaurus [22], [26].
Attempts were made to image the skull using X-ray photography and CT scanning. The abundance of pyrite present in the skull ( ) presents a strong limitation in the use of these techniques as pyrite is opaque to X-rays.
Mandible
A small dentary fragment extending for only four complete alveolae (AR-1-133/10) was preserved from the holotype skeleton ( ). However, a robust left dentary and splenial are preserved together (AR-1-3698/31) from the paratype specimen ( ). The splenial is not in its posteriomedial position relative to the dentary, but is fused across the posterior portion of the tooth row transversely. Additionally, an isolated left angular with a distinct highly sculptured scale along its ventral margin (AR-1-2945/31), was recovered ( ).
The dentary is 184.7 mm long with a minimum of 21 tooth positions, with no possibility of more than two unpreserved alveoli as determined by the position of the suture with the angular and surangular. As with the maxillary teeth, the alveoli are more than twice as large posteriorly. There is only 1.5 cm between the anteriormost alveoli and the symphysis, suggesting that there may have been premaxillary teeth as at least nine anterior teeth would have been positioned to oppose the premaxilla. The primitive ankylosaurs Sarcolestes [34], [98], Gargoyleosaurus, [85], Silvisaurus [76], Animantarx [97], Sauropelta [99], Anoplosaurus [17], Hungarosaurus [33] and Struthiosaurus [22] have a short anterior diastema, and thus a narrow predentary, whereas this diastema is longer in ankylosaurs with wide predentaries. However, the symphysis in Europelta is robust and dorsoventrally deeper (45.0 mm deep and 29.00 mm across) than in ankylosaurs [82], and is most similar to the deep symphysis of Hungarosaurus [32], further suggesting a reduced predentary with a rudimentary ventral process. The symphysis is marked by two deep anteroposteriorly directed grooves. A row of foramina extends posteriorly on the lateral surface of the dentary from just dorsal to the buccal recess to the notch for the surangular, whereas nutritive foraminae are not clearly visible ventral to the alveolae on the medial side of the dentary as in other ankylosaurs. The recessed tooth row is deflected medially and forms a convex arch in lateral view. The dentary of Hungarosaurus is deeper dorsoventrally than that of Europelta [33].
The splenial ( ) is a thin bone with a convex ventral margin 156.6 mm long that contacts the angular. It has the appearance of an obtuse triangle in medial view. There is large, well-developed intermandibular foramen (7 mm long and 5.3 mm wide) 50 mm from its anterior end.
The angular ( ) has a maximum length of 175 mm. The lateral margin is highly rugose, because the bone is textured and remodeled to support a large scale, extending about 10–12 mm ventral to the ventral margin of the angular for most of its length. A distinct ridge marks the dorsal limit of the mandibular ornament medially, where it is in contact with the ventral margin of the splenial. Dorsal to this contact the bone is smooth. The ventral extent of the textured bone supporting the mandibular scale is similar to that observed in ankylosaurids such as Euoplocephalus [95] and Minataurasaurus [100], rather than the more lateral orientation found in Gargoyleosaurus [93] and in nodosaurids like Sauropelta [99] and Panoplosaurus [101].
Teeth
A large number of teeth are preserved from both the holotype AR-1/10 (20+) and the paratype AR-1/31 (15+) although many have drifted away from the alvaeolae. We assume that the teeth associated with the holotype pertain to the maxilla (several are preserved in the palate and in the maxilla) and those of the paratype pertain to the dentary (several are preserved in the dentary). In general, the cutting surfaces of the teeth are not well preserved, but a few exceptions exist. Wear facets were not observed on any of the teeth. The roots for both dentary and maxillary teeth are swollen lingually, are three to four times the length of the crowns, and are subquadrate in cross-section. One small tooth (AR-1-343/10) is more highly asymmetrical mesiodistally and may represent a premaxillary tooth ( ).
The isolated maxillary teeth ( ) have a weakly developed labial cingulum and a strongly developed lingual cingulum. The best preserved right tooth AR-1-324/10 is 11.50 mm wide, 9.99 mm tall with seven to eight mesial denticles and five to six distal denticles ( ). A large right tooth AR-1-564/10 is 17.23 mm wide and 12.95 mm tall with eight to nine mesial denticles and ∼six to seven distal denticles ( ).
The isolated dentary teeth ( ) are identical to the maxillary teeth and have a weak lingual cingulum and a strongly developed labial cingulum. The best preserved tooth AR-1-3700/31 is 14.03 mm wide and 12.69 mm tall with eight to nine mesial denticles and six to seven distal denticles ( ). The largest dentary tooth AR-1-3650/31 is 16.58 mm wide and 13.50 mm tall ( ).
With their relatively large size and well-developed cingula, the teeth of Europelta are most comparable to those of other nodosaurids [72]. They similar to the teeth of Cedarpelta, Sauropelta [34], [97], [102], Edmontonia and Panoplosaurus [72], but are not as high crowned as in the Jurassic ankylosaurs Sarcolestes and Priodontognathus [103], the Jurassic polacanthids Gargoyleosaurus [93] and Mymoorapelta (Kirkland, pers. obs.), the nodosaurids Peloroplites [84] or Hungarosaurus [33]. Additionally, the large teeth of Gobisaurus are more inflated labiolingually than in Europelta and other ankylosaurs. The teeth of Gastonia and putative Polacanthus teeth are also inflated, but are smaller proportionally [83], [103]. The teeth of Europelta differ from an isolated tooth from the Cenomanian of France which is about half the size, and proportionally is longer mesiodistally with more deeply divided denticles forming ridges on the labiolingual surfaces of the tooth [104]. Likewise, lower Cenomanian teeth assigned to “Acanthopholis” have more deeply divided denticles in what is a proportionally taller tooth [17]. The teeth of Struthiosaurus languedocensis [31] from the lower Campanian of France also differ in size and in having longer, lower tooth crowns.
Axial skeleton
There are numerous ribs and vertebrae preserved from the holotype (AR-1/10) and the paratype specimen (AR-1/31). Vertebral measurements are presented in .
Table 1
Europelta VERTEBRAL MEASUREMENTS IN MMAnteriorPosteriorOverallNeuralNeuralTotalNeuralTransverseTransverseCentrum FaceCentrum FaceCentrumCanalCanalVertebralSpineProcessesProcessesWidthHeightWidthHeightLengthWidthHeightHeightHeightWidthLength(above canal)AR/10* estimated Cervical Vertebrae AR-1-431109.278.8--*85.230.630.6186.186.4203.879.2AR-1-449100.174.3--66.331.622.4185.590.9198.261.1AR-1-53394.969.9--*81.7*23.1*22.9*218.9*133.2*203.172.5AR-1-637*81.5*60.5*78.3*60.5*96.8*25.8*17.1---47.3AR-1-63893.168.9*85.0*73.875.620.630.3--*160.286.7AR-1-64973.270.199.961.262.028.819.1---77.4AR-1-650*81.4*57.780.662.5*61.0*26.8*14.4122.5*56.0*104.1*29.0 Dorsal Vertebrae AR-1-154----79.4------AR-1-155*60.0*69.875.5*68.4*79.2--*159.5129.3--AR-1-32289.976.394.678.982.514.824.0-133.1-*85.2AR-1-43091.477.797.583.684.6*20.1*25.2222.9-*175.8*76.5AR-1-44890.378.595.579.090.7*16.8*23.7--*120.0*73.3AR-1-47891.478.294.884.0*86.116.722.1219.8139.2114.485.7AR-1-53598.581.792.8*82.093.522.426.3239.9-142.591.2AR-1-556*88.4*79.2*83.0*76.3*70.7*21.0*27.4---*89.8 Caudal Vertebrae AR-1-56276.273.881.679.372.414.326.0178.885.8193.964.8AR-1-63582.279.892.492.179.419.426.7192.280.7240.674.0AR-1-63682.780.789.194.2*66.2*23.10*21.3*193.0*88.8*211.377.4AR/31 Cervical Vertebrae AR-1-3632*76.5*63.4*66.0*63.9*52.9*9.0*18.5*154.8*68.2*139.6*60.6AR-1-365767.953.576.0-*51.813.721.1--*151.650.1AR-1-366269.160.3*67.160.2*53.3------AR-1-3671*69.0*49.5*78.0*52.4*53.9*25.2*11.8*134.1*57.3*120.5*41.3AR-1-3676*52.1*55.6*60.3*60.0*60.4*8.8*19.5*136.1*51.1*89.8*26.9 Dorsal Vertebrae AR-1-348965.659.365.061.879.0*12.0*15.1178.9105.0-68.0AR-1-358675.661.672.761.854.614.319.6157.574.7140.962.9AR-1-363376.460.167.058.662.713.118.9178.594.7*139.173.2AR-1-367268.752.777.157.873.5------AR-1-367366.660.566.555.472.5*11.8*15.5--*119.658.0AR-1-3674*59.1*65.7*53.7*63.8*72.9--*168.5*88.7-85.1AR-1-3675*64.656.966.763.266.315.922.3*171.8*104.4*133.974.2AR-1-370467.062.8*64.7*59.979.1-*14.4--*154.663.3 Caudal Vertebrae AR-1-295031.125.328.724.250.26.14.937.09.0--AR-1-3204---------49.8-AR-1-320639.029.535.430.050.6*8.04*7.0----AR-1-324343.038.838.731.751.05.36.047.320.7--AR-1-326545.1*30.045.232.052.6------AR-1-3348*60.749.5*55.3*45.5*53.7--73.115.1*99.038.2AR-1-339842.534.632.435.351.93.36.250.113.242.5-AR-1-347848.934.846.738.252.24.88.052.2-46.1-AR-1-361551.437.149.026.156.16.511.247.7*8.4--AR-1-3616*48.9*42.8*45.2*39.7*56.9--*58.9-49.3-AR-1-371430.223.7--42.15.74.8----AR-1-3715--25.622.237.7--31.6---AR-1-3716--*48.143.9---*70.115.5--AR-1-371761.440.058.139.654.27.9*5.1--95.429.7
The complete atlas (AR-1-649/10) from the holotype has a total width of 195.6 mm ( ). The neural arch is divided dorsally with the left side fused to the centrum and the right side unattached. The anterior face of the atlantal intercentrum is 73.7 mm wide by 71.7 mm tall and its posterior face is 99.9 mm wide by 61.2 mm tall with a length of 62.0 mm. The axis is not present in either associated skeleton.
There are five post-axis cervical vertebrae (AR-1-431/10, 449, 533, 637, 650) preserved from the holotype skeleton ( ) and five from the paratype skeleton; of which four are illustrated (AR-1-3586/31, 3632, 3671, and 3676) ( ). Overall, they are typical of most other described ankylosaur cervical vertebrae. The centra are amphicoelus, wider than tall, anterorposteriorly short, and medially constricted. Anterior and mid-cervical vertebrae have the anterior faces of the centra dorsally elevated relative to the posterior faces. This is in contrast to the posterior cervical centra which have horizontally aligned faces. The ventral sides of the anterior centra are characterized by two anteroposteriorly-oriented paired fossae separated by a low keel ( ), as observed in the primitive nodosaurid Animantarx [97]. The dorsal ends of the neural spines are expanded transversely. AR-1-638/10 may either be the last cervical vertebra or the first dorsal vertebra based on the position of the parapophyses.
There are two complete cervical ribs preserved for the holotype. AR-1-450/10 is a relatively anterior cervical rib ( ) and AR-1-4452/10 is a posterior cervical rib. There is no evidence of fusion of cervical ribs to the cervical vertebrae as in the ankylosaurid Saichania [105], [106] or Ankylosaurus [107]. The cervical ribs are Y-shaped overall and much like the cervical ribs of other ankylosaurs such as Silvisaurus [76], [78], [82].
Several amphiplatan to amphicoelus dorsal vertebra are preserved: eight for the holotype AR-1/10 and nine for the paratype AR-1/31. The diapophyses originate at the level of the post-zygopophyses at the dorsal extent of the neural canal. The more anterior vertebrae have large cylindrical amphiplatan centra which lack a constricted ventral keel with circular neural canals and fused ribs (AR-1-448/10, 478, and 535). The broad transverse processes are T-shaped in cross-section and angled dorsally, unlike the laterally directed transverse processes in Polacanthus [10], [38]. Two dorsal vertebrae from the holotype appear to be pathological with the centra overgrown by about 0.5 cm of lumpy reactive bone ( ). One of these pathologic vertebrae (AR-1-535/10) has fused ribs ( ) although the other (AR-1-430/10) does not ( ). Two additional dorsal vertebrae (AR-1-478/10, 448) with fused ribs are not pathologic ( ). More posterior dorsal vertebrae have shorter, taller, more medially constricted centra, laterally compressed neural canals, more dorsally directed transverse processes, and lack fused ribs (AR-1-155/10, 322, and 556). The neural spines are thin and rectangular with narrowly expanded dorsal ends as in Sauropelta [99]. The neural spines are oriented dorsally as opposed to the posteriorly inclined neural spines of some other ankylosaurs such as Sauropelta [97]. None of the paratype vertebrae (AR- 1-3489/31, 3633, 3662, 3672, 3673, 3674, 3675, 3677 and 3704) have fused ribs ( ), suggesting that this character is ontogenetic because the paratype AR-1/31 represents a somewhat smaller (and presumably younger) individual than the holotype AR-1/10. More expanded neural spines are present in Shamosaurus [91].
There are a number of rib fragments preserved with AR-1/10, but there are only three (AR-1-331/10, 333, 476) relatively complete ribs ( ). As with most other ankylosaurs, the ribs are sharply arched and L-shaped in cross-section proximally in anterior ribs and broadly arched and T-shaped in cross-section proximally in more posterior ribs.
The sacrum is not preserved in AR-1/10 other than an anteriormost centrum (AR-1-154/10) of the synscacrum ( ). However, for the paratype, AR-1-3466/31, there is a largely complete but fragmented synsacrum ( ) that includes an interpreted anteriormost synsacral centrum (AR-1-3451/31), more of the anterior synsacrum composed of two dorsal centra (AR-1-3450/31), four sacral vertebrae with the sacral ribs from the left side (AR-1-3446/31), two sacral ribs from the right side (AR-1-3452/31, 3460), and one caudosacral vertebra (AR-1-3512/31). Given that at least one intermediate and one anterior fused synsacral dorsal vertebra are missing, the vertebral formula for the synsacrum would be five or more dorsosacral vertebrae, four sacral vertebrae, and one sacrocaudal vertebra. The entire synsacrum would have been over 50 cm long and measures about 44 cm across the sacral ribs. The middle section of the preserved dorsal synsacrum thins anteriorly from about 7 cm wide to about 5.5 cm wide. It then expands again anteriorly as indicated by the anteriormost centrum of the synsacrum. This differs from the sacrum of Euoplocephalus [108] and Saichania [106] in which each centrum making up the synsacrum is constricted medially. The sacrum is distinctive in being more strongly arched anteroposteriorly than other described ankylosaur sacra. The neural spines are dorsoventrally shorter than the height of the centra and are fused into a vertical sheet of bone along the length of the sacrum. The caudosacral neural spine is longer and unexpanded, transitional in form between the sacral neural spines and those of the proximal caudal vertebrae. The neural spines are broken off the anterior end of the synsacrum. The ventral side of the sacrum and anterior synsacrum is longitudinally depressed. The distal ends of the sacral ribs are expanded and the most robust medial sacral rib is about 50% taller (9.4 cm) than wide (6 cm) at its attachment with the ilium. There is no sign of expansion of the dorsal termination of the neural spine on the sacrocaudal vertebra. Additionally, the caudal rib is reduced compared to the sacral ribs.
The sacrum of Struthiosaurus languedocensis [31] is similar overall, but based on the description is not so strongly anteroposteriorly arched as in Europelta. Similarly, the sacrum of Hungarosaurus, as exhibited at the Hungarian Natural History Museum, appears to be moderately arched. The moderate angulation of the faces of the sacral centra (somewhat wedge-shaped in lateral view) in Anoplosaurus [17] indicates that a moderately arched sacram may have been present in this taxon as well. Among North American nodosaurids, we have observed only a moderate anteroposteriorly arching of the synsacrum of Silvisaurus, which appears to be restricted to the posterior part of the sacrum and two sacrocaudals. In other ankylosaurs, the downward flexure of the tail from the hips is taken up in the proximal caudal vertebrae as in Mymoorapelta [84], [109] and Euoplocephalus [70], [82].
Only three proximal caudal vertebrae (AR-1-562/10, 635, 636) are present ( ). The proximal-most caudal vertebrae are not preserved for the holotype. The preserved vertebrae probably represent caudal vertebrae positions in the interval of about 3–7. The centra are anteroposteriorly shorter than dorsoventrally tall and somewhat wedge-shaped in anterior and posterior views. The posterior chevron facets are well developed. The neural spines are inclined posteriorly and the dorsal ends of the neural spines are only slightly expanded transversely as in Gargoyleosaurus [95] and some other ankylosaurs such as Cedarpelta [86], Edmontonia [110], Hungarosaurus [32] and Euoplocephalus [70], [82]. The neural spines are strongly expanded in most polacanthids such as Mymoorapelta [84], [109], Gastonia [83], and Polacanthus [10], and some North American nodosaurids such as Sauropelta [99], and Silvisaurus [76]. The neural spine of AR-1-562/10 is broken, erroneously giving it the appearance of being strongly inclined posteriorly. The caudal ribs (transverse processes) in Europelta originate high on the sides of the centrum and angle ventrally proximal to flexing laterally, giving them a dorsally concave profile in anterior view like Hungarosaurus, Struthiosaurus, and Peloroplites, and unlike the ventrally flexed caudal ribs of many polacanthids [10], [84], [109] and the caudal vertebra assigned to “Acanthopholis” [17] or straight caudal ribs of Gargoyleosaurus [95], Cedarpelta, Peloroplites [86], and Edmontonia [87]. The proximal caudal ribs of Hylaeosaurus differ in being swept back posteriorly [111]. The lateral terminations of the caudal ribs do not expand dorsoventrally as they do in Peloroplites [86] and Struthiosaurus, which actually appear to bifurcate [25], [26].
Additionally, there are four chevrons preserved from about the same region of the tail (AR-1-560/10, 561, 569, and 4451) of which three are illustrated ( ). The proximal chevrons are approximately as long as the neural spines as in most other ankylosaurs. They are relatively straight and expanded into teardrop shapes distally in lateral view. Unlike in many ankylosaurs, there is no fusion of proximal chevrons to their respective caudal vertebrae as in Pinacosaurus and Saichania [105], [106], Ankylosaurus [107], [112], and Edmontonia (ROM 1215) [87].
Several more distal caudal vertebrae are preserved in the paratype. The two most proximal of these (AR-1-3348/31, AR-1-3717/31) have centra of nearly equal height, width, and length, with a ventral groove, and caudal ribs shorter than the diameter of the centrum that extend laterally and angle posteriorly ( ). The chevron facets are well developed with the posterior facets more strongly developed than the anterior facets. The neural spines are not developed and the zygapophyeses only extend a short distance beyond the anterior and posterior margins of the centra. These vertebrae are interpreted to represent mid-caudal vertebrae. Two more distal mid-caudal vertebrae (AR-1-3616/31, AR-1-3716/31) are similar in morphology except that the caudal ribs are reduced to anteroposteriorly directed ridges on the lateral margins of the centra ( ). Their neural spines incline posteriorly, merging with the postzygapophyses as posterior processes extending laterally past the faces of the centra to overlie and articulate between the paired prezygapophyses of the immediatly distal vertebra. This morphology is retained in the distal caudal vertebra. More distally, as in AR-1- 2950/31, 3206, 3243, 3265, 3478, and 3615, the caudal ribs are lost and the centra become more elongate ( ). Unlike many ankylosaurs, the faces of the centra maintain a well-rounded to heart-shaped surface distally down the caudal series [82]. For many of these vertebrae, ventrally anteroposteriorly elongated skid-shaped (inverted T) chevrons are fused to the posterior chevron facets. Fusion of distal chevrons to their respective vertebrae is widespread among ankylosaurs [84], [106], [110] although it is not present in some, such as Nodosaurus [113]. One pair of distal caudal vertebrae is fused by their mutually shared chevron ( ) such as has been documented in Mymoorapelta [84]. The most distal four caudal vertebrae ( ) and their chevrons are fused together in AR-1-3204/31 to form a tapering, terminal rod of bone at the end of the tail somewhat similar to that of Sauropelta [71].
Pectoral Girdle
Parts of the right scapulocoracoid are preserved. A portion of the distal scapular blade (AR-1-429/10) is preserved with a portion of the distal ventral margin missing with a curved section broken away. There is no evidence of any distal expansion of the scapular blade as in many nodosaurids [94].
The coracoid (AR-1-657/10) is preserved with only the most proximal portion of the scapula fused on ( ). It appears to have been sheared off just dorsal to the suture between the coracoid and the scapula, perhaps in the process of removing the overlying coal seam. The coracoid is relatively equidimensional (201.3 mm long by 186.5 mm tall) relative to the elongate coracoids characteristic of many other nodosaurids [114] such as Peleroplites [86], Texasites [77], [115], and Animantarx [97]. The medial surface is concave and the lateral surface is convex giving it a bowl-shaped appearance. The ventral margin is evenly convex as in many polacanthids and nodosaurids and there is no anteroventral process as in all ankylosaurids, including Shamosaurus [91], [94]. The articular surface of the ventrally directed glenoid is wide, bounded by a flange that extends beyond the medial surface of the coracoid.
Both xiphisternal plates are preserved ( ). The best preserved xiphisternal is approximately 350 mm long. They appear to be arcuate flat bones. Xiphisternal plates are only known in a few nodosaurids, but those of Europelta, whereas similar in overall shape to other nodosaurid xiphisterna, are not fenestrate or scalloped along their margins as in North American nodosaurids for which they are known [82], [87], [116].
Forelimb
Parts of both humeri are preserved. The right humerus (AR-1-655/10) is represented by the proximal end ( ). It is 249.2 mm wide with a well-developed proximal head 91.9 mm wide that extends onto the posterior side of the humerus. Distinct notches separate both the laterally directed deltopectoral crest as in nodosaurids such as Sauropelta [70], [71], [99] and the internal tuberosity from the humeral head. The deltopectoral crest extends lateraly from the humerus and is not flexed anteriorly as in polacanthids and ankylosaurids [94].
The left humerus (AR-1-327/10) is represented by a midshaft for which both the proximal and distal ends appear to have rotted off and the core of the shaft has rotted away ( ). The shaft is deeply waisted relative to the proximal and distal ends. Although relatively uninformative, enough of this humerus is preserved to indicate that the deltopectoral crest would have made up less than 50% of the length of the humerus as in nodosaurids [71], [117] and in the basal ankylosaur Mymoorapelta (Kirkland, pers. obs.) compared to the longer deltopectoral crests of ankylosaurids [70], [71]. Overall, the humerus of Europelta is similar in proportions to Niobrarasaurus [118], [119]. The wide proximal end of the humerus figured by Ősi and Prondvai [120] as cf. Struthiosaurus is similar to that of Europelta, whereas the humerus of co-occuring Hungarosaurusis is more slender proportionally.
Among the nine unguals preserved for AR-1/31, one specimen (AR-1-3711/31) may represent a manual ungual. It is more equidimensuional than the other eight more elongate unguals.
Pelvic Girdle
The right ilium of AR-1/10 is fused with its ischium and pubis (AR-1-479/10) which are flexed medially due to compaction ( ). The acetabulum is completely enclosed as in all derived ankylosaurs [70], [71], [82], [94], [108]. Only Mymoorapelta is known to retain an open acetabulum [84], [109]. The acetabulum is directed verntrally and is situated medially near the contact of the ilium with the sacrum so that the ilium extends far out beyond the acetabulum laterally for a distance nearly equal to its width. The lateral and anterior margins of the laterally oriented ilium are broken away. The prepubic portion of the ilium diverges from the midline of the sacrum at about 30 degrees and is thickened ventrally along its midline. Large, fairly equi-dimensional, closely appressed osteoderms (7-10 cm in diameter) cover the dorsal surface of the ilium posterior to and medial to the acetabulum. As discussed below, this morphology of sacral armor compares well with “Category 3” pelvic armor of Arbour and others [121]. Anteriorly, the smooth dorsal surface of the ilium is exposed. The pubis is fully fused to the anterior margin of the ischium with no visible sutures; its presence is indicated by a slot-shaped foramen along the anterior side of the ischium. This foramen represents the obturator notch between the postpubic process and the main body of the pubis as in Scelidosaurus and stegosaurs [122]. The distal end of the ischium is broken away. Additionally, AR-1-129/10 is a poorly preserved, proximal left ischium with the pubis fully fused to its anterior margin ( ).
Beyond some relatively uninformative fragments of the ilium ( ), AR-1/31 includes both the right (AR-1-3648/31) and the left (AR-1-3649/31) ischia with fully fused pubes ( ). Both exhibit the slot-shaped foramen along the anterior side of the ischium formed by the obturator notch. The proximal ends appear enrolled such that the anterior and posterior margins are nearly parallel due to compaction. Both display an anterior kink at their distal end as in Cedarpelta [86], [88], but overall are straight-shafted as in the Ankylosauridae [70], [82], [123] and the other European nodosaurids Struthiosaurus [31] and Hungarosaurus [32]. The distal end of the left ischium is the best preserved and measures 299.9 mm long along its anterior margin, including the fully fused pubis forming an ischiopubis. Given the asymmetry of the proximal end of the fused ischium and pubis and the position of the obturator foramen, it appears that the pubis still makes up some of the acetabular margin. The contact between the ilium and the fused ischiopubis is straight with about one-fourth to one-third of the acetabulum formed by the fused ischiopubis.
A straight ischium has been considered to be the primitive character state for ankylosaurs, with the bent ischium of Polacanthus and nodosaurids, a derived character [63], [82], [83], [94], [114], [123]. It is possible that as opposed to being primitive, a straight ischium may be secondarily acquired in the ankylosaurids and European nodosaurids. The only known ischium from the Jurassic ankylosaur (Mymoorapelta) is bent, a trait that is also observed in some stegosaurs such as Kentrosaurus [124]. Stegosaur ischia, even when straight, have an angular thickening near the mid-point of the posterior margin [124] that is shared by the polacanthids Mymoorapelta (Kirkland pers. obs.) and Gastonia [83]. Europelta is the oldest known ankylosaur preserving a straight ischium. The slight kink in the distal end of the ischium of Europelta suggests the straight ischium in European nodosaurids and ankylosaurids is achieved by shortening the ischium distal to the bend.
Hindlimb
The right femur, tibia, and fibula were closely associated ( ). The robust right femur (AR-1-3244/31) is 502.9 mm long and 178.9 mm wide at the proximal end and has been flattened anteroposteriorly, with the most distortion to the mid-shaft region. The femoral head is distinct with much of its articular surface directed dorsally and only somewhat medially. It forms an angle of about 115° with the long axis of the femur. The femoral head is directed more dorsally under the ilium in polacanthids [7], [12], [82], [95], [125], and several nodosaururids. In addition, the femoral head of Europelta is expanded such that it overhangs the femoral shaft both anteriorly and posteriorly. The greater trochanter is well demarcated from the femoral head by a constriction across the proximal end of the femur, and the anterior trochanter forms a ridge ventral to the greater trochanter that is fully fused to the femur. The robust fourth trochanter overlaps the midpoint of the femoral shaft and its midpoint is located proximal at the midpoint of the femur. Polacanthids and nodosaurid ankylosaurs have this configuration, whereas in ankylosaurids the fourth trochanter is distal to the middle of the shaft [63], [82], [95], [120], [125]. The distal end of the femur is flattened and forms a planar articular surface relative to the straight femoral shaft. The intercondylar notch is not expressed ventrally, and is better developed posteriorly than anteriorly
The right tibia (AR-1-3237/31) and fibula (AR-1-3238/31) were closely associated ( ) and post-depositionally compressed. Compression has distorted the distal end of the tibia such that the wide posterior surface is twisted counterclockwise in line with the wide lateral side of the anterior end relative to the orientation of the proximal and distal ends of the tibia in most other ankylosaurs, such as Mymoorapelta [84] (Kirkland, pers. obs.). The fibula was taphonomically displaced ventrally and with the ventral end rotated posteriorly relative to its position in life with the tibia.
The tibia is 458.8 mm long and robust for its entire length ( ) as in Cedarpelta [86]. The proximal end is 169.2 mm wide by 93.1 mm wide and its distal end is 146.8 mm wide by 70.2 mm. It is significantly more narrowly waisted in Mymoorapelta [84], Gastonia [83], Polacanthus [7], [12], [18], Sauropelta [69], [71], [99], [108], Peloroplites [86], and in Zhejiangosaurus [126] and ankylosaurids like Saichania [106]. The cnemial crest is broadly rounded. The even curvature of the distal end of the tibia suggests that the astragalus was fully fused to it with no evident sutural contact as in most ankylosaurs [63], [82], [121]. The astragalus is not fused to the distal end of the tibia in Mymoorapelta [84], Gastonia [83], Hylaeosaurus [11], and Peloroplites [86].
Generally, ankylosaurids have tibiae that are less than two-thirds the length of their femora, as opposed to nodosaurids which have proportionally longer lower leg elements [127]. With a tibia to femur ratio of 0.91, Europelta has the proportionally longest tibia of any ankylosaur for which this ratio is known. Both Cedarpelta and Peloroplites have relatively longer tibiae than other ankylosaurs [86], with a tibia to femur ratio of 0.82 in both. Peloroplites differs in its proportionally more narrowly waisted tibial shaft.
The fibula is 395.5 mm long ( ) and laterally flattened. The proximal end is not expanded anteroposteriorly, such that the slender fibula changes little in size and shape from the proximal to distal end. In lateral view, the proximal end is rounded and the distal end is concave. In cross-section, it is flattened medially and convex laterally. It is longer relative to the tibia than in most other ankylosaurs [108].
A calcaneum (AR-1-3289/31) was identified in association with the lower right leg of AR-1/31. It is laterally compressed, convex laterally and concave medially ( ). Its dorsal margin is flattened where it would articulate with the fibula. Calcanea are practically unknown in ankylosaurs, but one has been identified in the juvenile specimen of the derived ankylosaur Anodontosaurus [128]. The type of Niobrarasaurus coleii preserves an articulated lower hind limb, with an astragalus fully fused with the tibia and possessing an articulation with the distal end of the fibula and an unfused calcaneum of similar morphology to that of Europelta [118]. The calcaneum is fully fused to the distal end of the fibula in Saichania [106].
A number of metatarsals and phalanges are associated with AR-1/31. The metatarsals have subrectangular proximal ends, indicating that they were closely articulated in a well-integrated pes in life ( ). The pedal phalanges ( ) are short, as in other ankylosaurs. There are eight relatively large, elongate, spade-like unguals ( ) of a morphology similar to pedal unguals in other ankylosaurs in which the unguals are nearly as long as the digits[82], which indicates that portions of both feet are present in AR-1/31. We interpret that the pes of Europelta possesses four pedal phalanges as in most other nodosaurids [80]. Liaoningosaurus has three digits on the pes. The eight similar unguals are interpreted as pedal unguals and the smallest ungual ( ) is interpreted as an isolated manual ungual. The overall proportions of the preserved pedal elements are similar to those of Niobrarasaurus [119], which also has pedal unguals nearly as large as its metatarsals.
Armor
There was an abundance of dermal armor recovered with both AR-1/10 and AR-1/31. On comparison with the quarry maps, none of the osteoderms appears to be preserved in situ with any of the skeletal elements or with each other, and there is no fusion between any of the osteoderms recovered. Therefore, the armor has been divided into several broad morphotypes for the purpose of description and comparison to armor described for other ankylosaurs. Although morphotypes and terminologies have been proposed [129], [130], no system fits for all armor types in all ankylosaurs. A number of researchers have divided armor into types as in Type 1, 2, etc. [131]; for this discussion the armor types are alphabetized to ensure minimal confusion with previous descriptions. The term osteoderm is used to describe relatively larger dorsal and lateral armor elements with the presence of an external keel or tubercle, whereas the term ossicle describes relatively smaller dermal armor lacking a keel, in the sense of Blows [130]. It is recognized that a consistent methodology for describing armor is achievable, but must be done within a phylogenetic framework to be of maximum utility.
Osteoderm surface texture may be broadly useful in differentiating ankylosaurids from nodosaurids [132], [133]. The vast majority of the osteoderms examined in Europelta has a moderately rugose texture with sparse pitting more in keeping with nodosaurids and basal ankylosaurids rather than more derived ankylosaurids. Whereas histological studies have proven useful in the study of thyreophorans [132], [134], [135], that is beyond the scope of this study.
It is noteworthy that no portions of distinct cervical rings were recovered, although cervical vertebrae are known for both skeletons of Europelta. Additionally, only one spine from the cervical or pectoral region was tentatively identified. We postulate that these elements were lost through the process of coal removal or may have been taphonomically removed from the skeletal associations. Only the discovery of additional specimens of Europelta can further reveal the presence of cervical half-rings.
Type A armor
An isolated fragmentary spine (AR-1-128/10), possibly from the cervical or pectoral region, is recognized from the holotype ( ). It appears to represent only the anterior half and may have been cut in two as the overlying coal was removed. This sharp, broken margin reveals an asymmetric, Y-shaped cross-section. The base flares more and is is less excavated than in a Type 2 caudal plate, suggesting that it was positioned on a broad flank of the body. From the possible anterior margin, the spine slopes posteriorly 15 cm to the broken margin in a gradual arc. There is no indication that the spine could not have been longer. The spine is compressed as in the cervical spines of Sauropelta [77], [99] and Edmontonia [110], [136], and the pectoral spines of Gastonia [83] and Polacanthus [7], [10]. The base is asymmetrical in a manner similar to the elongate osteoderms in Mymoorapelta [84], with one side of the base extending lower anteriorly and the other posteriorly. There is no evidence of a basal plate incorporated into fusion of the cervical half-ring as in mature ankylosaurs like Mymoorapelta [84] Gargoyleosaurus [85], [95], Gastonia [83], Polacanthus [10], [130], and Sauropelta [77], [99]. This may relate to the anchoring of larger elements into the dermis in Gastonia and Polacanthus [130]. We tentatively interpret AR-1-128/10 as a pectoral spine. However, if the complete element extends beyond the break for more than twice the length of the preserved portion, it would fall into the category of Type B armor, although that is unlikely because it is more massive form than the Type B elements.
Type B armor
Dorsoventrally compressed, hollow, asymmetric-based plate-like osteoderms with sharp anterior and posterior edges and lateroposteriorly directed apices are identified for AR-1/10 ( ) and AR-1/31 ( ). Similar large osteoderms have been described as caudal plate ostederms in Mymoorapelta [84], [109], Gargoyleosaurus [85], [95], Gastonia [83], and Polacanthus [8]-[10], [38], [130]. Similar, more anterorposteriorly symmetrical caudal plate osteoderms are also known in Minmi [137], [138] and several Asian ankylosaurids [131]. The few plate-like osteoderms of this morphology that are identified in Europelta are mediolaterally shorter and anteroposteriorly longer with a more posteriorly swept apices. Two pairs of similar plates are known for the holotype of Sauropelta (AMNH 3032), with one of the larger plates being illustrated [99]. One plate from the Yale collections of Sauropelta has a unique double apex (YPM 5490). Given the rarity of Type B armor in Sauropelta and Europelta we hypothesize that caudal plates in these nodosaurids ran down the sides of the tail but decreased in size more rapidly, such that long-keeled osteoderms of Type E morphology made up the lateral armor down most of the length of the tail. It is also possible that these large plate-like osteoderms were on the lateral margin of the sacrum as has been documented by Carpenter and others [106] in Saichania. Struthiosaurus preserves several osteoderms of this morphology that have been reconstructed as in Polacanthus as being medial, dorsally-projecting caudal osteoderms [25], [26]. The relative rarity of these plate-like osteoderms suggests that they were restricted to the base of the tail as well.
Type C armor
Both AR-1/10 ( ) and AR-1/31 ( ) preserve fairly large (∼15–25 cm long) subrectangular to subtrapezoidal, solid osteoderms with low, evenly developed keels running down the long axis of the osteoderm either medially or to one side of the mid-line. Their distal and medial surfaces are subparallel and the entire plate may be slightly flexed across the short axis perpendicular to the crest. The straight, longer margins of these plates appear to have been tightly affixed but not fused to adjoining osteoderms. Armor of Type C morphology is not common but is most similar to medial cervical osteoderms of half-rings, and most distinctively, across the mid-line of the pectoral region in some nodosaurids such as Stegopelta [138], Niobrarasaurus [140], [141], Panoplosaurus [74], [101], and Edmontonia [74], [110].
Type D armor
Both AR-1/10 and AR-1/31 preserve large (∼10-20 cm long) asymmetric, diamond ( ; ) to tear-drop shaped ( ) osteoderms with a long keel rising to an apex medially to posteriorly and in some specimens extending past the posterior margin of the base. They are distinguished from Type E osteoderms because they are wider than 50% of their length. The wider osteoderms are thinner and more solid than the narrower osteoderms with small pockets under the apices. The more diamond-shaped forms may be more closely appressed to each other in anterior bands similar to Type C armor.
Type D Armor is widely known in the nodosaurids such as Sauropelta [99], Panoplosaurus [101], and Edmontonia. Gastonia is documented to have similar armor [142], although more solid in cross section with less basal excavation, which occurs in oblique rows anterior to the sacrum with each osteoderm separated by a single row of small Type H ossicles. This pattern is similar to the dorsal dermal ornamentation documented for the ankylosaur Tarchia by Arbour and others [130], except that in Tarchia most of the intermediate scales lacked ossified cores. Similar armor is known from the lateral sides of the legs in some ankylosaurs such as Saichania [106].
Type E armor
Both AR-1/10 and AR-1/31 preserve large (10-15 cm long) moderately asymmetric osteoderms more than twice as long as wide with a long keel higher on the assumed posterior end ( ; ). These osteoderms have proportionally more deeply excavated bases than Type D armor, have chevron-shaped cross-sections, and are distinguished from Type D armor by their width being less than 50% of the length. Type E armor is gradational with Type D armor ( ; ) and may represent lateral or distal armor from the trunk of the body and along the sides of the tail. This armor type is present in Sauropelta [99] and Texasetes [115]. Similar armor is present on the sides of the limbs in Scelidosurus and Saichania [106].
Type F armor
Medium to large (∼5-15 cm long) oval to circular osteoderms of low profile with a median keel extending into an apex near or overhanging the posterior margin of the osteoderm are represented in both AR-1/10 ( ) and AR-1/31( ). The basal surface of the osteoderm is generally solid except for a small pocket under the apex, reminiscent of Type D armor. Less commonly, the base may be more extensively excavated. Armor of this morphology is abundant in many nodosaurids and makes up the major elements of the armor of Sauropelta anterior to the sacrum in AMNH 3036 [142] and is present in Panoplosaurus [101]. These osteoderms may reside within more expansive spaces among the larger dorsal armor as in Edmontonia (AMNH, 5665) and the polacanthids [81], [82], [93], [107], or may be major armor elements on the posterior portion of the sacrum as in Sauropelta (AMNH 3036). They may also lie on the tail between the Type B caudal plate-like osteoderms, or could be arranged along the lateral side of the limbs as in Saichania [106].
Type G armor
One piece (AR-1-192/10) of flat, oval to subtriangular armor (AR-1-192/10) from AR-1/10 is about 12 cm long and 7 cm wide and is about 0.5 cm thick throughout ( ). A pair of similar, osteoderms from the Sauropelta specimen AMNH 3032 was curated with a note from the collector, Barnum Brown, stating that these distinct osteoderms were associated with the forelimbs. Therefore, we suggest a similar position for Type G armor in Europelta.
Type H armor
Small (∼1-4 cm long) solid ossicles are abundant, with 71 examples from both AR-1/10 ( ) and AR-1/31 ( ) illustrated. These ossicles range in shape from round, to oval and even irregularly shaped, and are probably filling in the spaces between larger osteoderms. Small interstitial ossicles are not known for every ankylosaur taxon, but appear to be present in many nodosaurid taxa such as Sauropelta [99], [143] and Edmontonia [74], [136], in polacanthid ankylosaurs such as Gastonia [83] and in some ankylosaurids such as Tarchia [131], in which epidermal scales interstitial to osteoderms do not preserve deeper, interstitial ossicles. Their absence may be real, in that they never form deep to the epidermal scales, taphonomic, in that they are selectively transported away because of their small size and low density, or ontogenetic; in that they only ossify late in ontogeny. The surface texture of Gastonia ossicles is smoother than those of Europelta.
Sacral armor
Armor is present on the posterior margin of the ilium AR-1-479/10. It is composed of large, subequal-sized (7-10 cm) osteoderms that are tightly sutured together ( ) as in the poorly known Stegopelta [139], Nodosaurus [113], Aletopelta [127], and Glyptodontopelta [132], [144]. These low-relief ossicles lack a central apex or keel. The boundary between the margins of the osteoderms and the area devoid of osteoderms on the ilium is sharply demarcated along the margins of unbroken osteoderms, suggesting the armor was not coossified as in Aletopelta [127] and unlike the fully fused sacral armor in the polacanthids Polacanthus and Gastonia [63], [83]. This form of pelvic armor fits that of Arbour and others' Category 3 pelvic armor [121].
Additionally, there is a unique osteoderm AR-1-653/10 that has a large, posteriorly-curved, plate-like keel extending out from the surface that, considered in isolation, is comparable in size and morphology to Type B armor ( ). The base is smooth and gently convex, suggesting it may have been closely appressed to the more anterior portion of the ilium. In overall morphology, this large osteoderm is comparable to the spine-bearing armor plate-like osteoderm identified in Hungarosaurus and interpreted to be present in Struthiosaurus [33].
Unique armor pieces
Some irregularly shaped armor specimens are not represented by more than one element among this material or in the armor from other taxa. At this time, we can offer no positional interpretation of this armor. AR-1-447/10 is an irregular mass of what we interpret as an osteoderm, although it could be sacral armor ( ). AR-1-438/10 is a small, cap-shaped shaped with a small excavation in the center of the external surface ( ). Two small, deeply basally excavated, oval osteoderms ( ) were collected from AR-1/31(AR-1-3239/31, 3721). These osteoderms lack the external excavation.
Discussion
Europelta ( ) can be distinguished from any of the ankylosaurs assigned to the Polacanthidae (sensu Kirkland's Polacanthinae [83] and Carpenter's Polacanthidae [63] from the Upper Jurassic and Lower Cretaceous as defined by Yang and others [64]; see Terminology) by its rounded, tear-drop shaped skull and a suborbital horn developed on the posterior portion of the jugal and the quadratojugal posterior to the orbit, as opposed to a triangular-shaped skull that is widest at the posterior margin and a suborbital horn developed exclusively on the jugal (as seen in polacanthids). Post-cranially, it can also be distinguished from polacanthids, by its elongate lower hind limbs, the apparent rarity of cervical, pectoral, and thoracic spines, and reduction in the number of caudal plate-like osteoderms. Likewise, it has an abundance of Type D, asymmetric, tear-drop shaped osteoderms like those observed in many nodosaurids and absent in all polacanthids.
Europelta is also distinguished from derived ankylosaurids by its weakly ornamented teardrop-shaped skull in which the lower temporal opening is visible in lateral view. The absence of a tail club also distinguishes the taxon from these ankylosaurids. More basal “shamosaurine grade” ankylosaurids [63], [86] are more similar to Europelta, but also have the lower temporal openings completely obscured laterally by expanding the lateral margin of their skulls. “Shamosaurine grade” ankylosaurids also possess skulls that are approximately as wide mediolaterally between the orbits as they are across the posterior margin.
Europelta shares a number of derived characters with nodosaurids [71], [72], [83], [94], [114]. It has a tear-drop shaped skull that is longer than wide with its greatest width dorsal to the orbits, whereas the short, boxy skulls of Minmi and all anklosaurids are essentially as wide at the posterior edge of the skull, as are the elongate skulls of “shamosaurine-grade” ankylosaurids. Grooves in the remodeled textured skull roof define epidermal scale impressions, with the largest covering the frontoparietal area. Although poorly preserved, the laterally extensive pterygoids are pressed up against the anterior face of the braincase. All known nodosaurid scapulae have a prominent acromion process extending on to the blade of the scapula that terminates in an expanded knob. Unfortunately, this portion of the scapula is as yet unknown in Europelta.
Some character states considered typical of nodosaurids are absent in Europelta. Instead of having a distinct hourglass-shaped palate typical of nodosaurids [70], [71], [82], [83], [114], the upper tooth rows show less lateral emargination and diverge posteriorly. This is also true of Silvisaurus, which also shares an expanded lateral wall of the skull [76], [77]. The coracoid of Europelta is nearly as long as it is tall, whereas in other nodosaurids, for which the corocoid is known, it is expanded anteriorly and longer than tall [71], [72], [83], [94], [114].
The only other Early Cretaceous nodosaurid to have large cranial scales as in Europelta is Propanoplosaurus, known only from an embryonic to hatchling specimen from the base of the Potomac Group of Maryland [145]. However, only the anterior cranial scales are well defined in Propanoplosaurus, whereas only the posterior scale pattern in Europelta. The unusual preservation and extremely small size of Propanoplosaurus lead us to suspect that the fossil preserves the actual scales overlying the skull and not the remodeled skull roof, because this is such a young specimen and remodeling of the cranial bones is not expected to have occurred so early in ontogeny [129], [146].
Additionally, a number of important characters traditionally used to define nodosaurids are not known in Europelta, as yet, because of the missing anteroventral half of the scapula and the absence of premaxilla and surangulars. Thus, the presence absence of premaxillary teeth, if the tooth row joined the margin of premaxillary beak, the morphology of the naris, the height of the coronoid process, and the morphology of the acromion process are unknown for Europelta.
Europelta is distinguishable from European nodosaurids from the Albian through the Cenomanian. The juvenile Anoplosaurus from the Albian Gault Clays of southern England differs in a number of characters, such as possessing a proportionally longer coracoid, a narrower proximal end of the humerus, and a femur with a separate anterior trochanter [17] although the latter two characters are consistent with the juvenile nature of Anoplosaurus. No pectoral spines of the morphology described for “Acanthopholis” from the Cenomanian Lower Chalk in southern England by Huxley [13] are known in Europelta. Additionally, the tall teeth assigned to “Acanthopholis” are distinct in the long apicobasal ridges extending from the denticles to the root on medial and lateral faces of the teeth, and in the presence of caudal ribs that extend laterally and flex ventrally, whereas the caudal ribs in Europelta extend ventrolaterally and flex laterally [16], [17]. Europelta is like other Late Cretaceous European nodosaurids in having a short symphysis for the predentary, a mediolaterally wide and anteroposteriorly thin quadrate, an anteroposteriorly arched sacrum, and a straight ischium [21], [32].
The domed skull and elongate cervical vertebrae in Struthiosaurus clearly distinguish it from Europelta. Likewise, Hungarosaurus also has more elongate cervical vertebrae [32]. Both Hungarosaurus and Struthiosaurus possess a pair of spines on the anterior portion of the pelvis [33], whereas we interpret the presence of a pair of upright plate-like armor elements in this position in Europelta ( ).
The lateral wall of the skull in most North American nodosaurids is typically narrow [82], whereas in Europelta it is relatively wider, although a broad notch along its posterior margin permits the caudal margin of the lower temporal opening to be observed in lateral view. This morphology in Europelta is similar to that in the nodosaurids Silvisaurus [76], [77] and Peloroplites [86]. Although, the skull of Struthiosaurus transylvanicus is highly reconstructed [22], it appears that the lateral wall of the skull is expanded laterally, whereas not completely obscuring the lower temporal opening. This character state is not known in other species of Struthiosaurus, but appears to be moderately developed in Hungarosaurus [32].
Comparisons of Europelta with the Asian”nodosaurids” Zhongyuansaurus [93] and Zhejiangosaurus [126] from the lower Upper Cretaceous of China hinges partially on the question of whether those taxa have been validly referred to Nodosauridae. Carpenter and others [86] noted that the skull of Zhongyuansaurus is morphologically similar to that of a “shamosaurine-grade” (like Shamosaurus and Gobisaurus) ankylosaurids and was the first shamosaurine-grade ankylosaurid documented to not have a tail club. However, its distal tail is modified into a stiffened structure of the same morphology as the “handle” of the tail club in more derived ankylosaurids [147], [148]. Zhejiangosaurus was assigned to the nodosaurids based on characteristics of the femur and sacrum, together with the lack of a tail club [126]. We hypothesize that it lacked a knob as in basal ankylosaurids, polacanthids and nodosaurids because ankylosaurids with a full tail club have distal free caudal vertebrae bearing caudal ribs at the base of the handle. Most of the distal caudal vertebrae of Zhejiangosaurus have raised ridges on the sides of the centra as in the distal vertebrae of polacanthids and nodosaurids. Additionally, whereas the position of its most proximal preserved caudal vertebrae is not known, morphologically, they do not appear to represent the most proximal caudal vertebrae. Thus, while Zhejiangosaurus' 13 preserved caudal vertebra are more than the number of free caudals preserved in most ankylosaurs with tail clubs (10 in Saichania [106] and Dyoplosaurus [148]), the total number of free caudals in its tail would appear to be more than the 14 in Tarchia [130] and 15 in Pinacosaurus [129]. Unlike nodosaurids, Zhejiangosaurus has an exceedingly low ratio of femur to tibia length of 0.46 similar to that of with ankylosaurids and polacanthids rather than nodosaurids. Dongyangopelta [149] was described as a second nodosaurid from the same area and stratum as Zhejiangosaurus, which was found to be its sister taxon in their phylogeny [149]. With few overlapping elements, we feel that the proposed differences between these taxa may be due to preservation, individual variation, or ontogeny. Additionally, given the presence of a pelvic shield and numerous caudal plate-like osteoderms in Dongyangopelta, we suggest that both specimens may pertain to the same taxon and represent the first polacanthid described from Asia. Given the recent description of the polacanthid Taohelong from the upper portion of the Lower Cretaceous of Gansus Province in western China [64], this hypothesis has added support. We also do not think that the partial ankylosaur skull reported from the lower Upper Cretaceous of Hokkaido, Japan [150] can be diagnosed as a nodosaurid with any confidence at this time, due to the incomplete nature of the specimen. Thus, we do not presently recongnize the presence of true nodosaurids in Asia.
In his seminal paper defining a bipartite division of the Ankylosauria into Ankylosauridae and Nodosauridae, Coombs [71] hypothesized that Acanthopholis (as a nomen dubium in which he would have included Anoplosaurus) and Struthiosaurus might represent a separate lineage of European nodosaurids. Unlike Hylaeosaurus (in which he included Polacanthus), these taxa had a well-developed supraspinus fossa developed anteriorly on the scapula as did all North American nodosaurids. This European lineage was hypothesized based on their small body size, presence of premaxillary teeth, and their possessing an unfused scapula and corocoid. Although, none of the characters are valid in defining such a group, our research on Europelta has resulted in supporting the taxonomic hypothesis of Coombs [71], [72] as correct, just for the wrong reasons.
Relationships to Other Taxa
We use Struthiosaurinae to define the clade of European nodosaurs. Nopcsa [25] proposed Acanthopholidae as a family of relatively lightly built thyreophorans, that included Acanthopholis ( = Anoplosaurus), Polacanthus, Stegopelta, Stegoceras, and Struthiosaurus. In 1923, he divided the Acanthopholidae into an Acanthopholinae and a Struthiosaurinae without comment [69]. Subsequently, he relegated the Acanthopholidae to a subfamily of the Nodosauridae, in which he also included Ankylosaurus and restricted the Acanthopholinae to Acanthopholis, Hylaeosaurus, Rhodanosaurus, Struthiosaurus, Troodon [26], [151]. This artificial grouping included a polacanthid ankylosaur [72], [83], a pachycephalosaur [152] and Acanthopholis, now considered a nomen dubium [17], [82]. Thus, the term Acanthopholinae is not acceptable for this newly recognized clade of nodosaurids. Thus, Struthiosaurinae is the next published term available to use for this clade and is derived from the first described and youngest member of this clade. Struthiosaurinae is defined as the most inclusive clade containing Europelta but not Cedarpelta, Peloroplites, Sauropelta or Edmontonia.
In order to determine the systematic position of Europelta, it was found that previous cladistic analyses [71], [72], [82], [83], [114], did not include many of the character states that we identify as significant in our research on Upper Jurassic and Lower Cretaceous ankylosaurs. A major weakness of these analyses is the limited recognition of postcranial skeletal and dermal characters that restricts the testing the phylogenetic relationships for taxa for which skulls are either poorly known or not known at all.
We present a character based definition of Struthiosaurinae as: nodosaurid ankylosaurs that share a combination of characters including: narrow predentary; a nearly horizontal, unfused quadrate that is oriented less than 30° from the skull roof, and mandibular condyles that are 3 times transversely wider than long; premaxillary teeth and dentary teeth that are near the predentary symphysis; dorsally arched sacrum; an acromion process dorsal to midpoint of the scapula-coracoid suture; straight ischium, with a straight dorsal margin; relatively long slender limbs; a sacral shield of armor; and erect pelvic osteoderms with flat bases. This suite of characters unites Europelta with the European nodosaurids Anoplosaurus, Hungarosaurus and all species assigned to Struthiosaurus. This clade of European nodosaurids has not been previously recognized. Europelta represents the earliest member of the European clade Struthiosaurinae.
Biogeogeographic Implications
The near simultaneous appearance of nodosaurids in both North America and Europe is worthy of consideration ( ). Europelta is the oldest nodosaurid known in Europe, it derived from strata in the lower Escucha Formation that is dated to early Albian. The oldest nodosaurid from western North America is Sauropelta, which in the lower part of its range is in the lower Albian Little Sheep Mudstone Member (B interval) of the Cloverly Formation in northern Wyoming and southern Montana [99], [153] with an ash bed 75 meters above the base near the top of the member providing an age of 108.5±0.2 Ma [154]. Nodosaurid remains from eastern North America appear to be older. Teeth of a large nodosaurid Priconodon crassus are known from the Arundel Clay of the Potomac Group [77], [155], which palynology dates as near the Albian-Aptian stage boundary [156]. The hatchling Propanoplosaurus is from the base of the underlying Patuxent Formation of the Potomac Group of Maryland, which has been dated as late Aptian [157], [158], making Propanoplosaurus the oldest known nodosaurid. Polacanthid ankylosaurs characterize pre-Aptian faunas in both Europe [11], [12], [37]-[39] and North America [70], [95], [159]. We have not been able to document a specific example of Polacanthus in the Lower Aptian Vectis Formation of the Wealden Group, although Polacanthus has been reported to occur in those strata [10]-[12], [82], [160]. However, polacanthids are present in the lower Aptian Morella Formation of northeastern Spain [40]. Blows [10] illustrated a block with ankylosaur dorsal vertebrae with the uninformative ventral portion of a pelvic shield fragment and noted it as being from Charmouth, suggesting that there were upper Albian polacanthids in England [160]. However, the specimen NMW 92.34G.2 was actually found on the beach further west at Charton Bay and may have come from either the Aptian (Lower Greensand) or Albian (Upper Greensand). Only preparation of the dorsal surface of the pelvic shield would reveal if the specimen is a polacanthid or nodosaurid. A large polacanthid (BYU R254) occurs in the Poison Strip Sandstone Member of the Cedar Mountain Formation [156]. It is not a nodosaurid close to Sauropelta as reported by Carpenter and others [97], but a polacanthid that was initially described as cf. Hoplitosaurus [161]. These rocks have been dated as lower to middle Aptian by laser ablation of detrital zircons and by U-Pb dating of early diagenetic carbonate [162]. A fragmentary large nodosaurid with massive cervical spikes that may be referred to as cf. Sauropelta (DMNS 49764) has been recovered from the overlying Ruby Ranch Member about 20 m up section in the same region [163] in strata interpreted to be of Lower Albian age [162]. Thus, the youngest polacanthids occur in the lower to possibly mid-Aptian and the oldest documented nodosaurids occur in the upper Aptian or lower Albian in both Europe and North America with no discernible s
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Agnatha
Genus: Nova (1)
Stephaspis GAI & ZHU, 2007
Species: Nova (2)
Gigantaspis minima PERNEGRE & GOUJET, 2007
Stephaspis dipteriga GAI & ZHU, 2007
Synonym: Nova (1)
Gigantaspis laticephala (BLIECK & GOUJET, 1983) PERNEGRE & GOUJET, 2007
= Zascinaspis laticephala BLIECK & GOUJET, 1983
Placoderm
Genus: Nova (2)
Erikaspis DUPRET, GOUJET & MARK-KURIK, 2007
Materpiscis LONG, TRINAJSTIC, YOUNG & SENDEN, 2007
Species: Nova (1)
Materpiscis attenboroughi LONG, TRINAJSTIC, YOUNG & SENDEN, 2007
Synonym: Nova (1)
Erikaspis zychi (STENSIO, 1945) DUPRET, GOUJET & MARK-KURIK, 2007
= Kujdanowiaspis zychi STENSIO, 1945 (nomen nudum)
Blom, H., Clack, J. A., Ahlberg, Per E., and Friedman, M., 2007, Devonian vertebrates from East Greenland: a review of faunal composition and distribution: Geodiversitas, v. 29, n. 1, p. 119-141. Link to pdf
Dupret, V., Goujet, D., and Mark-Kurik, E., 2007, A new genus of placoderm (Arthrodira: 'Actinolepida') from the Lower Devonian of Podolia (Ukraine): Journal of Vertebrate Paleontology, v. 27, n. 2, p. 266-284. (Erikaspis zychi = Kujdanowiaspis zychi)
Janvier, P., Desbiens, S., and Willett, J. A., 2007, New evidence for the controversial "Lungs" on the Late Devonian antiarch Bothriolepis canadensis (Whieteaves, 1880) (Placodermi: Antiarcha): Journal of Vertebrate Paleontology, v. 27, n. 3, p. 709-710.
Long, J. A., Trinajstic, K., Young, G. C., and Senden, T., 2007, Live birth in the Devonian Period: Nature, v. 453, p. 650-652. (Materpiscis attenboroughi)
Randon, C., Derycke, C., Blieck, A., Perri, M. C., and Spalletta, C., 2007, Late Devonian-Early Carboniferous vertebrate microremains from the Carnic Alps, northern Italy: Geobios, v. 40, p. 809-826.
Suarez Soruco, R., 2007, Bolivia y su paleobiodiversidad: In: 4th European Meeting on the Paleontology and Stratigraphy of Latin America, edited by Diaz-Martinez, E., and Rabano, I., Cuademos del Museo Geomienro, n. 8, Instituto Geologico y Minero de Espana, Madrid, 2007, p. 375-382.
Trinajstic, K., and Hazelton, M., 2007, Ontogeny, phenotypic variation and phylogenetic implications of arthrodires from the Gogo Formation, Western Australia: Journal of Vertebrate Paleontology, v. 27, n. 3, p. 571-583.
Trinajstic, K., Marshall, C., Long, J., and Bifield, K., 2007, Exceptional preservation of nerve and muscle tissues in Late Devonian placoderm fish and their evolutionary implications: Biology Letters, v. 3, p. 197-200.
Wilson, M. V. H., Hanke, G. F., and Marss, T., 2007, Paired fins of jawless vertebrates and thier homologies across the "Agnathan"- Tnathostome transition: In: Major transitions in vertebrate evolution, edited by Anderson, J. S., and Sues, H.-S., p. 122-149.
Fish
Genus: Nova (7)
Australopachycormus KEAR, 2007
Baoqingichtys WANG, JIN, WANG & ZHU, 2007
Paraperleidus ZHAO & LU, 2007
Prosantichthys ARRATIA & HERZOG, 2007
Sangiorgioichthys TINTORI & LOMBARDO, 2007
Tycheroichthys HAY, CUMBAA, MURRAY & PLINT, 2007
Zhejiangichthys WANG, JIN, WANG & ZHU, 2007
Species: Nova (7)
Australopachycormus hurleyi KEAR, 2007
Baoqingichtys microdontus WANG, JIN, WANG & ZHU, 2007
Paraperleidus changxingensis ZHAO & LU, 2007
Prosantichthys buergini ARRATIA & HERZOG, 2007
Sangiorgioichthys aldae TINTORI & LOMBARDO, 2007
Tycheroichthys dunveganensis HAY, CUMBAA, MURRAY & PLINT, 2007
Zhejiangichthys zhaoi WANG, JIN, WANG & ZHU, 2007
Alvarado-Ortega, J., Espinosa-Arrubarrena, L., Blanco, A., Vega, F. J., Benammi, M., and Briggs, D. E. G., 2007, Exceptional preservation of soft tissues in Cretaceous fishes from the Tlayua Quarry, Centarl Mexico: Palaios, v. 22, p. 682-685.
Azevedo, R. P. F. de, Vasconcellos, P. L. de, Canderio, C. R. dos A., and Bergqvist, L. P., 2007, Restos microscopicos de vertebrados fosseis do Grupo Bauru (Neocertaceo), no oest do estado de Sao Paulo, Brasil: In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 533-541.
Arratia, G., and Herzog, A., 2007, A new halecomorph fish from the Middle Triassic of Switzerland and its systematic implications: Journal of Vertebrate Paleontology, v. 27, n. 4, p. 838-849. (Prosantichthys buegini)
Arratia, G,. and Schultz, H.-P., 2007, Eurycormus - Eurypoma, two Jurassic actinopterygia genera with mixed identity: Fossil Record, v. 10, n. 1, p. 17-31.
Blanco-Pinon, A., and Alvarado-Ortega, J., 2007, Review of Vallecillichthys multivertebratum (Teleostei: Ichthyodectiformes), a Late Cretaceous (early Turonian) "Bulldog fish" from northeastern Mexico: Revista Mexicana de Ciencias Geologicas, v. 24, n. 3, p.450-466. Link to pdf
Blom, H., Clack, J. A., Ahlberg, Per E., and Friedman, M., 2007, Devonian vertebrates from East Greenland: a review of faunal composition and distribution: Geodiversitas, v. 29, n. 1, p. 119-141. Link to pdf
Brito, P. M., 2007, The Crato Formation fish fauna: In: The Crato Fossil Beds of Brazil, window into an Ancient World, edited by Martin, D. M., Bechly, G., and Loveridge, R. F., p. 429-443.
Cavin, L., and Forey, P. L., 2007, Using ghost lineages to identify diversification events in the fossil record: Biology Letters, v. 3, p. 201-204.
Cavin, L., Suteethorn, V., Buffetaut, E., Claude, J., Cuny, G., Le Loeuff, J., and Tong, H., 2007, The first sinamiid fish (Holostei, Halecomorpha) from Southeast Asia (Early Cretaceous of Thailand): Journal of Vertebrate Paleontology, v. 27, n. 4, p. 827-837. (Siamamia naga)
Chahud, A., Fairchild, T. R., 2007, Vertebrados paleozoicos do estado de Sao Paulo: In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 101-110.
Dietze, K., 2007, Redescription of Dastilbe crandalli (Chanidae, Euteleostei) from the Early Cretaceous Crato Formation of north-eastern Brazil: Journal of Vertebrate Paleontology, v. 27, n. 1, p. 8-16.
Dzik, J., and Sulej, T., 2007, A review of the Early Late Triassic Krasiejow Biota from Silesia, Poland: Palaeontologica Polonica, n. 64, p. 1-27.
Everhart, M. J., 2007, Remains of a pycnodont fish (Actinopterygii: Pycnodontiformes) in a coporlite; an uppermost record of Micropycnodon kansasensis in the Smoky Hill Chalk, western Kansas: Transcations of the Kansas Academy of Science, v. 110, n. 1/2, p. 35-43.
Forey, P. L., and Cavin, L., 2007, A new speices of Cladocyclus (Teleostei: Ichthyodectiformes) from the Cenomanian of Morocco: Palaeontologia Electronica, n. 10.3.12a, 10 pp. (Cladocyclus pankowskii)
Fürsich, F. T., Mäuser, M., Schneider, S., and Werner, W., 2007, The Wattendorf Plattenkalk (Upper Kimmeridgian) - a new conservation lagerstätte from the northern Franconian Alb, southern Germany: Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 245, n. 1, p. 45-58.
Gallo, V., 2007, Parsimony analysis of endemicity of enchodontoid fishes from the Cenomanian: Notebooks on Geology, Letter 2007/01 (CG2007_L01), 8pp.
Garrison jr, J. R., Brinkman, D., Nichols, D. J., Layer, P., Burge, D., and Thayn, D., 2007, A multidisiplinary study of the Lower Cretaceous Cedar Mountain Formation, Mussentuchit Wash, Utah: a determination of the paleoenvironment and paleoecology of the Eolambia caroljonesa dinosaur quarry: Cretaceous Research, v. 28, p. 461-494.
Hay, M. J., Cumbaa, S. L., Murray, A. M., and Plint, A. G., 2007, A new paraclupeid fish (Clupeomorpha, Ellimmichthyiformes) from a muddy marine prodelta environment: middle Cenomanian Dunvegan Formation, Alberta, Canada: Canadian Journal of Earth Sciences, v. 44, p. 775-790. (Tycheroichthys dunveganensis)
Ifrim, C., Stinnesbeck W., and Frey, E., 2007, Upper Cretaceous (Cenomanian-Turonian and Turonian-Coniacian) open marine plattenkalk deposits of New Mexico: Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 245, n. 1, p. 71-81.
Ivanov, A., and Tatyanaklets, 2007, Triassic fishes from Siberia, Russia: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural & Science, Bulletin 41, p. 108-109.
Janvier, P., 2007, Homologies and evolutionary transitions in early vertebrate history: In: Major transitions in vertebrate evolution, edited by Anderson, J. S., and Sues, H.-S., p. 56-121.
Jurkovsek, & Kolar-Jurkovsek, T., 2007, Fossil assemblages from the Upper Cretaceous Komen and tomaj Limestones of Kras (Slovenia): Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 245, n. 1, p. 83-92.
Kear, B. P., 2007, First record of a pachycormid fish (Actinopterygii: Pachycormiformes) from the Lower Cretaceous of Australia: Journal of Vertebrate Paleontology, v. 27, n. 4, p. 1033-1038. (Australopachycormus hurleyi)
Lopez-Arbarello, A., and Codorniu, L., 2007, Semionotids (Neopterygi, semionotiformes) from the Lower Cretaceous Lagarcito Formation, San Luis Province, Argentina: Journal of Vertebrate Paleontology, v. 27, n. 4, p. 811-826.
Lucas, S. G., Heckert, A. B., Spielmann, J. A., Tanner, L. H., and Hunt, A. P., 2007, Second Day: Early and Middle Triassic stratigraphy, sedimentology and paleontology in northeastern Arizona: In: Triassic of the American West, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural History & Science, Bulletin 40, p. 181-187.
Lucas, S. G., Heckert, A. B., Spielmann, J. A., Tanner, L. H., and Hunt, A. P., 2007, Third Day: Triassic stratigraphy and paleontology in northeastern Arizona: In: Triassic of the American West, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural History & Science, Bulletin 40, p. 186-197.
Lucas, S. G., and Tanner, L. H., 2007, Tetrapod biosratigraphy and biochronology of the Triassic-Jurassic transition on the southern Colorado Plateau, USA: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 244, p. 242-256.
Martill, D. M., 2007, The age of the Cretaceous Santana Formation fossil konservat lagerstätte of north-east Brazil: a historical reivew and an appraisal of the biochronostratigraphic utility of its palaeobiota: Creaceous Research, v. 28, p. 895-920.
Martill, D. M., Bechly, G, and Heads, S. W., 2007, Appendix: species list for the Crato Formation: In: The Crato Fossil Beds of Brazil, window into an Ancient World, edited by Martin, D. M., Bechly, G., and Loveridge, R. F., p. 582-607.
Milner, A. R. C., and Kirkland, J. I., 2007, The case for fishing dinosaurs at the St. George dinosaur discovery site at Johnson Farm: Utah Geological Survey, Survey Notes, v. 39, n. 3, p. 1-3.
Newbrey, M. G., Wilson, M. V. H., and Ashworth, A. C., 2007, Centrum growth patterns provide evidence for two small taxa of Hiodontidae in the Cretaceous Dinosaur Park Formation: Canadian Journal of Earth Sciences, v. 44, p. 721-732.
Oheim, K. B., 2007, Fossil site predicition using geographic information systems (GIS) and suitability analysis: the Two Medicine Formation, MT, a test case: Paleogeography, Palaeoclimatology, Palaeoecology, v. 251, p. 354-365.
Parris, D. C., Grandstaff, B. S., and Gallagher, W. B., 2007, Fossil fish from the Pierre Shale Group (Late Cretaceous): clarifying the biostratigraphic record: In: The Geology and Paleontology of the Late Cretaceous marine deposits of the Dakotas, edited by Martin, J. E., and Parris, D. C., The Geological Society of America, Special Paper 427, p. 99-109.
Randon, C., Derycke, C., Blieck, A., Perri, M. C., and Spalletta, C., 2007, Late Devonian-Early Carboniferous vertebrate microremains from the Carnic Alps, northern Italy: Geobios, v. 40, p. 809-826.
Sanchez-Hernandez, B., Benton, M. J., and Naish, D., 2007, Dinosaurs and other fossil vertebrates from the Late Jurassic and Early Cretaceous of the Galve area, NE Spain: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 249, p. 180-215.
Schein, J. P., and Lewis, R. D., 2007, Actinopterygian fishes from Upper Cretaceous rocks in Alabama, with emphasis on the Teleostan genus Enchodus: Paludicola, v. 6, n. 2, p. 41-86.
Tintori, A., and Lombardo, C., 2007, A new early Semionotidae (Semionotiformes, Actinopterygii) from the Upper Ladinian of Monte San Giorgio area (Southern Switzerland and northern Italy): Rivista Italiana di Paleontologia e Stratigrafia, v. 113, n. 3, p. 369-381. (Sangiorgioichthys aldae) Link to pdf
Viohl, G., and Zapp, M,. 2007, Schamhaupten, an outstanding Fossil-Lagerstätte in a silicified Plattenkalk around the Kimmeridgian-Tithonian boundary (Southern Franconian Alb, Bavaria): Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 245, n. 1, p. 127-142.
Wang, N.-Z., Jing, F., Wang, W., and Zhu, X.-S., 2007, Actinopterygian fishes from the Permian-Triassic boundary beds in Zhejiang and the Jiangzi Provinces, South China and fish mass extinction, recovery and radiation: Vertebrata PalAsiatica, v. 45, n. 4, p. 307-329. (Baoqingichthys microdontus, Zhejiangichtys zhaoi)
Wilson, M. V. H., Hanke, G. F., and Marss, T., 2007, Paired fins of jawless vertebrates and thier homologies across the "Agnathan"- Tnathostome transition: In: Major transitions in vertebrate evolution, edited by Anderson, J. S., and Sues, H.-S., p. 122-149.
Zhao, L.-J., and Lu, L.-W., 2007, A new genus of early Triassic perleidid fish from Changzing, Zhejiang, China: Acta Palaeontologica Sinica, v. 46, n. 2, p. 238-243. (Paraperleidus changzingensis)
Blom, H., Clack, J. A., Ahlberg, Per E., and Friedman, M., 2007, Devonian vertebrates from East Greenland: a review of faunal composition and distribution: Geodiversitas, v. 29, n. 1, p. 119-141. Link to pdf
Holland, T., Warren, A., Johanson, Z., Long, J., Parker, K., and Garvey, J., 2007, A new species of Barameda (Rhizodontida) and heterochrony in the Rhizodontid pectoral fin: Journal of Vertebrate Paleontology, v. 27, n. 2, p. 295-315. (Barameda mitchelli)
Janvier, P., Clement, G., and Cloutier, R., 2007, A primitive megalichthyid fish (Sacropterygii, Tetrapodomorpha) from the Upper Devonian of Turkey and its biogeographical implications: Geodiversitas, v. 29, n. 2, p. 249-268. (Sengoerichthys ottoman) Link to pdf
Johanson, Z,. Long, J. A., Talent, J. A., Janvier, P., and Warren, J. W., 2007, New onychodontiform (Osteichthyes; Sacropterygii) from the Lower Devonian of Victoria, Australia: Journal of Paleontology, v. 81, n. 5, p. 1031-1043. (Bukkanodus jesseni)
Laurin, M., Meunier, F. J., Germain, D., and Lemoine, M., 2007, A microanatomical and histological sudy of the paired fin skeleton of the Devonian sacropterygian Eusthenopteron foordi: Journal of Paleontology, v. 81, n. 1, p. 143-153.
Newman, M. J., and den Blaauwen, J. L., 2007, The synonymy of the Scottish Devonian osteolepid fish Thursius macrolepidotus: Scottish Journal of Geology, v. 43, n. 2, p. 101-106. (Thursius macrolepidotus = Thursius moythomasi) Link to pdf
Witzmann, F., and Schoch, R. R., 2007, A megalichthyid sacropterygian fish from the Lower Permian (Autunian) of the Saar-Nahe Basin, Germany: Geobios, v. 45, p. 241-248. (Palatinicnthys laticeps) Link to pdf
Apesteguia, S., Agnolin, F. L., and Claeson, K., 2007, Review of Cretaceous dipnoans from Argentina (Sarcopterygii: Dipnoi) with descriptions of new species: Rev. Mus. Argentno Cienc. Nat. N. S., v. 9, n. 1, p. 27-40. (Chaoceratodus portezuelensis, Ameghinoceratodus iherigi, Ceratodus argentinus, Ceratodus kaopen, Ptychoceratodus cionei, Ptychoceratodus wichmanni)
Blom, H., Clack, J. A., Ahlberg, Per E., and Friedman, M., 2007, Devonian vertebrates from East Greenland: a review of faunal composition and distribution: Geodiversitas, v. 29, n. 1, p. 119-141. Link to pdf
Campbell, K. S. W., and Barwick, R. E., 2007, The structure and stratigraphy of Speonesydrion from New South Wales, Australia, and the dentition of primitive dipnoans: Paläontologische Zeitschrift, v. 81, Heft 2, p. 146-159.
Campbell, K. S. W., Barwick, R. E., and Blaauwen, J. L. den, 2007, Structure and function of the shoulder girdle in Dipnoans: new material from Dipterus valenciennesi: Senckenbergiana lethaea, v. 86, n. 1, p. 77-91.
Cavin, L., Suteethorn, V., Buffetaut, E., and Tong, H., 2007, A new Thai Mesozoic lungfish (Sarcopterygii, Dipnoi) with an insight into post-Palaeozoic dipnoan evolution: Zoological Journal of the Linnean Society, v. 149, P. 141-177. (Ferganoceratodus martini)
Chahud, A., Fairchild, T. R., 2007, Vertebrados paleozoicos do estado de Sao Paulo: In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 101-110.
Cione, A. L., Gouirie, S., Goin, F., and Poire, D., 2007, Atlantoceratodus, a new genus of lungfish from the Upper Cretaceous of South America and Africa: Revista del Museo de La Plata, v. 10, p. 1-12. (Atlantoceratodus iheringi = Ceratodus iheringi, Atlantoceratodus madagascariensis = Ceratodus madagascariensis)
Dzik, J., and Sulej, T., 2007, A review of the Early Late Triassic Krasiejow Biota from Silesia, Poland: Palaeontologica Polonica, n. 64, p. 1-27.
Friedman, M., 2007, Cranial structure in the Devonian lungfish Soederberghia groenlandica and its implications for the interrelationships of 'rhynchodipterids': Earth and Environmental Science Transcations of the Royal Society of Edinburgh, v. 98, p. 179-198.
Lucas, S. G., Heckert, A. B., Spielmann, J. A., Tanner, L. H., and Hunt, A. P., 2007, Second Day: Early and Middle Triassic stratigraphy, sedimentology and paleontology in northeastern Arizona: In: Triassic of the American West, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural History & Science, Bulletin 40, p. 181-187.
Lucas, S. G., Heckert, A. B., Spielmann, J. A., Tanner, L. H., and Hunt, A. P., 2007, Third Day: Triassic stratigraphy and paleontology in northeastern Arizona: In: Triassic of the American West, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural History & Science, Bulletin 40, p. 186-197.
Milner, A. R. C., and Kirkland, J. I., 2007, The case for fishing dinosaurs at the St. George dinosaur discovery site at Johnson Farm: Utah Geological Survey, Survey Notes, v. 39, n. 3, p. 1-3.
Newman, M. J., and Blaauwen, J. L. D., 2007, A new dipnoan fish from the Middle Devonian (Eifelian) of Scotland: Palaeontology, v. 50, part 6, p. 1403-1419. (Pinnalongus saxoni)
Coelocanths
Brito, P. M., 2007, The Crato Formation fish fauna: In: The Crato Fossil Beds of Brazil, window into an Ancient World, edited by Martin, D. M., Bechly, G., and Loveridge, R. F., p. 429-443.
Friedman, M., 2007, Styloichthys as the oldest coelacanth: implications for early osteichthyan interrelationships: Journal of Systematic Palaeontology, v. 5, n. 3, p. 289-343.
Lucas, S. G., and Tanner, L. H., 2007, Tetrapod biosratigraphy and biochronology of the Triassic-Jurassic transition on the southern Colorado Plateau, USA: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 244, p. 242-256.
Martill, D. M., 2007, The age of the Cretaceous Santana Formation fossil konservat lagerstätte of north-east Brazil: a historical reivew and an appraisal of the biochronostratigraphic utility of its palaeobiota: Creaceous Research, v. 28, p. 895-920.
Martill, D. M., Bechly, G, and Heads, S. W., 2007, Appendix: species list for the Crato Formation: In: The Crato Fossil Beds of Brazil, window into an Ancient World, edited by Martin, D. M., Bechly, G., and Loveridge, R. F., p. 582-607.
Milner, A. R. C., and Kirkland, J. I., 2007, The case for fishing dinosaurs at the St. George dinosaur discovery site at Johnson Farm: Utah Geological Survey, Survey Notes, v. 39, n. 3, p. 1-3.
Szrek, P., 2007, Coelacanths (Actinistia, Sacropterygii) from the Famennian (Upper Devonian) of the Holy Cross Mountains, Poland: Acta Geologica Polonica, v. 57, p. 403-413. Link to pdf
Viohl, G., and Zapp, M,. 2007, Schamhaupten, an outstanding Fossil-Lagerstätte in a silicified Plattenkalk around the Kimmeridgian-Tithonian boundary (Southern Franconian Alb, Bavaria): Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 245, n. 1, p. 127-142.
Fish Ichnology
Fish Burrows
Fish Coprolites
Everhart, M. J., 2007, Remains of a pycnodont fish (Actinopterygii: Pycnodontiformes) in a coporlite; an uppermost record of Micropycnodon kansasensis in the Smoky Hill Chalk, western Kansas: Transactions of the Kansas Academy of Science, v. 110, n. 1/2, p. 35-43.
Schwanke, C., Souto, P. R. de F., 2007, Coprolitos espiralados da Formacao pedra do fogo, Bacia do Parnaiba: In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 111-120.
Anderson, J. S., 2007, Incorporating ontogeny into the matrix: a phylogenetic evaluation of developmental evidence for the origin of modern amphibians: In: Major transitions in vertebrate evolution, edited by Anderson, J. S., and Sues, H.-S., p. 182-227.
Barycka, E., 2007, Morphology and ontogeny of the humerus of the Triassic temnospondyl amphibian Metoposaurus diagnosticus: Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 243, n. 3, p. 351-361.
Blieck, A., Clement, G., Blom, H., Lelievre, H., Luksevics, E., Streel, M., Thorez, J., and Young, G. C., 2007, The biostratigraphical and palaeogeographical framework of the earliest diversification of tetrapods (Late Devonian): In Devonian Events and Correlations, edited by Becker, R. T., & Kirchgasser, W. T., Geological Society, london, Speical Publications, v. 278, p. 219-235.
Blom, H., Clack, J. A., Ahlberg, Per E., and Friedman, M., 2007, Devonian vertebrates from East Greenland: a review of faunal composition and distribution: Geodiversitas, v. 29, n. 1, p. 119-141.
Carroll, R. L., 2007, The Palaeozoic ancestry of Salamanders, Frogs and Caecilians: Zoological Journal of the Linnean Society, v. 150 (suppl. 1), p. 1-140.
Chahud, A., Fairchild, T. R., 2007, Vertebrados paleozoicos do estado de Sao Paulo: In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 101-110.
Dzik, J., and Sulej, T., 2007, A review of the Early Late Triassic Krasiejow Biota from Silesia, Poland: Palaeontologica Polonica, n. 64, p. 1-27.
Huttenlacker, A. K., Pardo, J. D., and Small, B. J., 2007, Plemmyradytes shintoni, gen. et sp. nov., an Early Permian Amphibamid (Temnospondyli: Dissorophoidea) from the Eskridge Formation, Nebraska: Journal of Vertebrate Paleontology, v. 27, n. 2, p. 316-328. (Plemmyradytes shintoni)
Konietzko-Meier, D., and Wawro, K., 2007, Mandibular dentition in the Late Triassic temnospondyl amphibian Metoposaurus: Acta Palaeontologica Polonica, v. 52, n. 1, p. 213-215.
Langer, M. C., Ribeiro, A. M., Schultz, C. L., and Ferigolo, J., 2007, The continental tetrapod-bearing Triassic of South Brazil: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural & Science, Bulletin 41, p. 201-218.
Lucas, S. G., Heckert, A. B., Spielmann, J. A., Tanner, L. H., and Hunt, A. P., 2007, Second Day: Early and Middle Triassic stratigraphy, sedimentology and paleontology in northeastern Arizona: In: Triassic of the American West, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural History & Science, Bulletin 40, p. 181-187.
Lucas, S. G., Heckert, A. B., Spielmann, J. A., Tanner, L. H., and Hunt, A. P., 2007, Third Day: Triassic stratigraphy and paleontology in northeastern Arizona: In: Triassic of the American West, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural History & Science, Bulletin 40, p. 186-197.
Lucas, S. G., Hunt, A. P., Heckert, A. P., and Spielmann, J. A., 2007, Global Triassic tetrapod biostratigraphy and biochronology: 2007 status: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural and Science, bulletin 41, p. 229-240.
Lucas, S. G., Spielmann, J. A., and Hunt, A. P., 2007, Biochronological significance of the Late Triassic tetrapods from Krasijow, Poland: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural and Science, bulletin 41, p. 248-258.
Markey, M. J., and Marshall, C. R., 2007, Terrestrial-style feeding in a very early aquatic tetrapod is supported by evidence from experimental analysis of suture morphology: Proceedings of the National Academy of Science, v. 104, n. 17, p. 7134-7138.
Milner, A. R., Klembara, J., and Dostal, O., 2007, A zatrachydid temnospondyl from the Lower Permian of the Boskovice furrow in Moravia (Czech Republic): Journal of Vertebrate Paleontology, v. 27, n. 3, p. 711-715.
Moser, M., and Schoch, R., 2007, Revision of the type material and nomenclature of Mastodonsaurus giganteus (Jaeger) (Temnospondyli) from the Middle Triassic of Germany: Palaeontology, v. 50, part 5, p. 1245-1266.
Mueller, B. D., 2007, Koskinonodon Branson and Mehl, 1929, a replacement name for the preoccupied temnospondyl Buettneria Case, 1922: Journal of Vertebrate Paleontology, v 27, n. 1, p. 225. (Koskinonodon perfectus = Buettneria perfecta, Koskinonodon priniceps)
Novikov, I. V., 2007, New data on trematosauroid labyrinthodonts of Eastern Europe: 1. Genus Inflectosaurus Shishkin, 1960: Palaeontological Journal, v. 41, n. 2, p. 167-174.
Pawley, K., 2007, The postcranial skeleton of Trimerorhachis insignis Cope, 1878 (Temnospondyli: Trimerorhachidae): a plesiomorphic temnospondyl from the Lower Permian of North America: Journal of Paleontology, V. 81, n. 5, p. 873-894. Link to pdf
Pineiro, G., Mariscano, C. A., and Damiani, R., 2007, Mandibles of mastodonsaurid temnopsondyls from the Upper Permian-Lower Triassic of Uruguay: Acta Palaeontologica Polonica, v. 52, n. 4, p. 695-703. Link to pdf
Pineiro, G., Marsicano, C., and Lorenzo, N., 2007, A new temnospondyl from the Permo-Triassic Buena Vista Formation of Uruguay: Palaeontology, v. 50, part 3, p. 627-640. (Uruyiella liminea)
Pineiro, G., Marsicano, C. A., Goso, C., and Morosi, E., 2007, Temnospondyl diversity of the Permian-Triassic Colonia Orozco local fauna (Buena Vista Formation) of Uruguay: Revista Brasileria de Paleontologia, v. 10, n. 3, p. 169-180. Link to pdf
Ruta, M., and Coates, M. J., 2007, Dates, nodes and character conflect: addressing the lissamphibian origin problem: Journal of Systematic Palaeontology, v. 5, n. 1, p. 69-122.
Ruta, M., Pisani, D., Lloyd, G. T., and Benton, M. J., 2007, A supertree of Temnospondyli: cladogenetic patterns in the most species-rich group of early-tetrapods: Procedings of the Royal Society, Series B, v. 274, p. 2087-2095.
Schoch, R. R., Fichter, F. M., and Keller, T., 2007, Anatomy and relationships of the Triassic temnospondyl Sclerothroax: Acta Palaeontologica Polonica, v. 52, n. 1, p. 117-136.
Schultz, C. L., and Langer, M. C., 2007, Tetrapodes Triassicos do Rio Grande do Sul, Brasil: In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 269-282.
Shishkin, M. A., 2007, Patterns of recovery of amphibian diversity in the Triassic: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural and Science, bulletin 41, p. 369-370.
Sidor, C. A., Steyer, J., S., and Damiani, R., 2007, Parotosuchus (Temnospondyl: Mastodonsauridae) from the Triassic of Antarctica: Journal of Vertebrate Paleontology, v. 27, n. 1, p. 232-235.
Spielmann, J. A., Lucas, S. G., and Heckert, A. B., 2007, Tetrapod fauna of the UpperTriassic (Revuletian) Owl Rock Formation, Chinle Group, Arizona:In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural and Science, bulletin 41, p. 371-383.
Sulej, T., 2007, Osteology, variability and evolution of Metoposaurus, a temnospondyl from the Late Triassic of Poland: Palaeontologia Polonica, n, 64, p. 29-143.
Urban, M., and Berman, D. S., 2007, First occurrence of the Late Paleozoic amphibian Zatrachys serratus (Temnospondyli, Zatrachydidae) in the Eastern United States: Annals of Carnegie Museum, v. 76, n. 3, p. 157-164.
Warren, A., 2007, New data on Ossinodus pueri, a stem tetrapod from the Early Carbonifeorus of Australia: Journal of Vertebrate Paleontology, v. 27, n. 4, p. 850-862.
Werneburg, R., 2007, Timeless design: colored pattern on skin in Early Permian brachiosaurids (Temnospondyli: Dissorophoidea): Journal of Vertebrate Paleontology, v. 27, n. 4, p. 1047-1050.
Werneburg, R., Ronchi, A., and Schneider, J. W., 2007, The Early Permian branchiosaurids (Amphibia) of Sardinia (Italy): systematic palaeontology, palaeoecology, biostratigraphy and palaeobiogeographic problems: Palaeogeography, Paleoclimatology, Palaeontology, v. 252, p. 383-404.
Werneburg, R., Steyer, J. S., Sommer, G., Gand, G., Schneider, J. W., and Vianey-Liaud, M., 2007, The earliest tupilakosaurid amphibian with diplospondylous vertebrae from the Late Permian of Southern France: Journal of Vertebrate Paleontology, v. 27, n. 1, p. 26-30.
Witzmann, F., 2007, The evolution of the scalation pattern in temnospondyl amphibians: Zoological Journal of the Linnean Society, v. 150, p. 815-834.
Witzmann, F., 2007, A hemivertebra in a temnospondyl amphibian: the oldest record of scoliosis: Journal of Vertebrate Paleontology, v. 27, n. 4, p. 1043-1046.
Witzman, F., and Scholz, H., 2007, Morphometric study of allometric skull growth in the temnospondyl Archegosaurus decheni from the Permian/Carboniferous of Germany: Geobios, v. 40, n. 4, p. 541-554.
Anderson, J. S., 2007, Direct evidence of the rostral anatomy of the Aistopod Phlegethontia, with a new cranial reconstruction: Journal of Paleontology, v. 81, n. 2, p. 408-410.
Carroll, R. L., 2007, The Palaeozoic ancestry of Salamanders, Frogs and Caecilians: Zoological Journal of the Linnean Society, v. 150 (suppl. 1), p. 1-140.
Markey, M. J., and Marshall, C. R., 2007, Terrestrial-style feeding in a very early aquatic tetrapod is supported by evidence from experimental analysis of suture morphology: Proceedings of the National Academy of Science, v. 104, n. 17, p. 7134-7138.
Reisz, R. R., 2007, The cranial anatomy of basal diadectomorphs and the origin of amniotes: In: Major transitions in vertebrate evolution, edited by Anderson, J. S., and Sues, H.-S., p. 228-252.
Ruta, M., and Coates, M. J., 2007, Dates, nodes and character conflect: addressing the lissamphibian origin problem: Journal of Systematic Palaeontology, v. 5, n. 1, p. 69-122.
Saber, H., Wartit, M. El., Hmich, D., and Schneider, J. W., 2007, Tectonic evolution from the Hercynian shortening to the Triassic extension in the Paleozoic sediments of the Western High Atlas (Morocco): Journal of Iberian Geology, v. 33, n. 1, p. 31-40.
Lower Reptiles (“Anapsids”)
Botha, J., Modesto, S. P., and Smith, R. M. H., 2007, Extended procolophonoid reptile survivorship after the end-Permian extinction: South African Journal of Science, v. 103, p. 54-56.
Chahud, A., Fairchild, T. R., 2007, Vertebrados paleozoicos do estado de Sao Paulo: In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 101-110.
Dias-da-Silva, S., Modesto, S. P., and Schultz, C. L., 2006 (published 2007), New material of Procolophon (Parareptilia: Procolophonoidea) from the Lower Triassic of Brazil, with remarks on the ages of the Sanga do Cabral and Buena Vista Formation of South America: Canadian Journal of Earth Sciences, v. 43,p. 1685-1693.
Harris, S. R., Pisani, D., Gower, D. J., and Wilkinson, M., 2007, Investigating stagnation in morphological phylogenetics using consensus data: Systematic Biology, v. 56, n. 1, p. 125-129.
Karl, H.-V., Groning, E., and Brauckmann C., 2007, The Mesosauria in the collection of Gottingen and Clausthal: implicatiosn for a modifed classification: Clausthaler Geowissenschaften, v. 6, p. 63-78.
Langer, M. C., Ribeiro, A. M., Schultz, C. L., and Ferigolo, J., 2007, The continental tetrapod-bearing Triassic of South Brazil: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural, p. 201-218.
Larsson, H. C. E., 2007, MODEs of developmental evolution: an example wiht the origin and definition of the Autopodium: In: Major transitions in vertebrate evolution, edited by Anderson, J. S., and Sues, H.-S., p. 150-181.
Modesto, S. P., and Damiani, R., 2007, The procolophonid reptile Sauropareion anoplus from the Lowermost Triassic of South Africa: Journal of Vertebrate Paleontology, v. 27, n. 2, p. 337-349.
Modesto, S. P., Scott, D. M., Berman, D. S., Muller, J., and Reisz, R. R., 2007, The skull and the palaeoecological significance of Labidosaurus hamatus, a captorhinid reptile from the Lower Permian of Texas: Zoological Journal of the Linnean Society, v. 149, p. 237-262.
Müller, J., and Tsuji, L. A., 2007, Impendance-matching hearing in Paleozoic reptiles: evidence of advanced senory perception at an early state of amniote evolution: Public Library of Science (PLOS), One, v. 9, 7 pp.
Reisz, R. R., 2007, The cranial anatomy of basal diadectomorphs and the origin of amniotes: In: Major transitions in vertebrate evolution, edited by Anderson, J. S., and Sues, H.-S., p. 228-252.
Reisz, R. R., and Modesto, S. P., 2007, Heleosaurus scholtzi from the Permian of South Africa: a varanopid synapsid, not a diapsid reptile: Journal of Vertebrate Paleontology, v. 27, n. 3, p. 734-739.
Reisz, R. R., Muller, J., Tsuji, L., and Scott, D., 2007, The cranial osteology of Belebey vegrandis (Parareptilia: Bolosauridae), from the Middle Permian of Russia, and its bearing on reptilian evolution: Zoological Journal of the Linnean Society, v. 151, p. 191-214.Link to pdf
Saber, H., Wartit, M. El., Hmich, D., and Schneider, J. W., 2007, Tectonic evolution from the Hercynian shortening to the Triassic extension in the Paleozoic sediments of the Western High Atlas (Morocco): Journal of Iberian Geology, v. 33, n. 1, p. 31-40.
Schultz, C. L., and Langer, M. C., 2007, Tetrapodes Triassicos do Rio Grande do Sul, Brasil: In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 269-282.
Voigt, S., Berman, D. S., and Henrici, A. C., 2007, First well-established track-trackmaker association of Paleozoic tetrapods based on Ichniotherium trackways and diadectid skeletons from the Lower Permian of Germany: Journal of Vertebrate Paleontology, v. 27, n. 3, p. 553-570.
Turtles
Genus: Nova (2)
Chubutemys GAFFNEY, RICH, VICKERS-RICH, CONSTANTINE, VACCA, & KOOL, 2007
Linderochelys FUENTE, CALVO & GONZALEZ RIGA, 2007
Species: Nova (3)
Chubutemys copelloi GAFFNEY, RICH, VICKERS-RICH, CONSTANTINE, VACCA, & KOOL, 2007
Linderochelys rinconensis FUENTE, CALVO & GONZALEZ RIGA, 2007
Ordosemys brinkmania DANILOV & PARHAM, 2007
Synonym: Nova (1)
Xinjiangchelys wuerhoensis (YEH, 1977) emend DANILOV & PARHAM, 2007
= Sinemys wuerhoensis YEH, 1977
Batista, D. L., and Carvalho, I. de S., 2007, O genero Araripemys (Chelonii, Pleurodira) no Cretaceo Brasileiro: In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 283-289.
Candelro, C. R. dos A., Azevedo, R. P. de, and Silva, P. M. da., 2007, Preliminary approach on depositonal environmental of the Uberaba Formation (Upper Cretaceous, Peiropolis site, Minas Gerais State, Brazil: an introduduction: Caminhos de Geografia, v. 8, n. 2, p. 81-85.
Carrino, M. H., 2007, Taxonomic comparison and stratigraphic distribuion of Toxochelys (Testudines: Cheloniidae) of South Dakota: In: The Geology and Paleontology of the Late Cretaceous marine deposits of the Dakotas, edited by Martin, J. E., and Parris, D. C., The Geological Society of America, Special Paper 427, p. 111-132.
Danilov, I. G., and Parham, J. F., 2007, The type series of 'Sinemys' wuerhoensis, a problematic turtle from the Lower Cretaceous of China, includes at least three taxa: Palaeontology, v. 50, part 2, p. 431-444. (Ordosemys brinkmania)
Difley, R., 2007, Biostratigraphy of the North Horn Formation at North Horn Mountain, Emery County, Utah: UGA Publication n. 36, p. 439-454.
De Laparent de Broin, F,. de la Fuente, M. S., and Frenandez, M. S., 2007, Notoemys laticentralis (Chelonii, Pleurodira), Late Jurassic of Argentina: New examination of the anatomical structures and comparisons: Revue de Paleobiologie, n. 26, n. 1, p. 99-136. Link to pdf
de Fuente, M. S., 2007, Testudines: In: Patagonian Mesozoic Reptiles, edited by Gasparini, Z., Salgado, L., and Coria, R. A., Indiana University Press, p. 50-86.
de Fuente, M. S., Calvo, J. O., and Gonzalez Riga, B. J., 2007, A new Cretaceous chelid turtle from the northern Neuquen Basin, Agentina: Ameghiana, v. 44, n. 2, p. 485-492. (Linderochelys rinconensis)
de Fuente, M. S., Salgado, L., Albino, A., Baez, A. M., Bonaparte, J. F., Calvo, J. O., Chiappe, L. M., Codorniu, L. S., Coria, R. A., Gasparini, Z., Gonzalez Riga, B. J., Novas, F. E., and Pol, D., 2007, Tetrapodos continentales del Cretacico de la Argentina: una sintesis actualizada: Ameghiana, 50 anviersario, p. 137-153.
Gaffney, E. S., Rich, T. H., Vickers-Rich, P., Constantine, A, Vacca, R., and Kool, L., 2007, Chubutemys, a new eucryptodiran turtle from the Early Cretaceous of Argentina, and the relationships of the Meiolaniidae: American Museum Novitates, n. 3599, 35pp. (Chubutemys copelloi)
Gasparini, Z., Fernandez, M., Fuente, M. de la, and Salgado, L., 2007, Reptiles marinos jurasicos y cretacicos de la Patagonia argentina: sup aporte al conocimiento de la herpetofauna mesozoica: Ameghiniana, 50 anivesario, p. 125-136. Link to pdf
Harris, S. R., Pisani, D., Gower, D. J., and Wilkinson, M., 2007, Investigating stagnation in morphological phylogenetics using consensus data: Systematic Biology, v. 56, n. 1, p. 125-129.
Hoganson, J. W., Erickson, J. M., and Holland, Jr., F. D., Amphibian, reptilian, and avian remains from the Fox Hills Formation (Maastrichtian): shoreline and estuarine deposits of the Pierre Sea in south-central North Dakota: In: The Geology and Paleontology of the Late Cretaceous marine deposits of the Dakotas, edited by Martin, J. E., and Parris, D. C., The Geological Society of America, Special Paper 427, p. 239-256.
Ifrim, C., Stinnesbeck W., and Frey, E., 2007, Upper Cretaceous (Cenomanian-Turonian and Turonian-Coniacian) open marine plattenkalk deposits of New Mexico: Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 245, n. 1, p. 71-81.
Joyce, W. G., 2007, Phylogenetic relationships of Mesozoic Turtles: Bulletin of the Peabody Museum of Natural History, v. 48, n. 1, 3-102.
Lucas, S. G., and Tanner, L. H., 2007, Tetrapod biosratigraphy and biochronology of the Triassic-Jurassic transition on the southern Colorado Plateau, USA: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 244, p. 242-256.
Martill, D. M., Bechly, G, and Heads, S. W., 2007, Appendix: species list for the Crato Formation: In: The Crato Fossil Beds of Brazil, window into an Ancient World, edited by Martin, D. M., Bechly, G., and Loveridge, R. F., p. 582-607.
Matze, A. T., 2007, An almost complete juvenile specimen of the cheloniid turtle Ctenochelys stenoporus (Hay, 2905) from the Upper Cretaceous Niobrara Formation of Kansas, USA: Palaeontology, v. 50, part 3, p. 669-691.
Naish, D., 2007, Turtles of the Crato Formation: In: The Crato Fossil Beds of Brazil, window into an Ancient World, edited by Martin, D. M., Bechly, G., and Loveridge, R. F., p. 452-457.
Oheim, K. B., 2007, Fossil site predicition using geographic information systems (GIS) and suitability analysis: the Two Medicine Formation, MT, a test case: Paleogeography, Palaeoclimatology, Palaeoecology, v. 251, p. 354-365.
Oliveira, G. R. de, 2007, Aspectos tafonomicos de testudines da Formacao Santana (Cretaceo (Inferior), Bacia do Araripe, Nordeste do Brazil: Anuario do Instituto de Geociencias, v. 30, n. 1, p. 77-87.
Oliveria, G. R. de., and Kellner, A. W. A., 2007, Taxonomic status of Araripemys "arturi" fielding, Martill & Naish, 2005 (Testudines, Pleurodira, Araripemyididae): In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 385-391.
Sanchez-Hernandez, B., Benton, M. J., and Naish, D., 2007, Dinosaurs and other fossil vertebrates from the Late Jurassic and Early Cretaceous of the Galve area, NE Spain: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 249, p. 180-215.
Scheyer, T. M., and Sander, P. M., 2007, Shell bone histology indicates terrestrial palaeoecology of basal turtles: Proceedings of the Royal Socity, Series B,. v. 274, p. 1885-1893.
Sterli, J., de la Fuente, M. S., and Rougier, G. W., 2007, Anatomy and relationships of Palaeochersis talampayensis, a Late Triassic Turtle from Argentina: Palaeontographica, Abt. A., v. 281, Lfg. 1-3, p. 1-61. Link to pdf
Sterli, J., and Joyce, W. G., 2007, The cranial anataomy of the Early Jurassic turtle Kayentachelys aprix: Acta Palaontologica Polonica, v. 52, v. 4, p. 675-694.
Viohl, G., and Zapp, M,. 2007, Schamhaupten, an outstanding Fossil-Lagerstätte in a silicified Plattenkalk around the Kimmeridgian-Tithonian boundary (Southern Franconian Alb, Bavaria): Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 245, n. 1, p. 127-142.
Diapsida (Lower Lepidosauromorphs)
Genus: Nova (1)
Miodentosaurus CHENG, WU & SATO, 2007
Species: Nova (1)
Anshusaurus huangnihensis CHENG, CHEN & WANG, 2007
Miodentosaurus brevis CHENG, WU & SATO, 2007
Lizards and Snakes
Genus: Nova (3)
Komensaurus CALDWELL & PALCI, 2007
Tripennaculus NYDAM, & VOCI, 2007
Xianglong LI, GAO, HOU & XU, 2007
Species: Nova (9)
Adriosaurus microbrachis PALCI & CALDWELL, 2007
Dicothodon cifellii NYDAM, EATON & SANKEY, 2007
Globidens schumanni MARTIN, 2007a
Hainosaurus newmilleri MARTIN, 2007b
Komensaurus carrolli CALDWELL & PALCI, 2007
Mensicognathus molybrochoros NYDAM, & VOCI, 2007
Plioplatecarpus nichollsae CUTHBERTSON, MALLON, CAMPIONE, & HOLMES, 2007
Tripennaculus eatoni NYDAM, & VOCI, 2007
Xianglong zhaoi LI, GAO, HOU & XU, 2007
Synonym: Nova (3)
Dicothodon bajaensis (NYDAM, 1999) NYDAM, EATON & SANKEY, 2007
= Polyglyphanodon bajaensis NYDAM, 1999
Peneteius saueri (McCORD, 1998) NYDAM, EATON & SANKEY, 2007
=Manangyasaurus saueri McCORD, 1998
Taniwhasaurus antarcticus (NOVAS, FERENANDEZ, GASPARINI, LIRIO, NUNEZ & PUERTA, 2002) MARTIN, & FERNANDEZ, 2007
= Lakumasaurus antarcticus NOVAS, FERENANDEZ, GASPARINI, LIRIO, NUNEZ & PUERTA, 2002
Albino, A., 2007, Lepidosauromorpha: In: Patagonian Mesozoic Reptiles, edited by Gasparini, Z., Salgado, L., and Coria, R. A., Indiana University Press, p. 87-115.
Buchy, M.-C., Frey, E., Stinnesbeck, W., and Lopez-Oliva, J. G., 2007, Cranial anatomy of a Maastrichtian (Upper Cretaceous) mosasaur (Squamata, Mosasauridae) from north-east Mexico: Revista Mexicana de Ciencias Geologics, v. 24, n. 1, p. 89-103. Link to pdf
Caldwell, M. W., 2007, Ontogeny, anatomy and attachment of the dentition in mosasaurs (Mosasauridae: Squamata): Zoological Journal of the Linnean Society, v. 149, p. 687-700.
Caldwell, M. W., 2007, Snake phylogeny, origins, and evolution: the role, impact, and importance of fossils, (1869-2006): In: Major transitions in vertebrate evolution, edited by Anderson, J. S., and Sues, H.-S., p. 253-302.
Caldell, M. W., and Konishi, T., 2007, Taxonomic re-assignment of the first-known mosasaur specimen from Japan, and a discussion of circum-Pacific mosasaur paleobiogeography: Journal of Vertebrate Paleontology, v. 27, n. 2, p. 517-520.
Caldwell, M. W., and Palci, A., 2007, A new basal mosasauroid from the Cenomanian (U. Cretaceous) of Slovenia with a review of mosasauroid phylogeny and evolution: Journal of Vertebrate Paleontology, v. 27, n. 4, p. 863-880. (Komensaurus carrolli)
Chatterjee, S., and Scotese, C., 2007, Biogeography of the Mesozoic lepidosaurs on the wandering Indian plate: In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 551-579.
Conrad, J. L., and Norell, M. A., 2007, A complete Late Cretaceous Iguanian (Squamata, Reptilia) from the Gobi and identification of new iguaninan clade: American Museum Novitates, n. 3584, p. 47pp. (Saichangurvel davidsonae emend Saichangurvel davidsoni) Link to pdf
Note: The authors incorrectly used the wrong ending. They named it after a female preparator, Amy Davidson, and the ending should actually be -ae not -i.
Cuthbertson, R. S., Mallon, J. C., Campione, N. E., and Holmes, R. B., 2007, A new species of mosasaur (Squamata: Mosasauridae) from the Pierre Shale (Lower Campanian) of Manitoba: Canadian Journal of Earth Sciences, v. 44, n. 5, p. 593-606. (Plioplatecarpus nichollsae)
Diez Diaz, V., and Ortega, F., 2007, Un nuevo ejemplar de mosasaurio halisaurino del Cretacico Superior (Maastrichtiense) de la cuenca de Khourigba (Morocco): In: Cantera Paleontológica: 143-156. Diputación Provincial de Cuenca, Cuenca: 398 pp.
Difley, R., 2007, Biostratigraphy of the North Horn Formation at North Horn Mountain, Emery County, Utah: UGA Publication n. 36, p. 439-454.
Dutchak, A. R., and Caldwell, M. W., 2006 (published in 2007), Redescription of Aigialosaurus dalmaticus Kramberger, 1892, a Cenomanian mosasauroid lizard from Hvar Island, Croatia: Canadian Journal of Earth Sciences, v. 43, p. 1821-1834.
Evans, S. E., and Wang, Y., 2007, A juvenile lizard specimen with well-preserved skin impressions from the Upper Jurassic/Lower Cretaceous of Daohugou, Inner Mongolia, China: Naturwissenschaften, v. 94, p. 431-439. Link to pdf
Evans, S. E., Wang, Y., and Jones, M. E. H., 2007, An aggregation of lizard skeletons from the Lower Cretaceous of China: Senckenbergiana lethaea, v. 87, n. 1, p. 109-118. Link to pdf
de Fuente, M. S., Salgado, L., Albino, A., Baez, A. M., Bonaparte, J. F., Calvo, J. O., Chiappe, L. M., Codorniu, L. S., Coria, R. A., Gasparini, Z., Gonzalez Riga, B. J., Novas, F. E., and Pol, D., 2007, Tetrapodos continentales del Cretacico de la Argentina: una sintesis actualizada: Ameghiana, 50 anviersario, p. 137-153.
Gasparini, Z., Fernandez, M., Fuente, M. de la, and Salgado, L., 2007, Reptiles marinos jurasicos y cretacicos de la Patagonia argentina: sup aporte al conocimiento de la herpetofauna mesozoica: Ameghiniana, 50 anivesario, p. 125-136. Link to pdf
Harris, S. R., Pisani, D., Gower, D. J., and Wilkinson, M., 2007, Investigating stagnation in morphological phylogenetics using consensus data: Systematic Biology, v. 56, n. 1, p. 125-129.
Hoganson, J. W., Erickson, J. M., and Holland, Jr., F. D., Amphibian, reptilian, and avian remains from the Fox Hills Formation (Maastrichtian): shoreline and estuarine deposits of the Pierre Sea in south-central North Dakota: In: The Geology and Paleontology of the Late Cretaceous marine deposits of the Dakotas, edited by Martin, J. E., and Parris, D. C., The Geological Society of America, Special Paper 427, p. 239-256.
Ifrim, C., Stinnesbeck W., and Frey, E., 2007, Upper Cretaceous (Cenomanian-Turonian and Turonian-Coniacian) open marine plattenkalk deposits of New Mexico: Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 245, n. 1, p. 71-81.
Konishi, T., and Caldwell, M. W., 2007, New specimens of Platecarpus planifrons (Cope, 1874) (Squamata: Mosasauridae) and a revised taxonomy of the genus: Journal of Vertebrate Paleontology, v. 27, n. 1, p. 59-72.
Li, P.-P., Gao, K.-Q., Hou, L.-H., and Xu, X., 2007, A gliding lizard from the Early Cretaceous of China: Proceedings of the National Academy of Science, v. 104, n. 13, p. 5507-5509. (Xianlong zhaoi)
Lindgren, J., 2007, First record of Halisaurus (Squamata: Mosasauridae) from the Pacific coast of North America: PaleoBios, v. 27, n. 2, p. 40-47.
Lindgren, J., Jagt, J. W. M., and Caldwell, M. W., 2007, A fishy mosasaur: the axial skeleton of Plotosaurus (Reptilia, Squamata) reassessed: Lethaia, v. 40, p. 153-160.
Martill, D. M., 2007, Lizards of the Crato Formation: In: The Crato Fossil Beds of Brazil, window into an Ancient World, edited by Martin, D. M., Bechly, G., and Loveridge, R. F., p. 458-462.
Martill, D. M., Bechly, G, and Heads, S. W., 2007, Appendix: species list for the Crato Formation: In: The Crato Fossil Beds of Brazil, window into an Ancient World, edited by Martin, D. M., Bechly, G., and Loveridge, R. F., p. 582-607.
Martin, J. E., 2007a, A new species of the durophagous mosasaur Globidens (Squamata: Mosasauridae) from the Late Cretaceous Pierre Shale Group of Central South Dakota, USA: In: The Geology and Paleontology of the Late Cretaceous marine deposits of the Dakotas, edited by Martin, J. E., and Parris, D. C., The Geological Society of America, Special Paper 427, p. 177-198. (Globidens schurmanni)
Martin, J. E., 2007b, A North American Hainosaurus (Squamata: Mosasauridae) from the Late Cretaceous of southern South Dakota: In: The Geology and Paleontology of the Late Cretaceous marine deposits of the Dakotas, edited by Martin, J. E., and Parris, D. C., The Geological Society of America, Special Paper 427, p. 199-207. (Hainosaurus neumilleri)
Martin, J E., and Fernandez, M., 2007, The synonymy of the Late Cretaceous mosasaur (Squamata) genus Lakumasaurus from Antarctica with Taniwhasaurus from New Zealand and its bearing upon faunal similarity within the Weddellian Province: Geological Journal, v. 42, p. 203-311. (Taniwhasaurus antarcticus = Lakumasaurus antarcticus)
Martin, J. E., and Fox, J. E., 2007, Stomach contents of Globidens, a shell-crushing mosasaur (Squamata), from the Late Cretaceous Pierre Shale Group, Big Bend area of the Missouri River, central South Dakota: In: The Geology and Paleontology of the Late Cretaceous marine deposits of the Dakotas, edited by Martin, J. E., and Parris, D. C., The Geological Society of America, Special Paper 427, p. 167-176.
Meredith, R. W., Martin, J. E., and Wegleitner, P. N., 2007, The largest mosasaur (Squamata: Mosasauridae) from the Missouri River area (Late Cretaceous; Pierre Shale Group) of South Dakota and its relationships to Lewis and Clark: In: The Geology and Paleontology of the Late Cretaceous marine deposits of the Dakotas, edited by Martin, J. E., and Parris, D. C., The Geological Society of America, Special Paper 427, p. 209-214. Link to pdf
Nydam, R. L., Eaton, J. F., and Sankey, J., 2007, New taxa of transversely-toothed lizards (squamata: Scincomorpha) and new information on the evolutionary history of "Teiids": Journal of Paleontology, v. 81, n. 3, p. 538-549. (Dicothodon cifellii, Dicothodon bajaensis = Polyglyphanodon bajaensis, Peneteius sauri = Manangyasaurus saurei) Link to pdf
Nydam, R. L., and Voci, G. E., 2007, Teiid-like Scincomorphan lizards from the Late Cretaceous (Campanian) of Southern Utah: Journal of Herpetology, v. 41, n. 2, p. 211-219. (Tripennaculus eatoni, Mensicognathus molybrochoros). Link to pdf
Obata, I., Matsukawa, M., and Shibata, K., 2007, Geological age and environments of the plesiosaurs and the mosasaurs from Japan: Jubilee Publ. Commem. Prof. Kamei's 80th birthday, p. 155-177.
Oheim, K. B., 2007, Fossil site predicition using geographic information systems (GIS) and suitability analysis: the Two Medicine Formation, MT, a test case: Paleogeography, Palaeoclimatology, Palaeoecology, v. 251, p. 354-365.
Palci, A., and Caldwell, M. W., 2007 Vestigial forelimbs and axial elongation in a 95 million-year-old non-snake squamate: Journal of Vertebrate Paleontology, v. 27, n. 1, p. 1-7. (Adriosaurus microbrachis)
Patrick, D., Martin, J. E., Parris, D. C., and Grandstaff, D. E., 2007, Rare earth element (REE) analysis of fossil vertebrates from the Upper Cretaceous Pierre Shale Group for the purposes of paleobathymetric interpretations of the Western Interior Seaway: In: The Geology and Paleontology of the Late Cretaceous marine deposits of the Dakotas, edited by Martin, J. E., and Parris, D. C., The Geological Society of America, Special Paper 427, p. 71-83.
Patrick, D., Martin, J. E., Parris, D. C., and Grandstaff, D. E., 2007, Rare earth element determination of the stratigraphic position of the holotype of Mosasaurus missouriensis (Harlan), the first named fossil reptile from the American West: In: The Geology and Paleontology of the Late Cretaceous marine deposits of the Dakotas, edited by Martin, J. E., and Parris, D. C., The Geological Society of America, Special Paper 427, p. 155-165.
Salgado, L., Ferandez, M., and Talevi, M., 2007, Observaciones histologicas en reptiles marinos (Elasmosauridae y Mosasauridae) del Cretacico Tardio de Patagonia y Antartida: Ameghinana, v. 44, n. 3, p. 513-523.
Sanchez-Hernandez, B., Benton, M. J., and Naish, D., 2007, Dinosaurs and other fossil vertebrates from the Late Jurassic and Early Cretaceous of the Galve area, NE Spain: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 249, p. 180-215.
Scanferla, C. A., and Canale, J. I., 2007, The youngest record of the Cretaceous snake genus Dinilysia (Squamata, Serpentes): South American Journal of Herpetology, v. 2, n. 1, p. 766-81. Link to pdf
Shimada, K., Everhart, M. J., and Ewell, K., 2007, A unique reptilian (large dolichosaurid lizard?) tooth from the Upper Cretaceous Niobrara Chalk of western Kansas: Transactions of the Kansas Academy of Science, v. 110, n. 3/4, p. 213-219.
Shimada, K., and Ystesund, T. K., 2007, A dolichosaurid lizard, Coniasaurus cf. C. crassidens, from the Upper Cretaeous Carlile Shale in Russell County, Kansas: Transacations of the Kansas Academy of Science, v. 110, n. 3/4, p. 236-242.
Silva, M. C. da, Barreto, A. M. F., Carvalho, I. de S., and Carvalho, M. S. S., 2007, Relacao entre a morfologia da denticao e os habitos alimentares dos vertebrados da Bacia da Paraiba, Nordeste do Brasil: In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 433-440.
Sphenodonts
Genus: Nova (1)
Lamarquesaurus APESTEGUIA & ROUGIER, 2007
Species: Nova (1)
Lamarquesaurus cabazai APESTEGUIA & ROUGIER, 2007
Albino, A., 2007, Lepidosauromorpha: In: Patagonian Mesozoic Reptiles, edited by Gasparini, Z., Salgado, L., and Coria, R. A., Indiana University Press, p. 87-115.
Apesteguia, S., and Rougier, G. W., 2007, A Late Campanian Sphenodontid maxilla from Northern Patagonia: American Museum Novitates, n. 3581, 11pp. (Lamarquesaurus cabazai) Link to pdf
Chatterjee, S., and Scotese, C., 2007, Biogeography of the Mesozoic lepidosaurs on the wandering Indian plate: In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 551-579.
Gasparini, Z., Salgado, L., and Coria, R. A., 2007, Reptilian faunal succession in the Mesozoic and Patagonia, an updated overview: In: Patagonian Mesozoic Reptiles, edited by Gasparini, Z., Salgado, L., and Coria, R. A., Indiana University Press, p. 335-358.
Lucas, S. G., and Tanner, L. H., 2007, Tetrapod biosratigraphy and biochronology of the Triassic-Jurassic transition on the southern Colorado Plateau, USA: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 244, p. 242-256.
Spielmann, J. A., Lucas, S. G., and Heckert, A. B., 2007, Tetrapod fauna of the UpperTriassic (Revuletian) Owl Rock Formation, Chinle Group, Arizona:In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural and Science, bulletin 41, p. 371-383.
Viohl, G., and Zapp, M,. 2007, Schamhaupten, an outstanding Fossil-Lagerstätte in a silicified Plattenkalk around the Kimmeridgian-Tithonian boundary (Southern Franconian Alb, Bavaria): Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 245, n. 1, p. 127-142.
Ichthyosaurs
Species: Nova (1)
Callawayia wolonggangense CHEN, CHENG & SANDER, 2007
Synonym: Nova (1)
Stenopterygius cayi (FERNANDEZ, 1994) FERNANDEZ, 2007
= Chacaicosaurus cayi FERNANDEZ, 1994
Chen, X. H., Cheng, L., and Sander, P. M., 2007, A new species of Callawayia (Reptilia: Ichthyosauria) from the Late Triassic Guanling biota, Guizhou, China: Geological Bulletin of China, v. 22, p. 228-235. (Callawayia wolonggangense)
Fernandez, M., 2007, Redescription and phylogenetic position of Caypullisaurus (Ichthyosauria: Ophthalmosauridae): Journal of Paleontology, v. 81, n. 2, p. 368-375.
Fernandez, M., 2007, Ichthyosauria: In: Patagonian Mesozoic Reptiles, edited by Gasparini, Z., Salgado, L., and Coria, R. A., Indiana University Press, p. 271-291.
Gasparini, Z., Fernandez, M., Fuente, M. de la, and Salgado, L., 2007, Reptiles marinos jurasicos y cretacicos de la Patagonia argentina: sup aporte al conocimiento de la herpetofauna mesozoica: Ameghiniana, 50 anivesario, p. 125-136. Link to pdf
Jiang, D.-Y., Schmitz, L., Motani, R., Hao, W.-C., and Sun, Y.-L., 2007, The mixosaurid ichthyosaur Phalarodon cf. P. fraasi from the Middle Triassic of Guizhou Province, China: Journal of Paleontology, v. 81, n. 3, p. 602-605. Link to pdf
Lingham-Soliar, T., and Plodowski, G., 2007, Taphonomic evidence for high-speed adapted fins in thunniform ichthyosaurs: Naturwissenschaften, v. 94, p. 65-70.
Lucas, S. G., Siberling, N. J., Jenks, J. F., Spielmann, J. A., and Rinehart, L. F., 2007, Third day: Upper Triassic and Lower Jurassic stratigraphy and biostratigrphy in western Nevada: In: Triassic of the American West, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural History & Science, Bulletin 40, p. 23-32.
Reisdorf, A. G., 2007a, Der ichthyosaurier von Hauensteiner Nebelmeer wie eine kopflandung die wissenschaft kopf stehen lässt: Naturforschende Gesellschaft des Kantos Solothurn, heft 40, p. 7-22.
Reisdrof, A. G., 2007b, No Joke Movement mehr über den hauensteiner ichthyosaurier und rezente marine lungenatmer: Naturforschende Gesellschaft des Kantos Solothurn, heft 40, p. 23-49.
Viohl, G., and Zapp, M,. 2007, Schamhaupten, an outstanding Fossil-Lagerstätte in a silicified Plattenkalk around the Kimmeridgian-Tithonian boundary (Southern Franconian Alb, Bavaria): Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 245, n. 1, p. 127-142.
Sauropterygians and Placodontians
Genus: Nova (4)
Eopolycotylus ALBRIGHT, GILLETTE & TITUS, 2007a
Hydrorion GROßMANN, 2007
Pahaspasaurus SCHUMACHER, 2007
Palmulasaurus ALBRIGHT, GILLETTE & TITUS, 2007c
Species: Nova (2)
Eopolycotylus rankini ALBRIGHT, GILLETTE & TITUS, 2007a
Pahaspasaurus haasi SCHUMACHER, 2007
Synonym: Nova (2)
Hydrorion brachypterygius (HUENE, 1923) GROßMANN, 2007
= Plesiosaurus brachypterygius HUENE, 1923
= Microcleidus brachypterygius (HUENE, 1923), BAKKER, 1993
Palmulasaurus quadratus (ALBRIGHT, GILLETTE & TITUS, 2007a) ALBRIGHT, GILLETTE & TITUS, 2007c
= Palmula quadratus ALBRIGHT, GILLETTE & TITUS, 2007a
Albright III, L. B., Gillette, D. D., and Titus, A. L., 2007a, Plesiosaurs from the Upper Cretaceous (Cenomanian-Turonian) Tropic Shale of southern Utah, part 1: new records of the pliosaur Brachauchenius lucasi: Journal of Vertebrate Paleontology, v. 27, n. 1, p. 31-40.
Albright III, L. B., Gillette, D. D., and Titus, A. L., 2007b, Plesiosaurs from the Upper Cretaceous (Cenomanian-Turonian) Tropic Shale of southern Utah, part 2: polycotylidae Journal of Vertebrate Paleontology, v. 27, n. 1, p. 41-58. (Eopolycotylus rankini, Palmula quadratus) Link to pdf
Albright III, L. B., Gillette, D. D., and Titus, A. L., 2007c, Plesiosaurs from the Upper Cretaceous (Cenomanian-Turonian) tropic shale of southern Utah, part 2: Polycotylidae: repalcement names for the preoccupied genus Palmula and the subfamily Palmulainae: Journal of Vertebrate Paleontology, v. 27, n. 4, p. 1051. (Palmulasaurus quadratus = Palmula quadratus) Link to pdf
Arkhangelsky, M. S., Averianov, A. O., and Pervushov, E. M., 2007, Short-necked plesiosaurs of the family Polycotylidae from the Campanian of the Saratov Region: Palaeontological Journal, v. 2007, v. 41, n . 6, p. 656-660. Link to pdf
Everhart, M. J., 2007, Use of archival photographs to rediscover the locality of the Holyrood elasmosaur (Ellsworth County, Kansas): Transactions of the Kansas Academy of Science, v. 110, n. 1/2, p. 135-241.
Everhart, M. J., 2007, Historical note on the 1884 discovery of Brachauchenius lucasi (Plesiosauria; Pliosauridae) in Ottawa County, Kansas: Transactions of the Kansas Academy of Science, v. 110, n. 3/4, p. 255-258. Link to pdf
Farke, A. A., 2007, Reexamination of paleopathology in plesiosaurs and implications for behavioral interpretations: Journal of Vertebrate Paleontology, v. 27, n. 3, p. 724-726.
Gasparini, Z., 2007, Plesiosauria: In: Patagonian Mesozoic Reptiles, edited by Gasparini, Z., Salgado, L., and Coria, R. A., Indiana University Press, p. 292-313.
Gasparini, Z., Fernandez, M., Fuente, M. de la, and Salgado, L., 2007, Reptiles marinos jurasicos y cretacicos de la Patagonia argentina: sup aporte al conocimiento de la herpetofauna mesozoica: Ameghiniana, 50 anivesario, p. 125-136. Link to pdf
Großmann, F., 2007, The taxonomic and phylogenetic position of the plesiosauroidea from the Lower Jurassic Posidonia shale of south-west Germany: Palaeontology, v. 50, part 3, p. 545-564. (Hydrorion brachypterygius = Plesiosaurus brachypterygius) Link to pdf
Kear, B. P., 2007, A juvenile pliosauroid plesiosaur (Reptilia: Sauropterygia) from the Lower Creaceous of South Australia: Journal of Paleontology, v. 81, n. 1, p. 154-162.
Kear, B. P., 2007, Taxonomic clarification of the Australian elasmosaurid genus Eromangasaurus, with reference to other Austral elasmosaur taxa: Journal of Vertebrate Paleontology, v. 27, n. 1, p. 241-246.
Martin, J. E., Sawyer, J. F., Reguero, M., and Case, J. A., 2007, Occurrence of a young elasmosaurid plesiosaur skeleton from the Late Cretaceous (Maastrichtian) of Antarctica: U.S. Geological Survey and The National Academics, USGS of -2007-2047, Short Research Papers 066, doi:10.3133/of2007-1047.srp066, 4pp.
Noe, L. F., and Gomez-Perez, M., 2007, Postcript to Everhart, M. J., 2005. "Elasmosaurid remains from the Pierre Shale (Upper Cretaceous) of western Kansas. Possible missing elements of the type specimen of Elasmosaurus platyurus Cope, 1868?" - PalArch's Journal of Vertebrate Paleontology, 4, 3, p. 19-32: PalArch's Journal of Vertebrate Paleontology, 2, 1, p. 1-9.
Obata, I., Matsukawa, M., and Shibata, K., 2007, Geological age and environments of the plesiosaurs and the mosasaurs from Japan: Jubilee Publ. Commem. Prof. Kamei's 80th birthday, p. 155-177.
Salgado, L., Ferandez, M., and Talevi, M., 2007, Observaciones histologicas en reptiles marinos (Elasmosauridae y Mosasauridae) del Cretacico Tardio de Patagonia y Antartida: Ameghinana, v. 44, n. 3, p. 513-523.
Salgado, L., Parras, A., and Gasparini, Z., 2007, Un plesiosaurio de cuello corto (Plesiosauroidea, Polycotylidae) del Cretacico Superior del norte de Patagonia: Ameghiniana, v. 44, n. 2, p. 349-358.
Schumacher, B. A., 2007, A new polycotylid plesiosaur (Reptilia; Sauropterygia) from the Greenhorn Limestone (Upper Cretaceous; lower upper Cenomanian), Black Hills, South Dakota: In: The Geology and Paleontology of the Late Cretaceous marine deposits of the Dakotas, edited by Martin, J. E., and Parris, D. C., The Geological Society of America, Special Paper 427, p. 133-146. (Pahasapasaurus haasi).
Smith, A. S., 2007, The back-to-front plesiosaur Cryptoclidus (Apractocleidus) aldingeri from the Kimmeridgian of Milne Land, Greenland: Bulletin of the Geological Society of Denmark, v. 58, p. 1-7.
Thompson, W. A., Martin, J. E., and Reguero, M., 2007, Comparison of gastroliths within plesiosaurs (Elasmosauridae) from the Late Cretaceous marine deposits of Vega Island, Antarctic Peninsula, and the Missouri River area, South Dakota: In: The Geology and Paleontology of the Late Cretaceous marine deposits of the Dakotas, edited by Martin, J. E., and Parris, D. C., The Geological Society of America, Special Paper 427, p. 147-153.
Vincent, P., Bardet, N., and Morel, N., 2007, An Elasmosaurid plesiosaur from the Aalenian (Middle Jurassic) of Western France: Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 243, n. 3, p. 363-370.
Wahl, W. R., Ross, M., and Massare, J. A., 2007, Rediscovery of Wilbur Knight's Megalneusaurus rex site: new material from an old pit: Paludicola, v. 6, n. 2, p. 94-104. Link to pdf
Lower Archosauromorpha
Genus: Nova (1)
Mecistotrachelos FRASER, OLSEN, DOOLEY & RYAN, 2007
Species: Nova (3)
Anshunsaurus huangnihensis CHENG, CHEN & WANG, 2007
Macrocnemus fuyanensis LI, ZHAO, & WANG, 2007
Mecistotrachelos apeoros FRASER, OLSEN, DOOLEY & RYAN, 2007
Buffetaut, E., Li, J., Tong, H., and Zhang, H., 2007, A two-headed reptile from the Cretaceous of China: Biology Letters, v. 3, p. 80-81.
Casey, M. M., Fraser, N. C., and Kowalewski, M., 2007, Quantitative taphonomy of a Triassic reptile Tanytrachelos ahynis from the Cow Branch Formation, Dan River Basin, Solite Quarry, Virginia: Palaios, v. 22, p. 598-611.
Cheng, L., Chen, X., and Wang, C., 2007, A new species of Late Triassic Anshunsaurus (Reptilia: Thalattosauria) from Guizhou PRovince, Acta Geologica Sinica, v. 81, n. 10, p. 1345-1351. (Anshunsaurus huangnihensis) Link to pdf
Dzik, J., and Sulej, T., 2007, A review of the Early Late Triassic Krasiejow Biota from Silesia, Poland: Palaeontologica Polonica, n. 64, p. 1-27.
Fraser, N. C., Olsen, P. E., Dooley jr, A. C., and Ryan, T. R., 2007, A new gliding tetrapod (Diapsida: ?Archosauromorpha) from the Upper Triassic (Carnian) of Virginia: Journal of Vertebrate Paleontology, v. 27, n. 2, p. 261-265. (Mecistotrachelos apeoros)
Gao, K.-Q., Ksepka, D., Hou, L., Duan, Y., and Hu, D., 2007, Cranial morphology of an Early Cretaceous Monjurosuchid (Reptilia: Diapsida) from the Liaoning Province of China and evolution of the chrostoderan palate: Historical Biology, v. 19, n. 3, p. 215-224.
Gao, K.-Q., and Li, Q., 2007, Osteology of Monjurosuchus splendens (Diapsida: Christodera) based on a new specimen from the Lower Cretaceous of western Liaoning, China: Cretaceous Research, v. 28, p. 261-271.
Harris, S. R., Pisani, D., Gower, D. J., and Wilkinson, M., 2007, Investigating stagnation in morphological phylogenetics using consensus data: Systematic Biology, v. 56, n. 1, p. 125-129.
Hoganson, J. W., Erickson, J. M., and Holland, Jr., F. D., Amphibian, reptilian, and avian remains from the Fox Hills Formation (Maastrichtian): shoreline and estuarine deposits of the Pierre Sea in south-central North Dakota: In: The Geology and Paleontology of the Late Cretaceous marine deposits of the Dakotas, edited by Martin, J. E., and Parris, D. C., The Geological Society of America, Special Paper 427, p. 239-256.
Hone, D. W. E., and Benton, M. J., 2007, An evaluation of the phylogenetic relationships of the pterosaurs among archosauromorph reptiles: Journal of Systematic Palaeontology, v. 5, n. 4, p. 465-469.
Irmis, R. B., Parker, W. G., Nesbitt, S. J., and Liu, J., 2007, Early ornithischian dinosaurs: the Triassic record: Historical Biology, v. 19, n. 1, p. 3-22.
Katsura, Y., 2007, Fusion of sacrals and anatomy in Champsosaurus (Diapsida, Choristodera): Historical Biology, v. 19, n. 3, p. 263-271.
Kubo, T., and Benton, M. J., 2007, Evolution of hindlimb posture in archosaurs: limb stresses in extinct vertebrates: Palaeontology, v. 50, part 6, p. 1519-1529.
Langer, M. C., Ribeiro, A. M., Schultz, C. L., and Ferigolo, J., 2007, The continental tetrapod-bearing Triassic of South Brazil: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural, p. 201-218.
Li, C., 2007, A juvenile Tanystropheus sp. (Protosauria, Tanystropheidae) from the Middle Triassic of Guizhou, China: Vertebrata PalAsiatica, v. 45, n. 1, p. 37-42.
Li, C., Zhao, L.-J., and Wang, L.-T., 2007, A new species of Macrocnemus (Reptilia: Protosauria) from the Middle Triassic of southwestern China and its palaeogeographical implications: Science in China Series D-Earth Sciences, v. 50, p. 1601-1605. (Macrocnemus fuyanensis)
Lucas, S. G., Heckert, A. B., Spielmann, J. A., Tanner, L. H., and Hunt, A. P., 2007, Second Day: Early and Middle Triassic stratigraphy, sedimentology and paleontology in northeastern Arizona: In: Triassic of the American West, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural History & Science, Bulletin 40, p. 181-187.
Lucas, S. G., Heckert, A. B., Spielmann, J. A., Tanner, L. H., and Hunt, A. P., 2007, Third Day: Triassic stratigraphy and paleontology in northeastern Arizona: In: Triassic of the American West, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural History & Science, Bulletin 40, p. 186-197.
Lucas, S. G., and Tanner, L. H., 2007, Tetrapod biosratigraphy and biochronology of the Triassic-Jurassic transition on the southern Colorado Plateau, USA: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 244, p. 242-256.
Matsumoto, R., Evans, S. E., and Manabe, M., 2007, The choristoderan reptile Monjurosuchus from the Early Cretaceous of Japan: Acta Palaeontologica Polonica, v. 52, n. 2, p. 339-350. Link to pdf
Nesbitt, S. J., Irmis, R. B., and Parker, W. G., 2007, A critical re-evaluation of the Late Triassic Dinosaur Taxa of North America: Journal of Systematic Palaeontology, v. 5, n. 2, p. 209-243.
Nosotti, S., 2007, Tanystropheus longobardicus (Reptilia, Protorosauria): re-interpretations of the anatomy based on new specimens from the Middle Triassic of Besano (Lombardy, Northern Italy): Memorie della Societa Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, v. 35, fasc. III, 88pp. Link to pdf
Oheim, K. B., 2007, Fossil site predicition using geographic information systems (GIS) and suitability analysis: the Two Medicine Formation, MT, a test case: Paleogeography, Palaeoclimatology, Palaeoecology, v. 251, p. 354-365.
Renesto, S., and Dalla Vecchia, F. M., 2007, A revision of Longobardisaurus rossii Bizzarini and Muscio, 1995 from the Late Triassic of Friuli (Italy): Rivista Italiana di Paleontologi e Stratigrafia, v. 113, n. 2, p. 191-201.
Sanchez-Hernandez, B., Benton, M. J., and Naish, D., 2007, Dinosaurs and other fossil vertebrates from the Late Jurassic and Early Cretaceous of the Galve area, NE Spain: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 249, p. 180-215.
Scheetz, R. D., and Britt, B. B., 2007, Paleontological discoveries of James A. "Dinosaur Jim" Jensen in Central Utah: UGA Publication 36, p. 455-465.
Schultz, C. L., and Langer, M. C., 2007, Tetrapodes Triassicos do Rio Grande do Sul, Brasil: In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 269-282.
Spielmann, J. A., Lucas, S. G., and Heckert, A. B., 2007, Tetrapod fauna of the UpperTriassic (Revuletian) Owl Rock Formation, Chinle Group, Arizona: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural, p. 371-383.
Spielmann, J. A., Lucas, S. G., Heckert, A. B., Rinehart, L. F., and Hunt, A. P., 2007, Taxonomy and biostratigraphy of the Late Triassic archosauromorph Trilophosaurus: In: Triassic of the American West, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural History & Science, Bulletin 40, p. 231-240.
Vandermark, D., Tarduno, J. A., and Brinkman, D. B., 2007, A fossil champsosaur population from the high Arctic: implications for Late Cretaceous paleotemperatures: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 248, p. 49-59. Link to pdf
Vega-Dias, C., and Schultz, C. L., 2007, Evidence of archosaurifrom feeding on dicynodonts in the Late Triassic of southern Brazil: PaleoBios, v. 27, n. 2, p. 62-67.
“Thecodontia, Pseudosuchia"
Genus: Nova (3)
Adamanasuchus LUCAS, HUNT & SPIELMANN, 2007
Arganasuchus JALIL & PEYER, 2007
Heliocanthus PARKER, 2007
Species: Nova (2)
Adamanasuchus eisenhardtae LUCAS, HUNT & SPIELMANN, 2007
Arganasuchus dutuiti JAILIL & PEYER, 2007
Synonym: Nova (2)
Heliocanthus chamaensis (ZEIGLER, HECKERT & LUCAS, 2002) PARKER, 2007
= Desmatosuchus chamaensis ZEIGLER, HECKERT & LUCAS, 2002
Poposaurus langstoni (LONG & MURRY, 1995) WEINBAUM & HUNGERBUHLER, 2007
= Lythrosuchus langstoni LONG & MURRY, 1995
= Postosuchus kirkpatricki CHATTERJEE, 1985 (partim)
Desojo, J. B, and Baez, A. M., 2007, Cranial morphology of the Late Triassic South American archosaur Neoaetosauroides engaeus: evidence for aetosaurian diversity: Palaeontology, v. 50, part 1, p. 267-276.
Dzik, J., and Sulej, T., 2007, A review of the Early Late Triassic Krasiejow Biota from Silesia, Poland: Palaeontologica Polonica, n. 64, p. 1-27.
Harris, S. R., Pisani, D., Gower, D. J., and Wilkinson, M., 2007, Investigating stagnation in morphological phylogenetics using consensus data: Systematic Biology, v. 56, n. 1, p. 125-129.
Heckert, A. B., Lucas, S. G., Hunt, A. P., and Spielmann, J. A., 2007, Late Triassic aetosaur biochronology revisited: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural and Science, bulletin 41, p. 49-50.
Heckert, A. B., Spielmann, J. A., Lucas, S. G., and Hunt, A. P., 2007, Biostratigraphy utility of the Upper Triassic aetosaur Tecovasuchus (Archosauria: Stagonolepididae), anindex taxon of St. Johnsian (Adamanian: Late Carnian) time: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural and Science, bulletin 41, p. 51-57.
Irmis, R. B., 2007, Axial skeleton ontogeny in the parasuchia (Archosauria: Pseudosuchia) and its implications for ontogenetic determination in archosaurs: Journal of Paleontology, v. 27, n. 2, p. 350-361.
Irmis, R. B., Parker, W. G., Nesbitt, S. J., and Liu, J., 2007, Early ornithischian dinosaurs: the Triassic record: Historical Biology, v. 19, n. 1, p. 3-22. Link to pdf
Jalil, N.-E., and Peyer, K., 2007, A new rauisuchian (Archosauria, Suchia) from the Upper Triassic of the Argana basin, Morocco: Palaeontology, v. 50, part 2, p. 417-430. (Arganasuchus dutuiti)
Kubo, T., and Benton, M. J., 2007, Evolution of hindlimb posture in archosaurs: limb stresses in extinct vertebrates: Palaeontology, v. 50, part 6, p. 1519-1529.
Langer, M. C., Ribeiro, A. M., Schultz, C. L., and Ferigolo, J., 2007, The continental tetrapod-bearing Triassic of South Brazil: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural and Science, bulletin 41, , p. 201-218.
Lucas, S. G., Heckert, A. B., Spielmann, J. A., Tanner, L. H., and Hunt, A. P., 2007, Second Day: Early and Middle Triassic stratigraphy, sedimentology and paleontology in northeastern Arizona: In: Triassic of the American West, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural History & Science, Bulletin 40, p. 181-187.
Lucas, S. G., Heckert, A. B., Spielmann, J. A., Tanner, L. H., and Hunt, A. P., 2007, Third Day: Triassic stratigraphy and paleontology in northeastern Arizona: In: Triassic of the American West, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural History & Science, Bulletin 40, p. 186-197.
Lucas, S. G., Heckert, A. B., and Rinehart, L., 2007, A giant skull, ontogenetic variation and taxonomic validity of the Late Triassic phytosaur Parasuchus: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural and Science, bulletin 41, p. 222-228.
Lucas, S. G., Hunt, A. P., Heckert, A. P., and Spielmann, J. A., 2007, Global Triassic tetrapod biostratigraphy and biochronology: 2007 status: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural and Science, bulletin 41, p. 229-240.
Lucas, S. G., Hunt, A. P., and Spielmann, J. A., 2007, A new aetosaur from the Upper Triassic (Adamanian: Carnian) of Arizona: In: Triassic of the American West, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural History & Science, Bulletin 40, p. 241-247. (Adamanasuchus eisenhardtae).
Lucas, S. G., Spielmann, J. A., Heckert, A. B., and Hunt, A. P., 2007, Topotypes of Typothorax coccinarum, a Late Triassic aetosaur from the American southwest: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural and Science, bulletin 41, p. 241-247.
Lucas, S. G., Spielmann, J. A., and Hunt, A. P., 2007, Biochronological significance of the Late Triassic tetrapods from Krasijow, Poland: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural and Science, bulletin 41, p. 248-258.
Lucas, S. G., and Tanner, L. H., 2007, Tetrapod biosratigraphy and biochronology of the Triassic-Jurassic transition on the southern Colorado Plateau, USA: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 244, p. 242-256.
Nesbitt, S. J., 2007, The Anatomy of Effigia okeeffeae (Archosauria, Suchia), Theropod-like convergence, and the distribution of related taxa: Bulletin of the American Museum of Natural History, n. 302, 84pp.
Nesbitt, S. J., Irmis, R. B., and Parker, W. G., 2007, A critical re-evaluation of the Late Triassic Dinosaur Taxa of North America: Journal of Systematic Palaeontology, v. 5, n. 2, p. 209-243.
Parker, W. G., 2007, Reassessment of the aetosaur 'Desmatosuchus' chamaensis with a reanalysis of the phylogeny of the aetosauria (Archosauria: Pseudosuchia): Journal of Systematic Paleontology, v. 5, n. 1, p. 41-68. (Rioarribasuchus chamaensis = Desmatosuchus chamaensis = Heliocanthus chamaensis. Note: Desmatosuchus chamaensis was renamed by Lucas, S. G., Hunt, A. P., and Spielmann, J. A., 2006 as this paper was in press)
Prieto-Marquez, A., Gignac, P. M., and Joshi, S., 2007, Neontological evaluation of pelvic skeletal attributes purported to reflect sex in extinct non-avian archosaurs: Journal of Vertebrate Paleontology, v. 27, n. 3, p. 603-609.
Schoch, R. R., 2007, Osteology of the small archosaur Aetosaurus from the Upper Triassic of Germany: Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 246, n. 1, p. 1-35.
Spielmann, J. A., Lucas, S. G., and Heckert, A. B., 2007, Tetrapod fauna of the UpperTriassic (Revuletian) Owl Rock Formation, Chinle Group, Arizona: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural and Science, bulletin 41, p. 371-383.
Tanner, L. H., and Lucas, S. G., 2007, The Moenave Formation: Sedimentologic and stratigraphic context of the Triassic-Jurassic boundary in the Four Corners area, southwestern U.S.A.: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 244, p. 111-125.
Wedal, M., 2007, What pneumaticity tells us about 'prosauropods', and vice versa: Special Papers in Paleontology, v. 77, p. 207-222.
Weinbaum, J. C., and Hungerbühler, A., 2007, A revision of Poposaurus gracilis (Archosauria: Suchia) based on two new specimens from the Late Triassic of southwestern U.S.A: Paläontologische Zeitschrift, v. 81, Heft 2, p. 131-145. (Poposaurus langstoni = Lythrosuchus langstoni)
Crocodilia (Mesosuchia, Eusuchia)
Genus: Nova (9
Aktiogavialis VELEZ-JURABE, BROCHU, & SANTOS, 2007
Barinasuchus PAOLILLO & LINARES, 2007
Dollosuchoides BROCHU, 2007a
Iharkutosuchus OSI, CLARK & WEISHAMPEL, 2007
Langstonia PAOLILLO & LINARES, 2007
Montealtasuchus CARVALHO, VASCONCELLOS & TAVARES, 2007
Oceanosuchus HUA, BUFFETAUT, LEGALL & ROGRON, 2007
Voay BROCHU, 2007
Zulmasuchus PAOLILLO & LINARES, 2007
Species: Nova (11)
Aktiogavialis puertoricensis VELEZ-JURABE, BROCHU, & SANTOS, 2007
Barinasuchus arveloi PAOLILLO & LINARES, 2007
= Sebecus cf. huilensis BUFFETAUT & OFFSTETTER, 1977
Congosaurus compressus (BUFFETAUT, 1979) JOUVE, 2007
= Rhabdognathus compressus BUFFETAUT, 1979
Dollosuchoides densmorei BROCHU, 2007a
Eocaiman palaeocoenicus BONA, 2007
Iharkutosuchus makadii OSI, CLARK & WEISHAMPEL, 2007
Mariliasuchus robustus NOBRE, CARVALHO, VASCONCELLOS & NAVA, 2007
Montealtasuchus arrudacamposi CARVALHO, VASCONCELLOS & TAVARES, 2007
Neuquensuchus universitas FIORELLI & CALVO, 2007
Oceanosuchus boecensis HUA, BUFFETAUT, LEGALL & ROGRON, 2007
Rhabdognathus aslerensis JOUVE, 2007
Synonym: Nova (5)
Kentisuchus spenceri (BUCKLAND, 1837) BROCHU, 2007
= Crocodilus spenceri (BUCKLAND, 1837) BROCHU, 2007
= Crocodilus toliapicus OWEN, 1850
= Kentisuchus toliapicus (OWEN, 1850) MOOK, 1955
= Crocodilus delucii GRAY, 1831
= Crocodilus champsoides OWEN, 1850
Langstonia huilensis (LANGSTON, 1965) PAOLILLO & LINARES, 2007
= Sebecus huilensis LANGSTON, 1965
Shuvosaurus okeeffeae (NESBIT & NORELL, 2006) LUCAS, SPIELMANN & HUNT, 2007
= Effigia okeeffeae NESBITT & NORELL, 2006
Voay robustus (GRANDIDIER & VAILLANT, 1872) BROCHU, 2007
= Crocodilus robustus GRANDIDIER & VAILLANT, 1872
Zulmasuchus querejazus (BUFFETAUT & MARSHELL, 1991) PAOLILLO & LINARES, 2007
= Sebecus querejazus BUFFETAUT & MARSHALL, 1991
Azevedo, R. P. F. de, Vasconcellos, P. L. de, Canderio, C. R. dos A., and Bergqvist, L. P., 2007, Restos microscopicos de vertebrados fosseis do Grupo Bauru (Neocertaceo), no oest do estado de Sao Paulo, Brasil: In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 533-541.
Bona, P., 2007, Un nueva especie de Eocaiman Simpson (Crocodylia, Alligatoridae) del Paleoceno Inferior de Patagonia: Ameghiniana, v. 44, n. 2, p. 435-445. (Eocaiman palaeocenicus)
Brochu, C. A., 2007, Systematics and taxonomy of Eocene tomistomine crocodylians from Britain and northern Europe: Palaeontology, v. 50, part 4, p. 917-928. (Dollosuchoides densmorei)
Brochu, C. A., 2007, Morphology, relationships, and biogeographical significance of an extinct horned crocodilie (Crocodylia, Crocodylidae) from the Quaternary of Madagascar: Zoological Journal of the Linnean Society, v. 150, p. 835-863. (Voay robustus = Crocodylus robustus)
Brochu, C. A., Nieves-Rivera, A. M., Velez-Juarbe, J., Daza-Vaca, J. D., and Santos, H., 2007, Tertiary crocodylians from Puerto Rico: Evidence for Late Tertiary endemic crocodylians in the West Indies? Geobios, v. 40, p. 51-59.
Calvo, J. O., Porfiri, J. D., Gonzalez-Riga, B. J., and Kellner, A. W. A., 2007, A new Cretaceous terrestrial ecosystem from Gondwana with description of a new sauropod dinosaur: Anais da Academia Brasilieria de Ciencias, v. 79, n. 3, p. 529-541.
Carvalho, I. de S., and Vasconcellos, F. M. de, and Tavares, S. A., S., 2007, Montealtosuchus arrudacamposi, a new peirosaurid crocodile (Mesoeucrocodylia) from the Late Cretaceous Adamantina Formation of Brazil: Zootaxa, v. 1607, p. 35-46. (Montealtosuchus arrudacomposi) Link to pdf
Delfino, M., Bohme, M., and Rook, L., 2007, First European evidence for transcontinental dispersal of Crocodylus (Late Neogene of southern Italy): Zoological Journal of the Linnean Society, v. 149, p. 293-307. Link to pdf
Fiorelli, L. E., and Calvo, J. O., 2007, The first "Protosuchian" (Archosauria: Crocodyliformes) from the Cretaceous (Santonian) of Gondwana: Arquivos do Museu Nacional, Rio de Janeiro, v. 65, n. 4, p. 417-459. (Neuquensuchus universitas)
Frey, E. D., and Salisbury, S. W., 2007, Crocodilians of the Crato Formation: evidence for enigmatic species: In: The Crato Fossil Beds of Brazil, window into an Ancient World, edited by Martin, D. M., Bechly, G., and Loveridge, R. F., p. 463-474.
de Fuente, M. S., Salgado, L., Albino, A., Baez, A. M., Bonaparte, J. F., Calvo, J. O., Chiappe, L. M., Codorniu, L. S., Coria, R. A., Gasparini, Z., Gonzalez Riga, B. J., Novas, F. E., and Pol, D., 2007, Tetrapodos continentales del Cretacico de la Argentina: una sintesis actualizada: Ameghiana, 50 anviersario, p. 137-153.
Garcia, K. L., 2007, A familia Peirosauridae do Cretaceo do Gondwana: taxonomia comparada e implicacoes paleogeograficas: In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 393-404.
Garrison jr, J. R., Brinkman, D., Nichols, D. J., Layer, P., Burge, D., and Thayn, D., 2007, A multidisiplinary study of the Lower Cretaceous Cedar Mountain Formation, Mussentuchit Wash, Utah: a determination of the paleoenvironment and paleoecology of the Eolambia caroljonesa dinosaur quarry: Cretaceous Research, v. 28, p. 461-494.
Gasparini, Z., Fernandez, M., Fuente, M. de la, and Salgado, L., 2007, Reptiles marinos jurasicos y cretacicos de la Patagonia argentina: sup aporte al conocimiento de la herpetofauna mesozoica: Ameghiniana, 50 anivesario, p. 125-136. Link to pdf
Hoganson, J. W., Erickson, J. M., and Holland, Jr., F. D., Amphibian, reptilian, and avian remains from the Fox Hills Formation (Maastrichtian): shoreline and estuarine deposits of the Pierre Sea in south-central North Dakota: In: The Geology and Paleontology of the Late Cretaceous marine deposits of the Dakotas, edited by Martin, J. E., and Parris, D. C., The Geological Society of America, Special Paper 427, p. 239-256.
Hua, S., Buffetaut, E., Legall, C., and Rogron, P., 2007, Oceanosuchus boecensis n. gen, n. sp., a marine pholidosaurid (Crocodylia, Mesosuchia) from the Lower Cenomanian of Normandy, (Western France): Bulletin of the Geological Society of France, t. 178, n. 6, p. 503-513. (Oceanosuchus boecensis)
Jouve, S., 2007, Taxonomic revision of the dryosaurid assemblage (Crocodyliformes: Mesoeucrocodylia) from the Paleocene of the Iullemmeden basin, West Africa: Journal of Paleontology, v. 81, n. 1, p. 163-175. (Rhabdognathus keinensis, Rhabdognathus aslerensis, Congosaurus compressus = Rhabdognathus compressus)
Larsson, H. C. E., and Sues, H.-D., 2007, Cranial osteology and phylogenetic relationships of Hamadasuchus rebouli (Crocodyliformes: Mesoeucrocodylia) from the Cretaceous of Morocco: Zoological Journal of the Linnean Society, v. 149, p. 533-567.
Lucas, S. G., Heckert, A. B., Spielmann, J. A., Tanner, L. H., and Hunt, A. P., 2007, Third Day: Triassic stratigraphy and paleontology in northeastern Arizona: In: Triassic of the American West, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural History & Science, Bulletin 40, p. 186-197.
Lucas, S. G., Spielmann, J. A., and Hunt, A. P., 2007b, Taxonomy of Shuvosaurus, a Late Triassic archosaur from the Chinle Group, American southwest: In: The Global Triassic, Edited by Lucas, S. G., and Spielmann, J. A., New Mexico Museum of Natural and Science, bulletin 41, p. 249-261. (Shuvosaurus okeeffeae = Effigia okeeffeae) Link to pdf
Lucas, S. G., and Tanner, L. H., 2007, Tetrapod biosratigraphy and biochronology of the Triassic-Jurassic transition on the southern Colorado Plateau, USA: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 244, p. 242-256.
Marinho, T. da S., and Carvalho, I. de S., 2007, Revision of hte Sphagesauridae Kuhn, 1968 (Crocodyliformes, Mesoeurocodylia): In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 581-592.
Martill, D. M., Bechly, G, and Heads, S. W., 2007, Appendix: species list for the Crato Formation: In: The Crato Fossil Beds of Brazil, window into an Ancient World, edited by Martin, D. M., Bechly, G., and Loveridge, R. F., p. 582-607.
Martin, J. E., 2007, New material of the Late Cretaceous globidontan Acynodon iberoccitanus (Crocodylia) from southern France: Journal of Paleontology, v. 27, n. 2, p. 362-372.
Martinelli, A. G., Garrido, A. C., Forasiepi, A. M., Paz, E. R., and Gurovich, Y., 2007, Notes on fossil remains from the Early Cretaceous Lohan Cura Formation, Neuquen Province, Argentina: Gondwana Research, v. 11, p. 537-552.
Nesbitt, S., 2007, The Anatomy of Effigia okeeffeae (Archosauria, Suchia), Theropod-like convergence, and the distribution of related taxa: Bulletin of the American Museum of Natural History, n. 302, 84pp.
Nobre, P. H., Carvalho, I. de S., Vasconcellos, F. M. de, and Nava, W. R., 2007, Mariliasuchus robustus, um nuvo crocodylomorpha (Mesoeucrocodylia) da Bacia Bauru, Brazil: Anuario do Instituto de Geociencias, v. 30, n. 1p., 32-43. (Mariliasuchus robustus)
Oheim, K. B., 2007, Fossil site predicition using geographic information systems (GIS) and suitability analysis: the Two Medicine Formation, MT, a test case: Paleogeography, Palaeoclimatology, Palaeoecology, v. 251, p. 354-365.
Osi, A., Clark, J. M., and Weishampel, D. B., 2007, First report on a new basal eusuchian crocodyliform with multicuspid teeth from the Upper Cretaceous (Santonian) of Hungary: Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 243, n. 2, p. 169-177. (Iharkutosuchus makadii)
Paolillo, A., and Linares, O. J., 2007, Nuevos cocodrilos Sebecosuchia del Ceozoico Suramericano (Mesosuchia: Crocodylia): Paleobiologia Neotropical, n. 3, p. 25pp. (Barinasuchus arveloi, Langstonia huilensis = Sebecus huilensis, Zulmasuchus querejazui = Sebecus querejazui) Link to Pdf
Piras, P., Delfino, M., del Favero, L., and Kotsakis, T., 2007, Phylogenetic position of the crocodylian Megadontosuchus arduini and tomistomine palaeobiogeography: Acta Paleontologica Polonica, v. 52, p. 2, p. 315-328.
Pol, D., and Gasparini, Z., 2007, Crocodyliformes: In: Patagonian Mesozoic Reptiles, edited by Gasparini, Z., Salgado, L., and Coria, R. A., Indiana University Press, p. 116-142.
Prieto-Marquez, A., Gignac, P. M., and Joshi, S., 2007, Neontological evaluation of pelvic skeletal attributes purported to reflect sex in extinct non-avian archosaurs: Journal of Vertebrate Paleontology, v. 27, n. 3, p. 603-609.
Salas-Gismondi, R., Antoine, P.-O., Baby, P., Brusset, S., Benammi, M., Espurt, N., de Franceschi, D., Pujos, F., and Tegada, J, and Urbina, M., 2007, Middle Miocene crocodiles from the Fitzcarrald arch, Amazonian Peru: In: 4th European Meeting on the Paleontology and Stratigraphy of Latin America, edited by Diaz-Martinez, E., and Rabano, I., Cuademos del Museo Geomienro, n. 8, Instituto Geologico y Minero de Espana, Madrid, 2007, p. 355-360.
Sanchez-Hernandez, B., Benton, M. J., and Naish, D., 2007, Dinosaurs and other fossil vertebrates from the Late Jurassic and Early Cretaceous of the Galve area, NE Spain: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 249, p. 180-215.
Schweitzer, M. H., Elsey, R. M., Dacke, C. G., Horner, J. R., and Lamm, E.-T., 2007, Do egg-laying crocodilian (Alligator mississippiensis) archosaurs form medullary bone? Bone, v. 40, p. 1152-1158.
Shimada, K., and Parris, D. C., 2007, A long-snouted Late Cretaceous crocodyliform, Terminonaris cf. T. browni, from the Carlile Shale (Turonian) of Kansas: Transcations of the Kansas Academy of Science, v. 110, no. 1/2, p. 107-115.
Snyder, D., 2007, Morphology and systematics of two Miocene alligators from Florida, with a discussion of Alligator biogeography: Journal of Paleontology, v. 81, n. 5, p. 917-928.
Suarez Soruco, R., 2007, Bolivia y su paleobiodiversidad: In: 4th European Meeting on the Paleontology and Stratigraphy of Latin America, edited by Diaz-Martinez, E., and Rabano, I., Cuademos del Museo Geomienro, n. 8, Instituto Geologico y Minero de Espana, Madrid, 2007, p. 375-382.
Tanner, L. H., and Lucas, S. G., 2007, The Moenave Formation: Sedimentologic and stratigraphic context of the Triassic-Jurassic boundary in the Four Corners area, southwestern U.S.A.: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 244, p. 111-125.
Tumarkin-Deratzian, A. R., Vann, D. R., and Dodson, P., 2007, Growth and textural ageing in long bones of the American alligator Alligator mississippiensis (Crocodylia: Alligatoridae): Zoological Journal of the Linnean Society, v. 150, p. 1-39.
Vasconcellos, F. M. de., and Carvalho, I. de S., 2007, Cranial features of Baurusuchus salgadoensis Carvalho, Campos & Nobre 2005, a Baurusuchidae (Mesoeucrocodylia) from the Adamantina Formation, Bauru Basin, Brazil: paleoichnological, taxonomic and systematic implications: In: Paleontologia: Cenarios de Vida, v. 1, edited by Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M., and Oliveria, M. E. Q., p. 319-332. Link to pdf
Velez-Juarbe, J,. Brochu, C. A., and Santos, H., 2007, A gharial from the Oligocene of Puerto Rico: transoceanic dipersal in the history of a non-marine reptile: Proceedings of the Royal Society, v .274, p. 1245-1254. (Aktiogavialis puertoricensis) Link to pdf
Viohl, G., and Zapp, M,. 2007, Schamhaupten, an outstanding Fossil-Lagerstätte in a silicified Plattenkalk around the Kimmeridgian-Tithonian boundary (Southern Franconian Alb, Bavaria): Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 245, n. 1, p. 127-142.
White, M., 2009, When Crocs ruled: National Geographic, v. 216, no. 5, p. 128-141.
Wolff, E. D. S., Fowler, D. W., and Bonde, J. W., 2007, A possilbe case of necrotizing dermatitis in the crocodylian Diplocynodon, from the Oligocene of the Isle of Wight, United Kingdom: Historical Biology, v. 19, n. 2, p. 203-207.
Pterosaurs
Genus: Nova (3)
Aralazhdarcho AVERIANOV, 2007
Gegepterus WANG, KELLNER, ZHOU & CAMPOS, 2007
Ingridia UNWIN & MARTILL, 2007
Tupandactylus KELLNER & CAMPOS, 2007
Species: Nova (4)
Aralazhdarcho bostobensis AVERIANOV, 2007
Ctenochasma taqueti BENNETT, 2007b
Gegepterus changi WANG, KELLNER, ZHOU & CAMPOS, 2007
Huaxiapterus benxiensis LU, GAO, XING, LI, & JI, 2007
Synonym: Nova (1)
Tupandactylus imperator (CAMPOS & KELLNER, 1997) KELLNER & CAMPOS, 2007
= Tapejara imperator CAMPOS & KELLNER, 1997
= Ingridia imperator (CAMPOS & KELLNER, 1997) UNWIN & MARTILL, 2007
= Tapejara imperator CAMPOS & KELLNER, 1997
Tupandactylus navigans (FREY, MARTILL & BUCHY, 2003b) emend KELLNER & CAMPS, 2007
= Tapejara navigans FREY, MARTILL & BUCHY, 2003b
= Ingridia navigans (FREY, MARTILL & BUCHY, 2003b ) UNWIN & MARTILL, 2007
Note: Both Kellner, et al, 2007 and Unwin, et al., 2007 renamed Tapejara imperator. Kellner & Campos paper was publshed two months before Unwin & Martill and therefore has priority. Also, Unwin & Martill place Tapejara navigans into thier new genus Ingridia as a seperate species, though Kellner et al., 2007 referTapejara navigans to Tupandactylus imperator. I'm following both Unwin & Martill and place Tapejara navigans as a sperate species of Tupandactylus.
Averianov, A. O., 2007, Mid-Cretaceous ornithocheirids (Pterosauria, Ornithocheiridae) from Russia and Uzbekistan: Palaeontological Journal, v. 41, n. 1, p. 79-86.
Averianov, A. O., 2007, New records of Azhdarchids (Pterosauria, Azhdarchidae) from the Late Cretaceous of Russia, Kazakhstan, and Central Asia: Palaeontological Journal, v. 41, n. 2, p. 189-197. (Aralazhdarcho bostobensis)
Bennett, S. C., 2007a, Reassessment of Utahdactylus from the Jurassic Morrison Formation of Utah: Journal of Vertebrate Paleontology, v. 27, n. 1, p. 257-260.
Bennett, S. C., 2007b, A review of the pterosaur Ctenochasma: taxonomy and ontogeny: Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 245, n. 1, p. 23-31. (Ctenochasma taqueti)
Bennett, S. C., 2007c, Articulation and function of the pteroid bone of pterosaurs: Journal of Vertebrate Paleontology, v. 27, n. 4, p. 881-891.
Bennett, S. C., 2007d, A second specimen of the pterosaur Anurognathus ammoni: Paläontologische Zeitschrift, v. 81, n. 4, p. 376-398.
Calvo, J. O., Porfiri, J. D., Gonzalez-Riga, B. J., and Kellner, A. W. A., 2007, A new Cretaceous terrestrial ecosystem from Gondwana with description of a new sauropod dinosaur: Anais da Academia Brasilieria de Ciencias, v. 79, n. 3, p. 529-541.
Codorniu, L., and Gasparini, Z., 2007, Pterosauria: In: Patagonian Mesozoic Reptiles, edited by Gasparini, Z., Salgado, L., and Coria, R. A., Indiana University Press, p. 143-166.
Dzik, J., and Sulej, T., 2007, A review of the Early Late Triassic Krasiejow Biota from Silesia, Poland: Palaeontologica Polonica, n. 64, p. 1-27.
Elias, F. A., Bertini, R. J., and Medeiros, M. A. A., 2007, Pterosaur teeth from the Lage do Coringa, middle Cretaceous, Sao Luis-Grajau basin, Maranhao state, Northern-Northeastern Brazil: Revista Brasileria de Geociencias, v. 37, n. 4, p. 688-676. Link to pdf
Fastnacht, M., 2007, Tooth repacement pattern of Coloborhynchus robustus (Pterosauria) from the Lower Cretaceous of Brazil: Journal of Morphology, published on line, 17pp.
de Fuente, M. S., Salgado, L., Albino, A., Baez, A. M., Bonaparte, J. F., Calvo, J. O., Chiappe, L. M., Codorniu, L. S., Coria, R. A., Gasparini, Z., Gonzalez Riga, B. J., Novas, F. E., and Pol, D., 2007, Tetrapodos continentales del Cretacico de la Argentina: una sintesis actualizada: Ameghiana, 50 anviersario, p. 137-153.
Gasparini, Z., Fernandez, M., Fuente, M. de la, and Salgado, L., 2007, Reptiles marinos jurasicos y cretacicos de la Patagonia argentina: sup aporte al conocimiento de la herpetofauna mesozoica: Ameghiniana, 50 anivesario, p. 125-136. Link to pdf
Grellet-Tinner, G., Wroe, S., Thompson, M. B., and Ji, Q., 2007, A note on pterosaur nesting behavior: Historical Biology, v. 19, n. 4, p. 273-277.
Hargrave, J. E., 2007, Pteranodon (Reptilia: Pterosauria): Stratigraphic distribution and taphonomy in the lower Pierre Shale Group (Campanian) western South Dakota and eastern Wyoming: In: The Geology and Paleontology of the Late Cretaceous marine deposits of the Dakotas, edited by Martin, J. E., and Parris, D. C., The Geological Society of America, Special Paper 427, p. 215-225.
Hone, D. W. E., and Benton, M. J., 2007, Cope's Rule in pterosauria, and differing perceptions of Cope's Rule at different taxonomic levels: Journal of Expermential Biology, v. 20, p. 1164-1170.
Hone, D. W. E., and Benton, M. J., 2007, An evaluation of the phylogenetic relationships of the pterosaurs among archosauromorph reptiles: Journal of Systematic Palaeontology, v. 5, n. 4, p. 465-469.
Humphries, S., Bonser, R. H., Witton, M. P., and Martil, D M., 2007, Did pterosaurs feed by skimming? physical modelling and anatomical evaluation of an unusual feeding method: PLOS Biology, v. 5, issue 7, 9pp. Link to pdf
Ifrim, C., Stinnesbeck W., and Frey, E., 2007, Upper
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Welcome... to Juras- wait, no, wrong line. Welcome... to Dinopedia! This is a wiki all about prehistoric life including, you guessed it, dinosaurs. If you want to share your knowledge and passion for the time when reptiles ruled the Earth, join us today. This is a community site that anyone can edit, including you!
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It's #SciArt week on Twitter!
I think we often downplay or take for granted the role that art plays in science. High quality art is obviously a hugely important aspect of public science communication. A paper describing a new species of dinosaur will have much more impact on the public if it's accompanied by an excellent life restoration of that dinosaur. Astronomers and their spacey kin use illustrations to show us satellites, the solar system, and far-off planets we can't photograph. Biologists dealing with the very small need illustrators to show us the cells in our bodies, what's inside those cells, what DNA looks like and how it works – the list is endless.
But the #SciArt tweet storm happening this week got me thinking again about the role that art plays in my own daily scientific activities. While I don't consider myself an artist, I was always drawing while I was growing up (for a while I entertained the idea of becoming an animator!). And I'm still drawing! Every time I go to a museum, I draw pretty much everything I look at. Why draw when I've got easy access to digital photography? Well, I take tons of photos, too, but drawing makes me LOOK at the specimen.
LOOKING AROUND YOU IS VERY IMPORTANT.
Sketching slows me down, in a good way. What's that weird texture in this part of the bone, how far does this groove extend, what's with this unusual hole in this spot? Is there symmetry? Asymmetry? What's missing, and what's been filled in with plaster? What exactly was I measuring when I say 'length' or 'width'? I've filled many notebooks with drawings, stream-of-consciousness-style notes, measurements, and other bits of data. Mostly I use regular ol' pencils, but I also really like coloured pens and usually travel with a set for annotating my pencil drawings. I would love to be the kind of person that could do watercolour sketching, or proper graphite drawings.
These are some of my earliest notes from my MSc research, from a 2007 visit to the Royal Ontario Museum.
I think, as scientists, we do ourselves a disservice by not teaching students more about art skills and visual design. Being able to quickly and confidently sketch something in front of you is a useful skill to have! And understanding some of the principles of visual design – lines, shapes, negative space, colour combinations, and the like – can only make you a better communicator of science, especially in scientific papers. In addition to just being personally rewarding, drawing makes me a better scientist!
If you're a Twitterer, you should really check out the #SciArt hashtag this week (and into the future), to see the variety of techniques and approaches people take to science art.
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What's up at Wapiti River?
The world can always use some more Pachyrhinosaurus bonebeds. So hooray to my friends and colleagues Federico Fanti and Mike Burns, and my PhD supervisor Phil Currie, for publishing a description of the Wapiti River Pachyrhinosaurus bonebed (currently in 'early view' accepted manuscript form at the Canadian Journal of Earth Sciences).
A friendly Pachyrhinosaurus lakustai greets students at Grande Prairie Regional College!
Most of the time, dinosaur palaeontologists look for bones in dry, barren landscapes – the badlands of Alberta, the Gobi Desert, etc – places that have lots of rocks and not much covering them up, like inconvenient forests or cities. But sometimes, you don't have vast expanses of outcrop. In Nova Scotia, we dig up dinosaurs on the beach. In the area around Grande Prairie, Alberta, you look for bones in the outcrops along rivers and streams.
The very first summer I went out with the University of Alberta crew (way back in the halcyon days of 2007; the first Transformers movie was 'good', everybody read the last Harry Potter book overnight to avoid spoilers, and...apparently not much was happening in my musical spheres, but my, how time has flown), there wasn't a Wapiti River bonebed. We knew that there were bones coming out of the riverbank somewhere, but it took the better part of a day to trace them up the hill to the bone layer.
See if you can spot Phil for scale way up on the hill there, and remember that Phil is about 3x as tall as most humans. That's where the bone layer is!
It's a pretty steep hill, and so those first few days excavating the bone layer meant hacking out little footholds and gradually making enough of a ledge for us to sit on and walk around each other without plummeting to our death.
The last time I was there, in 2011, the ledge had expanded significantly, although you can see it's still a pretty narrow slice! It's a scenic place to work, with the river and boreal forest stretching away below; bear sightings were not uncommon (and occassionally closer than we'd all prefer), and I remember a hummingbird came down to check on us one day, buzzing around my head for a few moments!
In this bonebed, there's a layer of bones in a crazy, mixed-up layer of folded mudstones, and those are pretty easy to excavate.
Here's a dorsal vertebra. Nice and easy.
But down beneath that, the skulls and larger bones are encased within super hard ironstones. We can't really do much with these in the field, so we need to take them out in huge pieces.
And here's what the skulls look like. The circular depression down towards my left foot is the narial opening. The UALVP has like 15 of these suckers and they each take about 2 years to prepare with a crack hammer and chisel.
But the bonebed is also about halfway down into the river valley on a steep slope that's hard enough to just haul yourself up, let alone a huge boulder. So we've been very lucky to have helicopter support to carry out some of the heaviest pieces at the end of each field season.
Up, up and away!
Sometimes we were even visited by Aluk the Pachyrhinosaurus, mascot of the Arctic Winter Games in 2009!
This was probably the strangest day in the field.
There's still much more work to be done on this bonebed – we still aren't exactly sure what species of Pachyrhinosaurus is present. The age is right for P. canadensis, but only time will tell. And with two Pachyrhinosaurus bonebeds in Grande Prairie – the Pipestone Creek bonebed with P. lakustai, and the slightly younger Wapiti River bonebed – there's bound to be much more to learn about the evolution and biology of this unusual ceratopsian.
Previously in Pachyrhinosaurus:
Wapiti River Fieldwork, Part 1
Wapiti River Fieldwork, Part 2
And don't forget to check out:
Fanti F, Currie PJ, Burns ME. 2015. Taphonomy, age, and paleoecological implication of a new Pachyrhinosaurus (Dinosauria: Ceratopsidae) bonebed from the Upper Cretaceous (Campanian) Wapiti Formation of Alberta, Canada. Canadian Journal of Earth Sciences, early view.
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#MuseumWeek Retrospective!
Last week's #MuseumWeek tweetstorm was an awful lot of fun, especially following the #SciArt event just a few weeks earlier. I thought I'd share a couple of photos and thoughts for each day's theme – I didn't manage to post something for each day on Twitter, but I'll fill in some thoughts and photos here!
Day 1: Secrets
One of the nice things about working in the Paleontology & Geology Research Lab at the North Carolina Musuem of Natural Sciences is that "behind the scenes" is part of the scene. You can actually stare at me while I'm working away at my computer each day, if you desire to do such a thing. More interesting, probably, would be to watch our staff, students, and volunteers preparing fossils in the main lab space - secrets waiting to be revealed. But hey, whatever floats your boat!
If you're in Raleigh, stop by and say hi to Carnufex!
Day 2: Souvenirs
I am kind of a Stuff Person and also have a Thing for Museum Gift Shops. As such, I have loads of doodads from my various museum visits. One of the things I like picking up are postcards, especially those that have non-Tyrannosaurus dinosaurs featured on them. For a while, I had these up on my wall at my apartment in Edmonton. Those who have visited my UofA office will also be familiar with my embarassingly large collection of ankylosaur toys, or as I prefer to refer to them, 'scientific models for grown-ups'.
Recognize any museums from your own travels?
Day 3: Architecture
I had a lot of fun with this one on twitter because I LOVE interesting museum architecture. A couple of favourites:
Permian Hall at the Moscow Paleontological Museum:
...which also had custom door hinges, like plesiosaurs!
Dinosaur museum in an old castle in Lerici, Italy:
I wasn't sure about the ROM Crystal at first, but it's grown on me:
And I think the SECU Daily Planet at the NC Museum is pretty swell (on the inside, it's a theatre!):
Day 4: Inspiration
Some non-dinosaur stuff for inspiration day: I really like learning about Canadian art and its history, and one of my very favourite museums on the entire planet is the Museum of Anthropology at the University of British Columbia. If you're in Vancouver, DO NOT MISS THAT MUSEUM. It's an emotional experience to step into the exhibits at this museum and be surrounded by so much creativity and history and skill. Here's a sample to sharpen your brain.
Day 5: Family
I'm lucky to have had great parents that fed my dinosaur obsession as a kid with trips to museums near and far. I'd love to dig out some photos from the before time, but for now, I'll leave this day for my own memories. What are some of your favourite museum memories from your childhood?
Day 6: Favourites
I like busy museums that are crammed full of stuff, especially when that takes the form of a Wall of Stuff or a Hall of Stuff. Here's a few of my favourites.
Hall of Stuff at the Museo de La Plata
Wall of Stuff at the Natural History Museum of LA County
Day 7: Pose
I don't like posting pictures of myself very much, so I'll just include one here to finish off: here's Pinacosaurus (nee "Syrmosaurus") at the museum in Moscow, with me for scale.
That's it for now! What did you share for Museum Week?
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A Brontobyte of Sauropods
Palaeontology emergency alert! This is not a drill! Brontosaurus is back!
YES I FINALLY GOT TO USE THIS ON THE BLOG. Success!
I mean, Brontosaurus never really left. That's the nice thing about taxonomy – once a name is out there, it's there forever, even if we decide later on that it might represent the same kind of animal that another name does. And so every now and then, we get to bring an old name back from the dead. Today, Tschopp and colleagues have published some very good support to indicate that Brontosaurus really is distinct from Apatosaurus after all, and we can all use that name and stop telling people that Brontosaurus isn't real. OMG, WHAT A RELIEF.
To recap: Brontosaurus has not been an accepted name in the palaeontological community for more than 100 years, but because of its use in some museum exhibits, and things like the 1964 World's Fair and the "Rite of Spring" passage in Fantasia, for example, the name has become entrenched in the popular consciousness in a way few other dinosaur names have. It is very disappointing to learn that palaeontologists don't call that big dinosaur Brontosaurus, but the decidedly less evocative name Apatosaurus instead.
Click for sauropod-size. With many thanks to the authors and PeerJ for creating such a useful diagram, which I'm sure will be reproduced often and with much gratitude by palaeontologists, teachers, and other science communicators.
The new paper is staggering in its length (almost 300 pages!) and the amount of work it represents, and I'm not a sauropod specialist, so I'll summarize it here without delving into sauropod anatomy very much:
Two of the Big 3 diplodocids: Apatosaurus (in the back) and Diplodocus (foreground) face-off at the Carnegie Museum.
Tschopp et al. did a specimen-level phylogeny of diplodocids, the sauropods like Apatosaurus and Diplodocus, but not Brachiosaurus or Camarasaurus. This means that individual specimens were coded, rather than species. Often, phylogenetic studies have just looked at the 'classic' diplodocids Apatosaurus, Barosaurus, and Diplodocus (the 'Big 3', shall we say?). And most of those studies elide the many species represented by these three genera. So a specimen-level phylogeny is a much-needed approach to resolve some questions about diplodocid diversity.
They then used some techniques to quantify differences among specimens – pairwise dissimilarity, and apomorphy counts – that would help justify dividing clusters of individuals into different genera. There isn't a rule in palaeontology that individuals need to be a certain amount 'different' from each other in order to be a new genus or species, so the authors looked at how many unique characters separate some sauropods that everyone seems pretty comfortable calling different species and genera. Apatosaurus ajax and Apatosaurus louisae had 12 different features, and Diplodocus carnegii and Diplodocus hallorum had 11 different features. So 13 different characters was set as the baseline for separating out genera in the specimen phylogeny. Using the same approach, they also set 6 differences as the baseline for separating species within a given genus. These numbers only apply to this particular analysis, but it's an interesting approach that I think would be worth considering for other dinosaur phylogenies.
Using this, they wind up doing some taxonomic reshuffling:
a.Diplodocus longus lacks any diagnostic features at the species level and is a nomen dubium, which is bad because it's also the type species for Diplodocus. A petition to the ICZN to switch the type species to D. carnegii is in the works. Diplodocus includes the species D. carnegii and D. hallorum (née Seismosaurus)
b.Dinheirosaurus (from Portugal) is a junior synonym of Supersaurus, and so Supersaurus is a cross-continental genus represented by two species.
c.Diplodocus hayi passes the threshold for generic distinctiveness from Diplodocus and gets a new name, Galeamopus hayi. Specimens of Galeamopus are actually more complete than Diplodocus, which means that Diplodocidae is best represented by Galeamopus at present if you need a diplodocid for whatever you're working on.
d. And finally, and arguably most significantly, Brontosaurus passes the threshold for generic distinctiveness from Apatosaurus. There are three species within Brontosaurus: B. exelsus ('classic'Brontosaurus), B. parvus(née Elosaurus), and B. yahnahpin (née 'Eobrontosaurus').
The third of the Big 3 diplodocids, the iconic rearing Barosaurus at the American Museum of Natural History.
I really hope this taxonomic shuffling gains wide acceptance, because 1) I think their approach and reasoning are pretty sound, and 2) it's going to be SO MUCH EASIER not to have to constantly 'debunk'Brontosaurus with non-palaeontologists.The oft-repeated story that "Brontosaurus" wasn't real because it had the head of one animal and the body of another is wrong, but the real story, about the rules of taxonomy and how we define species, is much more difficult to explain. (It's interesting, but it's not as easily parsed to a lay audience.) And let's face it, Brontosaurus was a really good name and it was sad that it had to be synonymized. The story of Brontosaurus now has a new and interesting chapter – our ideas about the biology of Brontosaurus have changed, but now we can talk about changes to how we think Brontosaurus looked and lived, rather than just focusing on a quirk of taxonomy. So let your Brontosaurus flag fly high, dinosaur fans, because Brontosaurus is back and that's awesome.
Old-timey sauropod in the little diorama at the Smithsonian, back in 2011.
Big taxonomic revisions are hard and important but often don't feel as 'sexy' as some of the other research that gets publicized. I like thinking about alpha taxonomy (uh, perhaps obviously) and I like doing this kind of research, and I think it's really important that we recognize how important this kind of work is – alpha taxonomy is really foundational to a lot of other studies. If you don't know how many species you have, or where they lived, or what anatomy belongs with each species, how can you do projects that look at the evolution of certain features through time, or understand changing ecosystems?
For example, given that there's at least 14 species of diplodocid in only 11 million years of Morrison Formation, it's unlikely that there's a slice of time in there in which there's only one diplodocid species. (And remember, diplodocids weren't the only sauropods in the Morrison – this is also the home of Brachiosaurus and Camarasaurus and Suuwassea and who knows what else.) This is a pretty good reason to reject what I like to call the "Highlander hypothesis", i.e. There Can Only Be One ___(ankylosaur, tyrannosaur, whatever)___ in a given formation, something that I've encountered in conversations on occasion. It's understandable that we would feel unease at the idea of high species/generic diversity in such massive dinosaurs, because how are they dividing up ecosystem space? But over and over again it seems like lots of similarly-shaped dinosaurs were occupying similar times and spaces in terms of what we see in the rock record, which I find very interesting indeed. (Now what we need is a really good stratigraphic framework for putting all of these diplodocids into chronological and geographical context.) We can only do a good job of addressing these kinds of questions by having good data to put into those studies, and that data comes from taxonomic revisions like this one.
And revising taxonomy is probably a never-ending job, because we need to keep reassessing our definitions of genera and species as we get more information through new specimens. Let's make sure we all support this kind of research as palaeontology continues to evolve with new techniques, questions, and approaches. Bully for Brontosaurus, and bully for alpha taxonomy.
Stray observations:
The concept of a 'relatively small' animal that is 12-15 metres long amuses me. (re: Kaatedocus, page 2)
The 'brontobyte' image at the top of this post is an old joke from my Currie lab days; a brontobyte is actually 10^27 bytes. But I think it would be a good collective noun for sauropods, and it also feels appropriate given the large number of sauropod species recovered by Tschopp et al. In fact, we need more collective nouns for dinosaurs, and so I'd like to propose brontobyte for sauropods and armada for ankylosaurs, to join terror of tyrannosaurs.
Go read the paper! It's open access!: Tschopp E, Mateus O, Benson RBJ. 2015. A specimen-level phylogenetic analysis and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda). PeerJ 3:857.
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May your mountains dark and dreary be.
Just wanted to give a quick shout out to some old fossil friends of mine. Horton Bluff/Blue Beach is a pretty cool place and I have fond memories of field trips out there during my Dalhousie days. Between this new paper and the recent paper describing the Permian to Jurassic assemblage of tetrapods, it's been a good time for Nova Scotia palaeontology.
Your friendly neighbourhood ankylosaur palaeontologist, in the before time (i.e. 2003), at Horton Bluff, following in her tetrapod ancestor's footprints. It's goopy there.
Which of course makes me miss it all terribly.
Anderson JS, Smithson T, Mansky CF, Meyer T, Clack J. 2015. A diverse tetrapod fauna at the base of 'Romer's Gap'. PLOS ONE 10:e0125446.
Sues H-D, Olsen PE. 2015. Stratigraphic and temporal context and faunal diversity of Permian-Jurassic continental tetrapod assemblages from the Fundy rift basin, eastern Canada. Atlantic Geology 51:139-205.
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On Surprises
I love surprises. Which is unfortunate for me, because I am extremely bad at being surprised. And it's hard to be surprised by things as you get older, and as easier access to more and more information becomes available to us every day.
But boy, when a surprise comes along that actually takes me by surprise, what a thing to be able to savour.
An extraordinary painting of Yi qi by Emily Willoughby, CC-BY.
So, enter Yi qi. In many ways, it's hardly a surprise at all – numerous artists produced wonderful speculative art about scansoriopterygids predicting membranous wings and/or gliding abilities, and there was even this neat hypothetical Archaeopteryx ancestor that I found in a paper a few years ago. At the time, I wrote on Facebook: "I had not realized that a bat-winged proto-bird was an idea that was on the table!"
(I also wrote, "I like his smirk, lack of neck, and skinny skinny tail", and I agree with Past Victoria about all of those things.)
While Yi qimight not really be a proto-bird, it's still an amazing discovery that shows there was a lot of experimenting with flying and gliding going on back in the Mesozoic, which is perhaps unsurprising, given that lots of disparate groups of animals use gliding to their advantage today – fish, frogs, rodents, marsupials, dermopterans, you name it. And yet, even though there's lots of precedent for gliding vertebrates, and others had predicted something kind of like Yi qibefore, I was still genuinely shocked when I saw the paper and press images. What a great feeling.
What I love best about Yi qi, apart from it's extremely meme-able name, is that it's a great example of maybe what I'll call an 'expected surprise'. A surprise that, as soon as you see it, it seems so obvious and like it should have been there all along. It's like the opposite of a failure of imagination. Surely there is a long German word that captures this specific emotion? What other expected surprises are lurking out there in our futures? What things have we speculated on today, dismissed as being way too out there to take seriously, and yet will pop up as really-for-real things later on?
I guess I need to get to work on some ankylosaur speculative biology! Maybe we'll find the real Yee?
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On Failures of Imagination
Yesterday I talked about 'expected surprises' with regards to Yi qi. Yi qi is a surprise because its anatomy is so unlike other theropods, and it suggests that dinosaurs were experimenting with flight and/or gliding in some ways that were quite different from our current understanding of feather and bird wing evolution. But it was also not entirely unexpected, because scansoriopterygids had super weird anatomy to begin with that gave us enough information to speculate about possible gliding adaptations in those dinosaurs, even though the general consensus was that it was pretty far-fetched.
But today I wanted to talk about a related feeling, which I like to call the Failure of Imagination. Last summer I was working my way through a DVD set of classic sci-fi, fantasy, and adventure movies that I had picked up at some point. I wound up watching a lot of these with friends and basically Mystery Science Theatre 3000-ing the films, and in particular the old space adventure movies from the 40s-60s provided much entertainment. It's really fun to take a look back and see what sorts of things people envisioned the future holding for us – space travel, exoplanet exploration, robots. But what also struck me was the things that the filmmakers and storywriters couldn't even imagine.
They could imagine spaceships and robots, but they couldn't imagine wireless technology. Or storing information in digital form rather than on spools of tape.
They couldn't imagine non-button-and-dial-based instrumentation.
And they definitely couldn't imagine women in roles other than administrative assistants (or as the bad guys). SO MANY SPACE SECRETARIES.
I kept thinking to myself – what sorts of failures of imagination are we having in palaeontology today? We can imagine so many things. But I wonder what kinds of things we won't even know we don't know. When we try our hand at speculative biology, what will scientists 80 or 100 years from now think was charming, or quaint, or ahead of its time. Failures of imagination are one of those things that make me nervous as a scientist, because I don't like the idea that I won't even know what I'm not imagining.
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Crystal Geyser Quarry Quest
I just got back from my first stint of fieldwork for the year, and my first time doing fieldwork in the States. This was just a brief jaunt out to Utah and Colorado for two weeks, but it was a nice sampling of some interesting and different field localities compared to my previous experiences. Today's post: Crystal Geyser quarry in Utah!
So scenic, so majestic. Such altitude.
Crystal Geyser is a non-geothermal, carbon dioxide geyser near Green River, Utah; although we didn't visit the geyser itself, it lends its name to a series of quarries of a massive bonebed in the Yellow Cat Member of the Cedar Mountain Formation (about 125 million years ago). The bonebed is mostly composed of the early therizinosaur Falcarius. The bones in this quarry are incredibly delicate - sometimes even using just a brush through the sediment felt like it was too aggressive! Definitely a challenging site to work at.
Ominous clouds brewed up frequently and then dumped rain and hail on us.
But then sometimes there were rainbows, so I guess it was ok.
We camped in the Morrison Formation and walked up to the Cedar Mountain Formation each day, which was kind of fun.
I'm not accustomed to walking through such a dramatic shift in time and faunas: the Morrison is characterized by lots of classic dinosaurs like Allosaurus and Apatosaurus and Stegosaurus from about 156 to 146 million years ago, but the dinosaurs of the Cedar Mountain Formation have only recently begun to receive much attention and are still poorly known. There's a gap of about 20 million years between the two formations, and in the Yellow Cat Member we find dinosaurs like Falcarius, the ankylosaur Gastonia, the iguanodontians Hippodraco and Iguanacolossus, and dromaeosaurs like Utahraptor and Geminiraptor. The world was changing.
We'll be returning to Utah later in July to work in the Mussentuchit Member. Up next: jackhammering in the Mesaverde Formation of Colorado!
Epilogue: I made a friend at lunchtime one day. D-:
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Mad Max: Fury Quarry
There's a hadrosaur underneath this cliff. (Probably.)
For the second week of our field expedition, we drove from Utah to a site near Rangely, Colorado, to help the Colorado Northwestern Community College with a specimen poking out of a cliff. We've shifted into the Upper Cretaceous here, and are working in the Mesaverde Formation. There's not very much known about the dinosaurs from this formation, so hopefully this will shed some more light on the dinosaurs in this region!
In between Utah and Rangely, we spent a night in Grand Junction and went to see Mad Max: Fury Road, which was way more awesome than any of us had really anticipated and which was basically all we could talk about all week. And so I must also share this great photo that Lindsay took! According to buzzfeed, my Mad Max name is Roop Duststorm, which seems appropriate given the dustiness of working around a rock saw all day.
I did a lot of jackhammering last week, which was great fun if terrible for my lower back, but my favourite thing to do is to pop off the blocks made using the rock saw. You cut a grid into the rock, position your chisel at just the right point, give a couple of hearty whacks with a crack hammer, and off pop these incredibly satisfying 'brownies' of sandstone. It's still slow going, but you can move a lot of rock a lot more quickly this way. (Thanks to Lindsay again for snapping this fun photo!)
Early in the week we were plagued with constant large thunderstorms that rolled in every few hours and made things kind of cold and miserable. Thankfully, this was the last one and it missed us! Instead, it just looked dramatic, which is fine by me.
By last Saturday we had made a lot of progress, although there is still a long way to go to get down to the bone level and (hopefully) find a good dinosaur down there. Best of luck to the crew as they keep working furiously away!
*'Fury Quarry' is also shamelessly stolen from Lindsay.
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Cornelius says hello
Say hello to Cornelius! I got to meet him during a brief visit to the ROM last week, and he seems like a pretty nice guy.
This cool new ceratopsian is on display in the Age of Dinosaurs gallery in an exhibit called "New Dino Discovered", and was also featured in Dino Hunt Canada, which aired earlier this year. It should have a new name soon, but for now Canada voted to nickname it Cornelius. The really nice skeletal mount was put together by Research Casting International based on about a 50% complete disarticulated skeleton.
Here's a close-up of that winning smile. This new dude is a centrosaurine ceratopsid with some pretty neat ornamentation going on at the back of the frill.
I really liked the inclusion of a quarry map on the floor, which highlights some of the bones that are on display. The skeleton was found in southern Alberta in the Milk River area, and comes from the Oldman Formation.
The mounted skeleton is a cast, but there are some original bones on display, like the radius and ulna shown here.
In particular, I liked this set of panels on the wall showing differences in frill ornamentation between centrosaurines, and how we identify different species. On the right is the original frill material for Cornelius, the bottom left is Centrosaurus, and the top left is Styracosaurus.
And look, there was even an ankylosaur osteoderm on display! These are some of the fossils found in the Milk River area, which tell us a bit about the ecosystem that the new centrosaurine lived in.
It's a cool new dinosaur and a nice exhibit, so definitely don't miss it if you're visiting the ROM anytime soon!
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Why does Jurassic World hate dinosaurs?
I have some Thoughts and Feelings about Jurassic World! Spoiler alert, I'm going to talk about details and plot points and this post is really for people who have seen the film. Also, while I'm going to talk about the dinosaurs a bit, this isn't really a review of the science of the film, because that's already been done to death. Ok, onwards and upwards into something that wound up being way too long!
Does Jurassic World hate dinosaurs?
I think the answer to that question is yes. Jurassic World keeps making these little homages and throwbacks to the earlier films (there are lots of shots that echo iconic moments in the earlier films, and some of the plot points mirror the original film almost exactly), and yet I feel like we could consider the theme of Jurassic World to be about rejecting nostalgia and childhood. It's buried under an interesting discussion of the role of the military in funding scientific research, and why some kinds of research are prioritized over others, and it may actually be unintentional, but it's the theme I took away most immediately from this film.
There are two characters that I think are supposed to represent the audience, and neither are treated particularly well by the other characters. And by 'the audience', I'm going to be really self-centered and say that I mean the 30-somethings like myself who saw the original film when we were in that 8-12 year old bracket, or 'peak Jurassic Park' age, and who this film is clearly pandering to. Firstly, we have Gray Mitchell, a 10-ish year old who represents us when we first saw Jurassic Park: he's a dinosaur geek and is one of the only characters to show unrelenting enthusiasm for dinosaurs while visiting Jurassic World. Secondly, we have Lowery, the 30-something computer room dude, who wears an original Jurassic Park shirt and has dinosaur toys on his desk and is obviously super into the dinosaurs in the dinosaur theme park. He is us, now, grown up and nostalgic for the original film. Multiple times throughout the film, Gray's older brother tells him he needs to grow up, and points out that many of the things are for little kids. Claire makes fun of Lowery's shirt, and I think in general we're supposed to think he's kind of a weird man-child who hasn't really grown up.
There's a moment in the film where Gray and his brother Zach stumble upon the old Jurassic Park visitor center building. The T. rex cast skeleton lies on the ground covered in vegetation, and a little piece of the "When Dinosaurs Ruled the Earth" banner is visible. Zach uses it to make a torch so they can investigate the rest of the suspiciously-well-lit ruins. Visiting the old building felt like some gratuitous fan-service to me, but then burning the banner felt like a purposeful statement about rejecting the nostalgia of the original film.
Jurassic World is constantly setting up little nostalgic moments and then seemingly stomping all over them. It's like the filmmakers wanted to pay tribute to Jurassic Park but then were embarassed to show that they liked it – or maybe they didn't really like that movie at all, but wanted to make lots of money (success!). I don't know, but I find it thematically problematic and a bit sad, since the excitement over DINOSAURS! in the first movie is one of the defining aspects of that film, and that sense of wonder and grandeur has rarely been replicated. Jurassic World feels jaded, and like it's too cool for dinosaurs.
Can we talk about ladies in this movie for a moment?
Did we really need to introduce our main female character with the camera sweeping up her legs to her face? Was that absolutely necessary? Also, could we just not use the 'frigid, uptight workaholic woman needs to learn to loosen up and become sexually free with a man, and also needs to remember that all women will have children eventually' stereotype? COULD WE JUST NOT?
It's an intriguing throwback to the original Jurassic Park movie, which I feel successfully used the kids as a character development point for Alan Grant. But Sam Neill managed to portray Grant's discomfort with kids in a more organic way, and the movie gave that plotline a bit of breathing room to develop during some of its quieter moments AND its action sequences (see: sitting in the tree feeding Brachiosaurus; escaping the falling car in the tree; the fence). It's less believable with Claire Dearing, because she doesn't even spend any time with the kids in peril until almost the very end of the movie, at which point she basically worried herself into liking kids? Or something?
Look, not every movie is going to have (or should have) a Strong Female Character(TM), because there are lots of ways to be a lady just like there are lots of ways to be a dude. But the first two Jurassic Park movies had some cool female characters: Ellie Sattler, a palaeobotanist, who was brave and curious and smart! Lex Murphy, who knew those UNIX systems! Sarah Harding, who was a bit foolish but was also brave and curious! Kelly Curtis Malcolm, who gymnastic-ed a Velociraptor to death! In Jurassic World, we get a woman who has great power and authority (she runs a theme park full of dinosaurs!) being told she should be different at almost every opportunity, and we get a distracted babysitter who is killed in the most gratuitous, drawn-out sequence of all. Thanks, movie.
Ok, now let's actually talk about dinosaurs (and other prehistoric creatures) in Jurassic World.
Other palaeontologists have already beaten me to much of this, but I still had a few thoughts I wanted to share. Ultimately I don't have a big problem with the 'retro' dinosaurs of 1993 appearing in this film, because I'm willing to go with the flow in terms of continuity. But there were some pretty dumb things in this film:
· The pterosaur sequence was pretty godawful and brought the action to a screeching halt. I can't suspend disbelief that the pterosaurs would immediately rampage and murder a bunch of people, and I can't suspend disbelief over the physics of that sequence. Refrigerating that babysitter lady was also pretty awful. Sweet jeepers, Jurassic World, you're going to make me say something horrible: this sequence was better in Jurassic Park III. THERE. I hope you're happy.
·I never really bought Indominus rex as anything more than a really big Allosaurus or Saurophaganax. (Sorry, theropod people! Allosaurus is cool, but not, like, THAT cool.) I did, however, like the incorporation of the camouflage idea from the Carnotaurus in the Lost World book, something that I had missed from the film adaptation. Overall, I'm frustrated that Indominus exists mostly so they had a dinosaur they could trademark. Because that's totally what that is, and everything else is secondary to that, including its incorporation into the plot.
·That mosasaur is just so gigantic. I'm on board, but that was starting to stretch credulity as well.
·Why doesn't Rexy eat Blue after the fight? The mind boggles.
Ok, things I liked!
·The Ankylosaurus gives Indominus the old what-for and doesn't immediately die like everything else! Indominus needs to really work at murdering that poor fellow. The design of the Ankylosaurus themselves is pretty terrible (wrong osteoderms, tail too curly, nostrils in the wrong spot, head generally a bit off), although I think it's meant to be consistent with Jurassic Park III.
Here's what Ankylosaurus REALLY looks like!
·Dinosaur petting zoo! It should be for all ages!
·The big kaiju battle between Indominus and Tyrannosaurus was pretty well matched. I liked the little kick to JPIII when the Tyrannosaurus busts through the Spinosaurus skeleton on the way to the fight.
·"Are they safe?""Oh no, under no circumstances, not even a little."
Some final Thoughts and Feelings
I haven't decided yet if I liked Jurassic World. I can't help but think back to the original Jurassic Park with its iconic visual moments and charming, if hokey, dialogue. While it was fun to see an operational Jurassic Park with rides and attractions, I don't feel like Jurassic World had much visual flair. It's really hard to beat dramatic, symbolic visuals like this:
Interesting camera angles like this:
Or quiet moments of terror like this:
And I miss the yellow and green and red colour palette of the original park, replaced here with chrome and blue and silver like every other washed out movie in theatres lately. It is also interesting that all of the big sweeping themes from the original soundtrack are used not for the dinosaurs, but for the manmade structures of the park itself. It really does feel like Jurassic World doesn't care about dinosaurs.
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Dinosaurs Unearthed!
Growing up in Nova Scotia, despite its many excellent and significant palaeontological treasures, meant that there weren't many dinosaur fossils for me to gawp at regularly. The Nova Scotia Museum of Natural History (which I loved) had only a few small fossils on display, and besides an exciting appearance by the Dinosauroid when I was small, did not have any big traveling dinosaur exhibits come through. But when I was in Grade 1 or so, DINAMATION came to town and seared its robotic dinosaurs all over my brain forever. And so I think I will forever have a soft spot in my heart for animatronic dinosaur displays.
Mike Burns and I ran into some of the old Dinamation robots puffing away at the New Mexico Museum back in 2012!
Like Jurassic Forest and Dino Dino Dreampark, the traveling exhibit Dinosaurs Unearthed (at Telus World of Science in Edmonton for the summer) mostly features large animatronic dinosaurs, as well as some casts and interactive displays.
One of the main focuses of the exhibit is showcasing Chinese dinosaurs and fossils, and talking about recent research on the evolution of birds from dinosaurs. Probably one of the best parts of the exhibit is the large number of casts of feathered dinosaurs from China, including Sinosauropteryx, Caudipteryx, Microraptor, and Confuciusornis. These are still not particularly household names, so it's nice to see these on display, especially given the lack of feathers in Jurassic World's dinosaurs!
Just past the casts we have a diorama of Jehol Biota feathered dinosaurs, including (from left to right) the dromaeosaur Microraptor, the compsognathid Sinosauropteryx, the tyrannosaur Dilong, and the bird Confuciusornis. Some of the animatronics are better than others, and all are kind of weirdly oversized, but I think if there was a sign that said this was a diorama at 4x life size or something like that, that it would work pretty well.
My favourite cluster of dinosaurs was the set of Mongolian dinosaurs, including the first time I've ever seen Gigantoraptor anywhere! There's also a pretty dapper Alxasaurus in the front there.
I would have liked to see more cast fossils rather than sculpted reconstructions, and perhaps more fossils overall and a couple fewer animatronics. But generally the information presented in the exhibit was pretty good and had been recently updated, with references to the new research on Brontosaurus, and lots of recent behavioural, biomechanics, and ecology facts as well. Here's a nice display showcasing some of the cool imaging work done by the WitmerLab!
Until next time...watch out for that Shantungosaurus as you leave!
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Woe betide those who summon the Galactic Coelacanth
A couple of years ago I had an existential crisis when I realized that, in the time one of my papers had been in review (almost 8 months!), I could nearly have physically created an entirely new human being in my body, if I had so chosen. Thus began the saddest game in the universe that I like to play when I submit a paper: "What kind of animal could have been gestated in the time this paper has been in review?". And this became an even better running joke when one of my colleagues had a highly unusual review experience that lasted for several years, which completely exhausted the gestation times of real animals.
My amazing and lovely sister saw us talking about this on Facebook and went ahead and wrote an R script that tells you exactly what kind of animal you could have birthed while waiting for reviewer comments. And because I am always forgetting to save this amazing piece of code, I've gotten permission from Jessica to post it here for posterity. My sincere apologies to anyone who gets the Space Whale, and my deepest condolences to anyone who is graced by the presence of the Galactic Coelacanth.
Click here for the R script!
Updated 30 June 2015: If you don't have R, you can also download a text file to see the code!
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Know Your Ankylosaurs: China Edition
I'm in Utah digging up dinosaurs! But also, one of the last big chunks of my PhD thesis has just been published online at the Journal of Systematic Palaeontology. They are generously allowing free access to the paper through the end of August, so head on over and grab a copy while it's free! This time, I'm taking all of the knowledge gained from my previous taxonomic revisions, adding in some more taxa, and doing a revised phylogenetic analysis building on previous analyses to see how everyone shakes out and to learn a little bit more about ankylosaurid biogeography. I'll cover some of the taxonomic stuff over the next few posts, and finish off with the big picture of ankylosaurid evolution.
Pinacosaurus!
I've talked previously about the ankylosaurs of Mongolia, but I've also had the opportunity to study some of their friends from across the border in China. In particular, I got to see lots of specimens of Pinacosaurus, both from the Alag Teeg bonebed in Mongolia, and from Bayan Mandahu in China. Because Pinacosaurus specimens are relatively abundant and usually well preserved, there has already been lots of descriptive work on this taxon, including on the skull (and here, and here), hands and feet, and general postcrania.
Baby Pinacosaurus are so teeny tiny! This one is from Bayan Mandahu and was collected during the Canada-China Dinosaur Project back in the 1980s.
I've discussed just a few new points about Pinacosaurus, especially about how we tell the two species of Pinacosaurus apart. Pinacosaurus grangeri is known from lots of specimens, almost all of which are juveniles; it has relatively short horns at the back of its skull, a constriction in the snout between its nose and its eyes, and a notch in the rough ornamentation above each nostril. Pinacosaurus mephistocephalus is known from just one specimen (also a juvenile), and it has long squamosal horns, no constriction in its snout, and no notch in the ornamentation above each nostril (it looks like it does on one side, but I think this is just damage given that it is not present on the other side). Both species are known from Bayan Mandahu, and so it is reasonable to ask whether or not these could represent the same taxon – given the differences in skull morphology, I suspect we're not looking at intraspecific variation here, although more specimens of P. mephistocephalus would be very helpful in this regard!
Crichtonsaurus becomes Crichtonpelta
Crichtonsaurus is another cool ankylosaur that has received surprisingly little attention given its Jurassic Park affinities. Two species have been named: Crichtonsaurus bohlini (the type species), and Crichtonsaurusbenxiensis. Crichtonsaurus bohlini is, unfortunately, a very incomplete jaw that does not bear any diagnostic features, and so we argue that Crichtonsaurusshould be considered a nomen dubium. Crichtonsaurus benxiensis, on the other hand, is a great specimen with a really good skull and a fair bit of the postcrania, and the skull has some unique features that make it easy to distinguish from other taxa, most specifically the upturned quadratojugal horns. We've proposed the new name Crichtonpelta benxiensis for this material – Crichtonsaurus was a good name and we wanted to keep the replacement name similar, so now we have Crichton's shield instead of Crichton's lizard.
During the Flugsaurier symposium in 2010, while I was visiting Beijing and the IVPP, we took a field trip out to Liaoning and visited the Sihetun Fossil Site. It has a cool museum, including a mounted Crichtonpelta skeleton! I don't think this specimen has been described, but it does corroborate certain aspects of the holotype skull. Crichtonpelta seems to lack discrete caputegulae (tile-like ornamentation) on its skull, which gives it a similar appearance to Pinacosaurus. I don't think the osteoderms have been placed quite correctly on this skeletal mount – I think they've been tipped on their sides so that the keel forms part of the 'base', giving it a somewhat stegosaur-like appearance.
Liaoningosaurus and Chuanqilong
I'm going to talk more about Liaoningosaurus in a few months, but it is one cool little ankylosaur! At only about 30 cm long, the holotype is one of the smallest known ankylosaur specimens and probably represents a very young individual. There may be a few osteoderms in the cervical/scapular region, but that's about it. I've previously argued that the putative plastron in this specimen is more likely skin impressions, which is still pretty cool because we don't have a lot of belly skin for ankylosaurs.
Liaoningosaurus! YAY!
I also wanted to give a shout out to here to Chuanqilong, a larger ankylosaur from Liaoning that was described last summer and which didn't make it into my thesis but which I did include in the revised phylogenetic analysis in the final paper.
Here's Chuanqilong, from Han et al. (2014).
Dongyangopelta, Taohelong, and Sauroplites
Let's finish off this post today with a triad of interesting but enigmatic ankylosaurs. Dongyangopelta and Taohelong are relatively new entries to the world of ankylosaurs, with both taxa appearing in 2013. Neither are particularly complete, but they are interesting because both species possess chunks of fused osteoderms, which would have been found over the pelvis and which are most commonly encountered in nodosaurids and 'polacanthids/polacanthines', and are presently unknown in ankylosaurids – and indeed, Yang et al. described Taohelong as the first example of a polacanthine from Asia. Nodosaurids (including 'polacanthines' as basal taxa within this clade) have been tentatively identified from Asia previously (an interesting but fragmentary specimen from Japan may be a nodosaurid), but to find a Polacanthus-like animal in Asia is unexpected and very interesting. The two species can be differentiated based on the morphology of these pelvic shield pieces. Dongyangopelta comes from the Chaochuan Formation, and another ankylosaur, Zhejiangosaurus, had been named from that formation in 2007; it may eventually shake out that Dongyangopelta is a junior synonym of Zhejiangosaurus, but in the absence of overlapping diagnostic material we opted to keep these taxa separate for now.
Pelvic shield fragments - Dongyangopelta redrawn from Chen et al. (2013), Taohelong redrawn from Yang et al. (2013), and Sauroplites redrawn from Bohlin (1953).
Sauroplites, on the other hand, is a very old name that has been largely overlooked in recent assessments of ankylosaurs. The material was originally described by Bohlin in 1953, but sadly the whereabouts of the original material is unknown today (although there are casts at the American Museum of Natural History). I think Sauroplites was overlooked for a while because it's based off of osteoderms alone, and it's hard to assess diagnostic characters in osteoderms sometimes because they vary so much along the body. This is partly why I like cervical half rings and pelvic shields – in these structures, you can understand the positions of the osteoderms on the body and directly compare patterns and morphologies across different taxa. Supposedly, the osteoderms for Sauroplites were preserved in their original positions when the specimen was excavated, and if so, it's a bit surprising that more of the skeleton was not preserved. Bohlin correctly identified some of these pieces as elements of the sacral armour, and the morphology of these pieces can be used to differentiate Sauroplites from Taohelong and Dongyangopelta, and we consider Sauroplites to be a valid, but poorly known, taxon. It's good to revisit poorly figured and fragmentary taxa from time to time, because new discoveries might help put those pieces in context.
Next time: we head south! See you then!
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Know Your Ankylosaurs: Gondwana Edition
Last time, I talked about the ankylosaurids of China, and today we're talking about Gondwanan ankylosaurs. Gondwana basically refers to the continents of today's southern hemisphere; when the supercontinent Pangaea broke apart, it split into two large continents – Laurasia in the north, and Gondwana in the south. Gondwana includes South America, Africa, Australia, and Antarctica, and, somewhat nonintuitively, India (India kind of beelined into Asia from Australia and that's why we have the Himalayas). Almost all of the ankylosaurs we know about are from the Laurasian continents, which means that the few found in Gondwana are phylogenetically and biogeographically interesting: do they represent southern branches of the ankylosaur family tree, or new migrations into Gondwana from Laurasia? Let's take a closer look:
Minmi paravertebra and Minmi sp.
Minmi is the iconic Australian ankylosaur. Most people, when they think of such things, think of the spectacular referred skeleton with agood skull and in situ armour.
The Smithsonian has a cast of the specimen - here's a section of the ribcage, showing some of the osteoderms in their original arrangement.
Sadly, the holotype is extremely fragmentary and has few elements to make a diagnosis with. Originally, one of the most striking features of Minmi paravertebra was the presence of paravertebral elements, thin rod-shaped bones along the dorsal vertebrae. These were originally interpreted as ossified tendons of the dorsal muscles, and although these are cool to see in Minmi, they are not really unique to Minmi or even to ankylosaurs, since ossified tendons are ubiquitous throughout Ornithischia. One unusual aspect of these ossified tendons is that one set has a flattened, expanded front end. These were interpreted as possible ossified aponeuroses (aponeuroses are sheets of connective tissue in between muscles and tendons). This particular aspect of the ossified tendons IS very unusual, because ossified aponeuroses are extremely rare in animals. While I was hunting around for information about ossified aponeuroses, I came across a very odd case study about mouse deer (Tragulus)– the males completely ossify the aponeuroses above their pelvis and back, creating a carapace-like structure! This is super weird and I would love to investigate this further at some point.
Ossified aponeuroses have since been identified in the European nodosaur Hungarosaurus, which poses a bit of a problem for Minmi: since this feature was one of the only diagnostic characters for Minmi, and since it is now found in an animal that is very unlikely to be Minmi given the spatial and temporal distance between the two, Minmi paravertebra is left without diagnostic characters. A sticky situation that will hopefully be resolved in the future by people who have spent time with the original fossil material!
Antarctopelta
Did you know that the first dinosaur discovered in Antarctica was an ankylosaur? Cryolophosaurus might get all the buzz, but Antarctopelta was first to the press. Antarctopelta is a very interesting little ankylosaur, which I had the chance to study during my visit to Argentina back in 2011. The material is fragmentary but tantalizing, with some pieces of the pelvic armour that are reminiscent of ankylosaurs like Stegopelta and Glyptodontopelta from North America. Unfortunately, in the course of my research I noticed that some of the bones attributed to Antarctopelta and used to help diagnose the taxon didn't quite seem like they came from an ankylosaur. The material was found on an ancient beach strandline with some marine fossils mixed in, and it looks like some of the material originally interpreted as ankylosaurian might be better interpreted as belonging to a mosasaur and a plesiosaur. In the end, we weren't left with any diagnostic characters for Antarctopelta and we should consider that a nomen dubium for now, but there's definitely an Antarctic ankylosaur and I hope at some point some better material is recovered so we can determine the best name for this guy.
The Argentinian ankylosaur
Finally, I also had the chance to study the only described ankylosaur from Argentina. This is also a fairly fragmentary specimen, and it came from a channel lag deposit so it's possible that more than one individual is represented. There are osteoderms, some vertebrae, and a femur, and all are very small – about the same size as the juvenile Anodontosaurus (originally described by Coombs as Euoplocephalus) from Alberta. The femur is interesting because it has some very prominent ridges running lengthwise on it, which seem to be intermuscular lines; these are present but very faint on some other ankylosaurs, and I haven't encountered anything like that in other ankylosaurs. There also may be fragments of the cervical half rings preserved as part of this specimen, since there are some unusual curved osteoderms with multiple peaks and keels. These don't bear any resemblance to other half rings I've looked at, and cervical half ring morphology seems to be taxonomically informative for ankylosaurs. Together, the weird intermuscular lines and unusual cervical half ring fragments might be enough to diagnose the Argentinian specimen as a new taxon, although we withheld from doing so at present.
Here's the specimen on display at the Museo Carlos Ameghino in Cipoletti!
There have been reports of some possible ankylosaur material from India and Madagascar, although much of this material is either very fragmentary (a single tooth from Madagascar), or has not been described (material from India). Stay tuned to find out more about how these rare ankylosaurs fit into the big picture of ankylosaur evolution!
Next up: a grab bag of everybody else!
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Know Your Ankylosaurs: North American Odds and Ends Edition
I've covered many of the North American ankylosaurs in my previous papers and blog posts. In 2013, I argued that what we thought was Euoplocephalus was more likely 4 taxa– Anodontosaurus, Dyoplosaurus, Scolosaurus, and Euoplocephalus proper. Then in 2014 we described a newankylosaurid, Ziapelta, from New Mexico. There are a few other taxa that had previously been proposed to be ankylosaurids, so let's take a look at them here.
Aletopelta, Stegopelta and Glyptodontopelta
Aletopelta is one of the more tantalizingly enigmatic ankylosaurs from North America. It's from a weird place – California – which may have been much further south 75 million years ago compared to its current position. It was also found in marine sediments, and the decaying carcass had formed a little reef, with oysters encrusting the ribs. The only known specimen of Aletopelta is relatively complete, all things considered, with the osteoderms in situ over part of the pelvis, the legs partially articulated, and with various odds and ends like osteoderms and vertebrae. Unfortunately, the ends of the bones are often chewed apart, and some of the material is a bit hard to interpret.
Here's the articulated pelvis and hindlimbs, and some other armour pieces, on display at the San Diego Museum of Natural History.
Regardless, Aletopelta is a very interesting ankylosaur. It has an unusual osteoderm morphology over the pelvis, with small hexagonal osteoderms closely appressed to each other. Ankylosaur pelvic armour seems to come in two major flavours: fused rosettes, like we saw in Dongyangopelta and Taohelong (and perhaps most famously in Polacanthus), and interlocking hexagons, like in Stegopelta, Glyptodontopelta, and Aletopelta. Tracy Ford suggested that ankylosaurs with these hexagonal pelvic shields might represent a clade (dubbed Stegopeltinae) of ankylosaurids. Glyptodontopelta has since typically been interpreted as a nodosaurid, as has Stegopelta, but the most recent interpretation of Aletopelta was that it was an ankylosaurid. In the revised phylogeny in my new paper, we found Stegopelta and Glyptodontopelta as nodosaurids, but Aletopelta as a very basal ankylosaurid. However, although Ford and Kirkland reconstructed Aletopelta with the typical ankylosaurid tail club, I don't think that it possessed one: the preserved distal caudal vertebrae don't show any of the lengthening or other modifications that are characteristic of ankylosaurid handle vertebrae.
An updated restoration of the known elements in Aletopelta - the main differences between this and Ford and Kirkland's reconstruction are the absence of a tail club, and uncertainty over what the head should look like.
Cedarpelta
Cedarpelta is an important taxon for understanding the biogeography and evolution of ankylosaurids, and I wish we had more specimens! I don't have many new comments to add about this taxon, since Ken Carpenter published a great description of the disarticulated skull back in 2001. Cedarpelta has been interpreted as a shamosaurine ankylosaur, as a relative of taxa like Gobisaurus and Shamosaurus (which I'll talk about in the next post) from Asia, and thus may point towards a mid Cretaceous faunal interchange between these two continents. In our revised phylogenetic analysis, we didn't find Cedarpelta as the sister taxon to either Gobisaurus or Shamosaurus, but it does come out as a basal ankylosaurid in their general neighbourhood, and I honestly wouldn't be surprised if future analyses or new taxa show support for it as a shamosaurine ankylosaur after all.
Nodocephalosaurus
Nodocephalosaurus! What a fun ankylosaur. It's really quite unlike the other ankylosaurids from North America, which typically have flat, hexagonal cranial ornamentation. Instead, Nodocephalosaurus has bulbous, conical cranial ornamentation. Bulbous cranial ornamentation is typical of Campanian-Maastrichtian Mongolian ankylosaurs like Saichania and Tarchia, but in those taxa the ornamentation is pyramidal rather than conical. The front end of the snout in Nodocephalosaurus is also unusual, because there's no obvious narial opening and instead the ornamentation has a stepped appearance. Hopefully better specimens with more complete snouts will resolve this weird morphology. I've also reinterpreted the position of the quadratojugal horn compared to Sullivan's original figures – the horn should be rotated forward so that the bottom margin of the orbit is complete.
Nodocephalosaurus holotype skull in dorsal and left lateral views.
Tatankacephalus
I don't have much to say about Tatankacephalus because I didn't look at the original material myself, but the previous phylogenetic analysis by Thompson et al. recovered it as a nodosaurid rather than an ankylosaurid as originally suggested by Parsons and Parsons, and we found the same result. Overall, Tatankacephalus is VERY similar to Sauropelta, so this is perhaps not surprising.
Up next: More odds and ends, but after I return from Utah!
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Know Your Ankylosaurs: Mongolian Odds and Ends Edition
I'm back in civilization, so let's get back to ankylosaurs! Ready Set Go!
Gobisaurus, Zhongyuansaurus, and Shamosaurus
Shamosaurus is a really interesting ankylosaurid from the Zuunbayan Formation of Mongolia. Unlike later ankylosaurids, it still has a relatively long snout like you see in basal ankylosaurs and nodosaurids, and it lacks the distinctive tile-like skull ornamentation of ankylosaurs like Euoplocephalus or Saichania, instead just having a granular, pebbly texture on the skull surface. Gobisaurus, from the Ulansuhai Formation of China, is nearly identical in appearance, and only a few features distinguish these two taxa, namely the length of the tooth row relative to skull length and the orientation of the pterygoids. (Indeed, I think you could make an argument for subsuming Gobisaurus into Shamosaurus as Shamosaurus domoculus, but I'm generally reluctant to start making new combinations given that generic separation is pretty arbitrary anyway.)
Shamosaurus and its too-cool-for-school cervical half rings, on display in Moscow.
Gobisaurus and Shamosaurus are sister taxa; the name Shamosaurinae was proposed at one point and there's no reason to discard it at present even though it only contains two taxa. Shamosaurinae is the sister taxon to Ankylosaurinae. I also identified one new character that links Gobisaurus and Shamosaurus together which isn't present in other ankylosaurids: both taxa have a distinctive groove on each premaxilla, the purpose of which is unknown but there you go. There have been some suggestions that Cedarpelta (from North America) is also a shamosaurine ankylosaurid, and while I find the overall morphology of Cedarpelta to be pretty compelling for placing it in a clade with Gobisaurus and Shamosaurus, I didn't recover it with those taxa in my analysis (it came out more basally-positioned). However, I wouldn't be surprised if Cedarpelta winds up in Shamosaurinae at some point in the future as we find more specimens of both it and Gobisaurus and Shamosaurus.
Zhongyuansaurus was originally described as a nodosaurid ankylosaur partly because of its long snout, but it's indistinguishable from Gobisaurus (except for being smashed and flattened). The holotype is also a subadult (or at least not fully skeletally mature), since some of the cranial sutures are still visible towards the back of the skull. There are some interesting things going on with the postcrania of Zhongyuansaurus, but that's a story for a few weeks from now so STAY TUNED NO SPOILERS IF YOU'VE READ MY THESIS.
Tsagantegia
Of all of the more obscure ankylosaurs I looked at during my PhD, Tsagantegia might be my favourite for being the most surprising in person compared to what I had read about it. Tumanova included a line drawing of the specimen in her original description, which has been oft reproduced, but interestingly it doesn't really do justice to the original specimen (despite being a pretty nice drawing). The line drawing shows a long-snouted ankylosaur with amorphous cranial ornamentation, not dissimilar to Shamosaurus, but with a wider premaxillary beak more typical of later ankylosaurs. In person, however, the skull has distinct cranial caputegulae like we see in Euoplocephalus and Ankylosaurus! It's a pretty cool ankylosaur and I think it's probably really important to understanding the dispersal of ankylosaurs from Asia into North America and the diversification of ankylosaurids in the Campanian-Maastrichtian of Asia, but it's really hard to pin down the age of the Bayan Shiree Formation, and we don't have any postcrania for this taxon. I'm sure I'll be revisiting this guy in the future.
Heck yeah Tsagantegia!
Here it is again but in a more different view!
Talarurus
Way back when I originally started this blog in 2010, I had travelled to Korea to spend some time working in the Hwaseong paleo lab preparing Talarurus material and generally studying the ankylosaur material they had collected from the Gobi. Talarurus, like Tsagantegia, is also from the Bayan Shiree Formation but is clearly distinct. The holotype skull has very subtle cranial ornamentation that takes the form of small cones, rather than flat hexagonal tiles like Euoplocephalus, or bulbous pyramids like Saichania. Weirdly, this configuration is also present in the North American taxon Nodocephalosaurus – either this ornamentation style has convergently evolved, or, as I recovered in my analysis, these two taxa are closely related despite being fairly widely separated geographically and temporally. This is another ankylosaur that I'm sure we'll talk about again.
Talarurus butt in Moscow. The skeleton on display is a composite of several individuals from the same locality, and the skull is totally sculpted and a bit out of date.
Here's the holotype skull, with its weird, weird ornamentation.
Saichania
I've talked about Saichania fairly extensively here last year, but there were a few new things added in this most recent paper: Tianzhenosaurus and Shanxia (both from China) are, most likely, junior synonyms of Saichania, making this the most geographically widespread of the Asian ankylosaurids. Tianzhenosaurus has a nearly identical cranial ornamentation pattern when compared to Saichania, and I couldn't identify any differences that were outside of the usual ornamentation pattern variation we see in something like Euoplocephalus. Shanxia is known from the same formation but from a less well preserved skull, but the morphology of the squamosal horn is consistent with that of both Tianzhenosaurus and Saichania and therefore it probably represents the same taxon.
Next up: what's the big picture here, anyway?
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Know Your Ankylosaurs: Everybody's in this Together Edition
So with all of those posts about ankylosaur taxonomy over the last few weeks, what have we learned about the evolution of this group? Over the course of my PhD research, I was able to identify a bunch of new characters that seemed useful for understanding ankylosaur phylogenetic relationships, including characters related to the cranial ornamentation, pelvis, and osteoderms. Although ornamentation and osteoderms can be tricky, they can still yield useful information if you're careful about how you construct the characters.
Here's a sampling of some of the new characters from the supplementary file that goes along with the paper. Long live rainbow ankylosaur skulls.
With all the new information, here's what the results of the analyses gave us (click to embiggen):
From this, we can take away some interesting points:
1.Gondwanan ankylosaurs are probably not ankylosaurids, but they also don't form a single evolutionary group. Whatever "Minmi" is, it's a very basal kind of ankylosaur, possibly outside the split between Ankylosauridae and Nodosauridae. It's a little bit harder to say what's going on with "Antarctopelta" (previously considered an ankylosaurid), and the Argentinian ankylosaur: both came out as relatively derived nodosaurids, but my dataset isn't designed to test the interrelationships of nodosaurids. I wouldn't be surprised if future analyses incorporating more nodosaurids and more nodosaurid-based characters found that these two species were closely related. It would also be interesting to know which lineage of nodosaurids (probably a lineage from North America) dispersed into South America in the Late Cretaceous in order to give us these two ankylosaurs.
2.There are nodosaurids in the early-mid Cretaceous of Asia, but not necessarily the ones that have been proposed previously. Zhongyuansaurus, for example, was first described as a nodosaurid but is instead a junior synonym of the shamosaurine ankylosaurid Gobisaurus. However, a couple of taxa, like Taohelong, Sauroplites, and Dongyangopelta, are recovered as basal nodosaurids. At present, there doesn't seem to be much overlap between Asian nodosaurids and ankylosaurids, which is interesting! Why didn't nodosaurids hang on in Asia once ankylosaurids evolved, when the two groups seem to have coexisted pretty happily in North America later on?
3.The ankylosaurids from the Late Cretaceous of North America represent a dispersal of Asian ankylosaurines sometime during the early-mid Late Cretaceous. The earliest ankylosaurine is probably Crichtonpelta, from China, and North American ankylosaurines are a deeply nested clade within Ankylosaurinae. We propose the new name Ankylosaurini for the North American ankylosaurines (plus Talarurus, for now).
Here, have some frowny-faced rainbow ankylosaurs. Ankylosaurs are very serious dinosaurs.
4.Where do ankylosaurids first evolve? Unfortunately, that question isn't easy to answer right now: down at the base of Ankylosauridae, there's a mix of taxa from North America and Asia. The position of Gastonia as an ankylosaurid tips the scales slightly in favour of a North American origin for the clade, but some analyses recover this taxon as a nodosaurid, so I think we should be a little cautious about this result. One step up the tree, we've got a polytomy of Aletopelta and Cedarpelta (both from North America) and Liaoningosaurus and Chuanqilong (both from China). Does Ankylosauridae originate in North America with something like Cedarpelta, with a subsequent migration and diversification into Asia? Or does this group originate in Asia with something like Liaoningosaurus and Chuanqilong, and Cedarpelta represents an immigration into North America?
5.And finally, what's going on with ankylosaurids in the mid-Cretaceous of North America? Why don't we find any ankylosaurids between Cedarpelta and the later ankylosaurins? Did 'endemic' North American ankylosaurids go extinct during that time? And why does Aletopelta have such a weird basal phylogenetic position despite being from the Campanian? I don't really have answers for some of these questions, although if you come to the Society of Vertebrate Paleontology meeting in Dallas this October I'm going to try addressing some of them. For now, Aletopelta remains the biggest ankylosaurid enigma to me – it really shares very few things in common with the other Campanian ankylosaurids and I doubt it is an ankylosaurin from the Asian immigration into North America – could it represent a distinctive lineage of North American ankylosaurids stemming from things like Gastonia or Cedarpelta, for which we just don't have other representatives at the moment? Or, is it a nodosaurid masquerading as an ankylosaurid because I haven't sampled the right taxa or characters?
Darn you Aletopelta, why must you vex me so?
As usual, I wind up with more questions than answers every time I try to figure something out.
That wraps up the summaries for this paper, but stay tuned for some more cool research coming out in the next few weeks, and some summer fieldwork recaps!
Arbour VM, Currie PJ. In press. Systematics, phylogeny and palaeobiogeography of the ankylosaurid dinosaurs. Journal of Systematic Palaeontology.
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How the ankylosaur got its tail club
Ankylosaur tail clubs are odd structures, odder than they are usually given credit for. They represent substantial modifications to two different skeletal systems – the endoskeleton, in the form of the caudal vertebrae, and the dermal skeleton, in the form of the caudal osteoderms. The centra of the caudal vertebrae lengthen but stay robust, and the neural arches undergo huge changes, such that the prezygapophyses, postzygapophyses, and neural spine become a robust, V-shaped structure on the top of the centrum, and which creates a tightly interlocking vertebral series with almost no flexibility. We call this the handle of the tail club. The osteoderms at the tip of the tail smush together and two of them become huge: although the tail club knob is small in some species, there are colossal knobs exceeding 60 cm in width. The ankylosaur tail club represents one of the most extreme modifications to the tail in terrestrial tetrapods.
Look at that thing. That is a weird thing.
(This is UALVP 47273, a really nice club that I studied for my MSc work on tail club biomechanics.)
One of the questions I became interested in during my MSc research on ankylosaur tail club biomechanics was how the tail club evolved in the first place. Most ankylosaurs with tail clubs are known from a relatively narrow slice of time right at the end of the Cretaceous, but when and where did the tail club first evolve? Did the stiffening of the tail occur before the enlargement of the tail osteoderms, or vice versa? Or did both changes happen at about the same time? This was a fun question to address during my PhD research, once I had a fairly well resolved phylogeny of ankylosaurids, and once I had looked at tons of ankylosaurid fossils.
So, how did the ankylosaur get its tail club? Well, based on what we see in the fossil record, it looks like the changes to the vertebrae predate the changes to the osteoderms – in other words, the handle comes first and the knob comes later. There is at least one ankylosaur out there that seems to have a tail club handle but not a knob: Gobisaurus!
Hello Gobisaurus! Many many thanks to my friend and colleague Sydney Mohr for preparing this awesome illustration of Gobisaurus for me.
Gobisaurus, a shamosaurine ankylosaurid, has a really nice complete tail club handle that is indistinguishable from other ankylosaurid tail club handles, but does not have a knob. And it's not just because the knob is broken off – it seems as though the last vertebrae in the tail are preserved, because they look very similar to the terminal vertebrae in a CT scan of a tail club from the University of Alberta collections. It's likely that Gobisaurus had osteoderms along the sides of the tail like we see in most other ankylosaurs, but it doesn't appear that there were osteoderms tightly enveloping the tip of the tail.
An even earlier ankylosaur seems to show some changes towards acquiring a tail club handle, as well. Liaoningosaurus, a basal ankylosaurid known only from a very small juvenile, has distal caudal vertebrae where the prezyapophyses extend about 50% the length of the adjacent vertebra. This is what we see in ankylosaurid tail clubs, but not in more basal taxa like Mymoorapelta where the prezygapophyses are much shorter. Liaoningosaurus is missing the tip of the tail and also lacks osteoderms on most of its body because it's a juvenile, so it's harder to say whether or not it had a tail club knob based just on the fossil alone.
I also did a cool and relatively simple thing with my phylogenetic tree to see if I could better understand the likelihood that some ankylosaurs without preserved tail material had a tail club handle or full tail club with a knob. Unsurprisingly, all shamosaurine and ankylosaurine ankylosaurids probably had a tail club handle. Liaoningosaurus is part of a basal polytomy of ankylosaurids, and it was a bit more equivocal whether or not any of these taxa was likely to have a tail club handle or not, partly because another basal ankylosaurid in this region of the tree, Chuanqilong, does not have modified distal caudal vertebrae.
All ankylosaurine ankylosaurids more derived than Pinacosaurus (so including things like Tsagantegia, Saichania, Euoplocephalus, etc.) almost certainly had a tail club knob, and shamosaurine ankylosaurids probably did not. Crichtonpelta, the most basal ankylosaurine, may or may not have had a tail club – we'll need more data to know for sure. There is amounted skeleton of Crichtonpelta at the Sihetun visitor center in Liaoning, and it is shown with a tail club, but it isn't clear whether or not this is sculpted or original material belonging to this specimen, and a full description of this material is necessary.
Gobisaurus and Liaoningosaurus both lived much earlier than the more familiar tail-clubbed ankylosaurs: Gobisaurus is no younger than 92 million years old, and Liaoningosaurus is about 122 million years old. The earliest ankylosaurid with a tail club in the fossil record is Pinacosaurus(from the Campanian), although there is a caveat to this: Talarurus, which is a bit older than Pinacosaurus, should have a full tail club based on its position in the phylogenetic tree, and while a tail club handle is known for this taxon, we haven't found a tail club knob for Talarurus. Talarurus is in kind of a weird spot phylogenetically, since it's from Mongolia but comes out as closely related to North American ankylosaurines, so I think it's worth keeping an eye on this taxon in the future – perhaps Talarurus is another taxon with only a handle and not a knob, which would fit a bit better with its chronologic position if not its phylogenetic position.
Regardless, the changes to the vertebrae of ankylosaurs, starting with Liaoningosaurus at least 122 million years ago and continuing on towards Gobisaurus about 92 million years ago, seem to have occurred long before ankylosaurs evolved a huge osteodermal knob at the end of the tail. Was a stiff tail as good a weapon as a full tail club with a knob? What drove the evolution of the knob so long after the evolution of a stiff handle? And why did ankylosaurs even evolve a tail club at all? Now that I've had fun investigating how ankylosaurs might have used their tails, and how the tail club evolved, the next question feels like it should be 'why'....so stay tuned for more tail club fun over the next year or so as I make an attempt at that question!
Read it for yourself! Arbour VM, Currie PJ. In press. Ankylosaurid dinosaur tail clubs evolved through stepwise acquisition of key features. Journal of Anatomy.
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Snapshots from the Field Museum
Last week I got a chance to visit the Field Museum in Chicago for the first time! It's a great big museum with lots of cool stuff, so I figured I'd share a few impressions from my lunchtime jaunts through the exhibits. Let's get started with all the fossil exhibits outside of the main fossil hall (there are several, but some of them are kind of hidden away!).
SUE
Sue the Tyrannosaurusis most definitely not hidden away, and occupies a place of pride in the museum's main entrance hall. Sue is undeniably a great fossil, although I (and I suspect probably some other palaeontologists as well) have mixed feelings about this fossil: it's incredibly well preserved, but the intense backstory to Sue's acquisition is filled with several unpleasant twists and turns. I'm glad Sue found a home in a museum, but I wish it hadn't been placed up for auction - Sue's auctioning may not have directly led to the trend of putting dinosaurs up for auction for millions of dollars, but I feel like it set a bad precedent all the same.
One thing that's particularly enjoyable about this specific Tyrannosaurus skeleton are the abundant pathologies to be found. Sue has a busted/infected shin, holes in its jaw, and rough bumpy spots on its vertebrae. These vertebrae near the end of the tail have a big mass of crinkly bone around them. It's obvious Sue got up to some trouble during its life, and it's interesting to speculate on the causes of the various oddities in the skeleton (and indeed, others have!).
Extinct Madagascar
Sadly, this exhibit is tucked so far out of the way that basically nobody had wandered back there besides me (you need to go through the conservation gallery to reach it). It's also a little bit specimen-sparse, a trend I've noticed recently in many museums and which I find somewhat concerning. However, I feel like it makes up for the lack of 3D objects in its cool and unusual subject matter - the extinct fauna of Madagascar. The main point to the gallery was showcasing the social media response to new images of Madagascar's prehistory, and the scientific process that went into those images. It was an interesting way to approach the topic, but might have been more compelling with video, audio, or more fossils.
It was pretty cool to see an Aepyornis (elephant bird) egg and life-size silhouette. They really were terrifyingly large and strange birds.
A highlight for me was this Palaeopropithecus skeleton - a lemur that lived and looked like a sloth.
Tracking the Reptiles of Pangea
Tucked away in the African mammals area was a room devoted to palaeontological fieldwork in Tanzania, featuring the newly described silesaurid Asilisaurus! This isn't a skeleton you're going to see in most museums - I only wish more people had been stepping into this little exhibit room to check it out.
A nice touch was showing the original fossil material in its cabinet-ready storage foam. Those are some nice fossils.
And one last fossil....
Seriously, how were these machines not in constant use? They're in the hallway leading towards the bottom-floor cafeteria, and you can get yourself a freshly-made retro Triceratops, Brontosaurus, Tyrannosaurus, or Stegosaurus. I made a Brontosaurus and consider it $2 extremely well spent, especially since it meant I got rid of a bunch of dimes and nickels I didn't know what to do with:
Next time: Evolving Planet!
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Hylaeosaurus is an extinct genus of nodosaurid dinosaur which lived in southern England during the Early Cretaceous. On July 20, 1832, Hideon Mantell wrote Benjamin Silliman about reptilian bones recovered from a gunpowder explosion in Tilgate Forest, with a local fossil dealer assembling ~50...
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Hylaeosaurus is an extinct genus of nodosaurid dinosaur which lived in southern England during the Early Cretaceous.
History[]
On July 20, 1832, Hideon Mantell wrote Benjamin Silliman about reptilian bones recovered from a gunpowder explosion in Tilgate Forest, with a local fossil dealer assembling ~50 fragments, describing them as a "great consarn of bites and boanes". Mentell purchased these and assembled them into a full skeleton, doubting their value. It was partially-articulate. Mantell had grown satisfied with this, since previous Megalosaurus and Iguanodon had consisted of single fragments, becoming the most complete non-avian dinosaur skeleton at the time, strongly wanting to name it a species of the latter. However, a visit by William Clift and John Edward Gray had changed his mind; Clift had found the spikes to be body armour. Mantell (1832) assigned it a new genus, Hylaeosaurus, meaning "lizard of the wood". However, he later claimed it meant "forest lizard" and "Wealden lizard". Mantell (November 30) sent it to the Geological Society of London, then travelling to London on December 5, then being informed his publication was 1/3 too long. Instead of performing a rewrite, Charles Lyell suggested he publish a book with a single chapter dedicated to Hylaeosaurus. In the next 3 weeks, he composed this, with Henry Da le Beche telling him he needed a specific epithet. On the 19th of December, he chose armatus, stating "there appears every reason to conclude that either its back was armed with a formidable row of spines, constituting a dermal fringe, or that its tail possessed the same appendage". In May 1833, The Geology of the South-East of England was published and Hylaeosaurus had become a valid genus and species, also publishing a lithograph of his find in the same book and another in The Wonders of Geology (1840). It is the most obscure of the first three dinosaurs used to define dinosauria. Additionally, it joins the suite of extinct animals in the Crystal Palace sculptures and was never used as a wastebasket taxon, unlike the others. Owen (1840) thought the spikes were asymmetrical, rejected that they formed a row on the back and assumed they were gastralia.
The Natural History Museum acquired the original specimen in 1838, assigned BMNH R3775 and then NHMUK 3775. It is from the Valanginian-aged Tunbridge Wells Sand Formation. It is the front of the skeleton lacking most of the head and forelimbs, measuring 135x75 centimeters. It consists of the posterior skull, possible mandibular fragments, 10 vertebrae, scapulae, coracoids and some spikes and armour; it is viewed from below. No further preparation had taken place for a while beyond Mantell's assembly and chiseling, with the museum attempting to free the remains via chemical and mechanical methods in the 21st century. However, this is difficult because the acids tend to dissolve the glue and gypsum Mantell had used to repair it, causing the blocks to disarticulate. Mark Graham (since 2003) has given limited details of this, but it was published and redescribed in 2020. Several more specimens from mainland Britain have been referred to H. armatus. However, Paul Barrett and Susannah Maidment (2011) concluded only the holotype is referable, since Polacanthus are recovered from the same layers. Specimens from France may be of Polacanthus, with possibly remains from Germany (DLM 537, a spike; and GPMM A3D.3, a lower humeral end) were assigned H. sp..
Description[]
Gideon Mantell, based on lizards, estimated Hylaeosaurus was 7.6 meters (25 feet) long, with recent estimations finding up to 6 meters (20 feet). Gregory S. Paul (2010) estimates a length of 5 meters (16 feet), weighting 2 tonnes (2 long tons; 2.2 short tons). Darren Naish et al. (2001) find a length of 3-4 meters (9.8-13.1 feet). Hylaeosaurus has a mainly unknown build. Maidment finds two autapomorphies; the scapula does not fuse with the coracoid and 3 long spines are at the shoulder. Though, these do not distinguish it much. Mantell and Owen attributed the (lack of) fusion to ontogenetic causes, and find the total spine count can not be seen. It is often depicted as a typical nodosaur. Kenneth Carpenter (2001) described cranial and mandibular remains, but the damaged and warped skull provides little data. The quadrate laterally bows; the quadratojugal bears a high attachment on the quadrate shaft; a triangular postorbital horn is present. In 2020, however, the presumed quadrate was fount to have been a jugal. The 2020 study found new distinguishing traits. A 120° is between the acromion and proximal plate. The acromial process is shaped like a shelf at 1/3 from the top edge that obliquely projects below and sideways. The top edge of the proximal plate curves sideways. The cervical centra have a horizontal ridge at the sides. Not included in this list are the exceptionally-concave undersides of the side processes. The shoulder spines are curved at the posterior, elongate, flattened, narrow and pointed. A shallow trough resides on the underside. The front spine is 42.5 centimeters long, where the spines shrink in height and enlarge in width. A fourth skull of about the same build, being more forwards-pointing, sits just behind the skull. Sven Sachs and Jahn Hornung (2013) suggest a configuration where 5 lateral nick spines are present, with the German spine specimen assigned the 3rd position.
Classification[]
Hylaeosaurus is the first known ankylosaur, but its affinities are poorly known to this day. Coombs (1978) assigned it Nodosauridae, now seen as an unusual classification, as recent analyses place it a basal nodosaur and sometimes a polacanthine. In the 1990s, polacanthines were recovered as basal ankylosaurids since they believed they had small tail clubs, which is now recognized as incorrect. A 2012 study finds it to be a non-polacanthine basal nodosaurid:
Nodosauridae
Antarctopelta
Mymoorapelta
Hylaeosaurus
Anoplosaurus
Tatankacephalus
Horshamosaurus
Polacanthinae
Gargoyleosaurus
Hoplitosaurus
GastoniaGastonia
Peloroplites
Polacanthus
Struthiosaurus
Zhejiangosaurus
Hungarosaurus
Animantarx
Niobrarasaurus
Nodosaurus
Pawpawsaurus
Sauropelta
Silvisaurus
Stegopelta
Texasetes
Edmontonia
Panoplosaurus
References[]
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2009-04-25T19:33:56+00:00
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https://en.wikipedia.org/wiki/List_of_Asian_dinosaurs
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Name Year Formation Location Notes Images Abdarainurus 2020 Alagteeg Formation (Late Cretaceous, Santonian to Campanian) Mongolia Inconsistent in phylogenetic placement; could represent an unknown lineage of macronarians[1] Abrosaurus 1989 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China Had unusually large fenestrae Achillobator 1999 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia Its robust build suggests it was not a cursorial animal[2] Adasaurus 1983 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Its sickle claw was markedly reduced compared to other dromaeosaurids Aepyornithomimus 2017 Djadochta Formation (Late Cretaceous, Campanian) Mongolia The first ornithomimosaur named from a dry desert environment Agilisaurus 1990 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China The holotype specimen was discovered during the construction of the museum where it is now housed Albalophosaurus 2009 Kuwajima Formation (Early Cretaceous, Valanginian to Hauterivian) Japan Only known from fragments of a skull Albinykus 2011 Javkhlant Formation (Late Cretaceous, Santonian) Mongolia Preserved in a sitting position not unlike that of modern birds Alectrosaurus 1933 Iren Dabasu Formation (Late Cretaceous, Cenomanian to Santonian) China Had long legs which may be an adaptation to pursuit predation[3] Alioramus 1976 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Possessed an elongated snout with a row of short crests Almas 2017 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Preserved alongside eggshells which may have come from a troodontid[4] Altirhinus 1998 Khuren Dukh Formation (Early Cretaceous, Barremian to Albian) Mongolia Had a distinctive, elevated nasal bone which supported a large nasal cavity Alxasaurus 1993 Bayin-Gobi Formation (Early Cretaceous, Albian) China Most of the skeleton is known, which allowed researchers to connect therizinosaurs to other theropods Ambopteryx 2019 Unnamed formation (Late Jurassic, Oxfordian) China Preserves stomach contents containing gastroliths and fragments of bone, suggesting an omnivorous diet Amtocephale 2011 Bayan Shireh Formation (Late Cretaceous, Turonian to Santonian) Mongolia One of the oldest known pachycephalosaurs Amurosaurus 1991 Udurchukan Formation (Late Cretaceous, Maastrichtian) Russia One specimen may have come from an individual with a limp[5] Analong 2020 Chuanjie Formation (Middle Jurassic, Bajocian) China Originally described as a specimen of Chuanjiesaurus but assigned a new genus due to several morphological differences Anchiornis 2009 Tiaojishan Formation (Late Jurassic, Oxfordian) China Analysis of fossilized melanosomes suggest a mostly gray or black body, white and black patterns on its wings, and a red head crest[6] Anhuilong 2020 Hongqin Formation (Middle Jurassic, Aalenian to Callovian) China Closely related to Huangshanlong and Omeisaurus, all forming an exclusive clade of mamenchisaurids Anomalipes 2018 Wangshi Group (Late Cretaceous, Maastrichtian) China May have been closely related to Gigantoraptor despite its significantly smaller size[7] Anserimimus 1988 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Had powerful forelimbs with uniquely straight, flattened claws Aorun 2013 Shishugou Formation, (Late Jurassic, Oxfordian) China Potentially a basal member of the alvarezsaurian lineage[8] Aralosaurus 1968 Bostobe Formation, (Late Cretaceous, Santonian to Campanian) Kazakhstan Its crest has been interpreted as being arch-shaped as in kritosaurin hadrosaurs, but this cannot be confirmed Archaeoceratops 1997 Xinminbao Group (Early Cretaceous, Barremian) China Had no horns and only the beginnings of a frill Archaeornithoides 1992 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Known from only a partial skull with scratches that may have been created by a small mammal[9] Archaeornithomimus 1972 Bissekty Formation?, Iren Dabasu Formation (Late Cretaceous, Cenomanian to Turonian) China
Uzbekistan? Unlike other ornithomimosaurs, its feet were not arctometatarsalian Arkharavia 2010 Udurchukan Formation (Late Cretaceous, Maastrichtian) Russia Described from a series of vertebrae, several of which were found to not belong to this taxon[10] Arstanosaurus 1982 Bostobe Formation (Late Cretaceous, Santonian to Campanian) Kazakhstan Poorly known Asiaceratops 1989 Khodzhakul Formation, Xinminbao Group? (Late Cretaceous, Cenomanian) China?
Uzbekistan Potentially a leptoceratopsid[11] Asiatosaurus 1924 Öösh Formation, Xinlong Formation (Early Cretaceous, Barremian to Albian) China
Mongolia Two species have been named but both are only known from extremely scant remains Auroraceratops 2005 Xinminbao Group (Early Cretaceous, Aptian) China Known from more than eighty specimens, including complete skeletons Aurornis 2013 Tiaojishan Formation (Late Jurassic, Oxfordian) China If an avialan as originally described it would be one of the oldest members of the group Avimimus 1981 Barun Goyot Formation, Nemegt Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia Bonebed remains indicate a gregarious lifestyle; it may have formed age-segregated herds for lekking or flocking purposes[12] Bactrosaurus 1933 Iren Dabasu Formation, Majiacun Formation? (Late Cretaceous, Cenomanian to Santonian?) China Remains of at least six individuals are known, making up much of the skeleton Bagaceratops 1975 Barun Goyot Formation, Bayan Mandahu Formation, Djadochta Formation? (Late Cretaceous, Campanian to Maastrichtian) China
Mongolia May have been a direct descendant of Protoceratops which it physically resembles[13] Bagaraatan 1996 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Combines traits of several theropod groups, possibly due to being chimaeric[14] Bainoceratops 2003 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Its supposedly diagnostic features may fall within Protoceratops variation[15] Banji 2010 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Vertical striations adorned the sides of its crest Bannykus 2018 Bayin-Gobi Formation (Early Cretaceous, Barremian to Aptian) China Exhibited a transitional hand morphology for an alvarezsaur, having three fingers of roughly equal length with the first being robust Baotianmansaurus 2009 Gaogou Formation (Late Cretaceous, Cenomanian to Turonian) China Large but known from only a few bones Barsboldia 1981 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Possessed elongated neural spines particularly above the hips Bashanosaurus 2022 Shaximiao Formation (Middle Jurassic, Bajocian) China Its skeleton combines traits of stegosaurs and more basal thyreophorans Bashunosaurus 2004 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China Although described as a macronarian, this has yet to be rigorously tested[16] Batyrosaurus 2012 Bostobe Formation (Late Cretaceous, Santonian to Campanian) Kazakhstan Remains originally identified as Arstanosaurus Bayannurosaurus 2018 Bayin-Gobi Formation (Early Cretaceous, Aptian) China Known from a well-preserved, almost complete skeleton Beg 2020 Ulaanoosh Formation (Early Cretaceous to Late Cretaceous, Albian to Cenomanian) Mongolia Its preserved skull has a rugose texture Beibeilong 2017 Gaogou Formation (Late Cretaceous, Cenomanian to Coniacian) China Similar to but more basal than Gigantoraptor.[17] Known from only a single embryo still in its egg Beipiaosaurus 1999 Yixian Formation (Early Cretaceous, Aptian) China Preserves evidence of downy feathers as well as a secondary coat of simpler "elongated broad filamentous feathers" or EBFFs[18] Beishanlong 2010 Xinminbao Group (Early Cretaceous, Aptian to Albian) China Lacked the elongated claws of more derived ornithomimosaurs Bellusaurus 1990 Shishugou Formation (Late Jurassic, Oxfordian) China Known from a bone bed with the remains of seventeen juvenile specimens Bienosaurus 2001 Lufeng Formation (Early Jurassic, Sinemurian) China Potentially synonymous with Tatisaurus[19] Bissektipelta 2004 Bissekty Formation (Late Cretaceous, Turonian to Coniacian) Uzbekistan Analysis of its braincase suggests poor hearing and eyesight but good olfaction and taste; it has been suggested to be a filter feeder[20] Bolong 2010 Yixian Formation (Early Cretaceous, Aptian) China Originally known from only a skull; an almost complete skeleton was described in 2013[21] Borealosaurus 2004 Sunjiawan Formation (Late Cretaceous, Cenomanian to Turonian) China Its caudal vertebrae were distinctively opisthocoelous Borogovia 1987 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Had a uniquely straight and flattened sickle claw, which may have had a weight-bearing function Breviceratops 1990 Barun Goyot Formation (Late Cretaceous, Campanian) Mongolia Only known from juvenile remains but can be distinguished from other protoceratopsids Brohisaurus 2003 Sembar Formation (Late Jurassic, Kimmeridgian) Pakistan Possibly an early titanosauriform Byronosaurus 2000 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Two juvenile skulls were found in an oviraptorid nest and claimed to be evidence for nest parasitism in this taxon, but both their identity and taphonomy have been questioned[4][22] Caenagnathasia 1994 Bissekty Formation (Late Cretaceous, Turonian to Coniacian) Uzbekistan One of the oldest and smallest known caenagnathoids Caihong 2018 Tiaojishan Formation (Late Jurassic, Oxfordian) China Possessed platelet-shaped melanosomes that produced iridesence as in modern trumpeters Caudipteryx 1998 Yixian Formation (Early Cretaceous, Barremian) China Two species are known. At least C. zoui did not have secondary feathers attached to the lower arm Ceratonykus 2009 Barun Goyot Formation (Late Cretaceous, Campanian) Mongolia Several osteological features were described as similar to ornithischians[23] Changchunsaurus 2005 Quantou Formation (Early Cretaceous to Late Cretaceous, Aptian to Cenomanian) China Had wavy enamel on its leaf-shaped teeth that made them more resistant to wear; this feature is also present in hadrosaurs[24] Changmiania 2020 Yixian Formation (Early Cretaceous, Barremian) China Preserved in a curled-up position as if sleeping in a potential burrow Changyuraptor 2014 Yixian Formation (Early Cretaceous, Barremian) China The largest microraptorian dromaeosaurid known. Had tail feathers almost a foot long[25] Chaoyangsaurus 1999 Tuchengzi Formation (Late Jurassic, Tithonian) China Known by a number of alternate spellings (e.g. Chaoyangosaurus, Chaoyoungosaurus) before its formal description Charonosaurus 2000 Yuliangze Formation (Late Cretaceous, Maastrichtian) China May have had a long, backwards-arcing crest similar to that of Parasaurolophus Chialingosaurus 1959 Shaximiao Formation (Late Jurassic, Oxfordian to Kimmeridgian) China Had both large plates and smaller spines, similar to Kentrosaurus Chiayusaurus 1953 Hasandong Formation, Xinminbao Group (Early Cretaceous, Barremian to Albian) China
South Korea Two species have been named, both from teeth. Those of C. lacustris are apparently indistinguishable to those of Euhelopus[26] or Mamenchisaurus[27] Chilantaisaurus 1964 Ulansuhai Formation (Late Cretaceous, Turonian) China Had a particularly hooked claw on its first finger Chingkankousaurus 1958 Wangshi Group (Late Cretaceous, Santonian to Campanian) China Known from only a scapula. Possibly a tyrannosauroid[28] Chinshakiangosaurus 1992 Fengjiahe Formation (Early Jurassic, Hettangian) China Had a U-shaped snout that may have supported fleshy cheeks, an adaptation to bulk feeding Choyrodon 2018 Khuren Dukh Formation (Early Cretaceous, Albian) Mongolia It had an enlarged nose similar to its contemporary, Altirhinus, but it is most likely a separate taxon[29] Chuandongocoelurus 1984 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China A tetanuran of uncertain relationships Chuanjiesaurus 2000 Chuanjie Formation (Middle Jurassic, Bathonian) China One of the more derived mamenchisaurids[30] Chuanqilong 2014 Jiufotang Formation (Early Cretaceous, Barremian to Aptian) China May have been the adult form of the coeval Liaoningosaurus[31] Chungkingosaurus 1983 Shaximiao Formation (Late Jurassic, Oxfordian) China May have possessed at least six thagomizer spikes; the rearmost pair was mounted horizontally, directed outwards and backwards Chuxiongosaurus 2010 Lufeng Formation (Early Jurassic, Hettangian to Pliensbachian) China Potentially a synonym of Jingshanosaurus[32] Citipati 2001 Djadochta Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia Had a distinctive triangular crest. A referred specimen known as the Zamyn Khondt oviraptorid possessed the familiar rectangular domed crest in most depictions of Oviraptor, but likely does not belong to that genus or Citipati[33] Conchoraptor 1986 Barun Goyot Formation, Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Named for a hypothesized diet of shellfish, but this cannot be confirmed Corythoraptor 2017 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Its crest was vertical and rectangular, not unlike that of a cassowary Crichtonpelta 2015 Sunjiawan Formation (Late Cretaceous, Cenomanian) China Originally named as a second species of Crichtonsaurus Crichtonsaurus 2002 Sunjiawan Formation (Late Cretaceous, Cenomanian to Turonian) China Sometimes reconstructed with semicircular osteoderms vaguely similar to the plates of stegosaurs Daanosaurus 2005 Shaximiao Formation (Late Jurassic, Oxfordian to Tithonian) China Phylogenetic position is uncertain as it is only known from the remains of a juvenile Daliansaurus 2017 Yixian Formation (Early Cretaceous, Barremian) China Had an enlarged claw on the fourth toe comparable in size to the sickle claw on its second Dashanpusaurus 2005 Shaximiao Formation (Middle Jurassic, Callovian) China One of the basalmost and earliest known macronarians[34] Datanglong 2014 Xinlong Formation (Early Cretaceous, Barremian to Albian) China Had a uniquely pneumatized ilium similar to megaraptorans Datonglong 2016 Huiquanpu Formation (Late Cretaceous, Cenomanian to Campanian) China Precise dating uncertain Datousaurus 1984 Shaximiao Formation (Middle Jurassic to Late Jurassic, Bathonian to Oxfordian) China One of the rarer sauropods of the Shaximiao, known from only two skeletons and a large, deep skull Daurlong 2022 Longjiang Formation (Early Cretaceous, Aptian) China Preserves remains of an intestinal tract Daxiatitan 2008 Hekou Group (Early Cretaceous, Barremian) China Large and very long-necked Deinocheirus 1970 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Had a suite of unique features, most notably a hump supported by elongated neural spines Dilong 2004 Yixian Formation (Early Cretaceous, Barremian) China Preserves evidence of a coating of simple feathers Dongbeititan 2007 Yixian Formation (Early Cretaceous, Barremian) China A theropod tooth has been found encrusted in one of its ribs[35] Dongyangopelta 2013 Chaochuan Formation (Early Cretaceous to Late Cretaceous, Albian to Cenomanian) China Coexisted with Zhejiangosaurus but could be distinguished based on subtle osteological features[36] Dongyangosaurus 2008 Jinhua Formation (Late Cretaceous, Turonian to Coniacian) China Its phylogenetic placement is uncertain Dzharaonyx 2022 Bissekty Formation (Late Cretaceous, Turonian) Uzbekistan One of the oldest known parvicursorines Dzharatitanis 2021 Bissekty Formation (Late Cretaceous, Turonian) Uzbekistan Originally described as a rebbachisaurid[37] but later reinterpreted as a titanosaur with possible lognkosaurian affinities[38] Elmisaurus 1981 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia One of the most complete caenagnathids known Embasaurus 1931 Neocomian Sands (Early Cretaceous, Berriasian) Kazakhstan Known from only two vertebrae Enigmosaurus 1983 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia Had a large, backwards-pointing pelvis Eomamenchisaurus 2008 Zhanghe Formation (Middle Jurassic to Late Jurassic, Aalenian to Oxfordian) China One of the oldest mamenchisaurids Eosinopteryx 2013 Tiaojishan Formation (Late Jurassic, Oxfordian) China Described as lacking advanced tail feathers and long "hind wings", unlike other paravians, but this may be an artifact of preservation[39] Epidexipteryx 2008 Haifanggou Formation (Middle Jurassic, Callovian) China Supported four long feathers from an abbreviated tail Equijubus 2003 Xinminbao Group (Early Cretaceous, Albian) China A grazer that preserves the oldest known evidence of grass-eating[40] Erketu 2006 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia May have had the longest neck of any dinosaur relative to its body Erliansaurus 2002 Iren Dabasu Formation (Late Cretaceous, Cenomanian) China Had long, curved claws on its fingers Erlikosaurus 1980 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia Preserves the most complete skull known from any therizinosaur Eshanosaurus 2001 Lufeng Formation (Early Jurassic, Hettangian) China Has been suggested to be the oldest known therizinosaur Euhelopus 1956 Meng-Yin Formation (Early Cretaceous, Berriasian to Valanginian) China Originally believed to have lived in a marshy environment Euronychodon 1991 Bissekty Formation (Late Cretaceous, Turonian) Uzbekistan Type species was found in Portugal. The Asian species may represent a form taxon of improperly developed teeth[41] Ferganasaurus 2003 Balabansai Formation (Middle Jurassic, Callovian) Kyrgyzstan Claimed to have two hand claws, but this is disputed[42] Ferganocephale 2005 Balabansai Formation (Middle Jurassic, Callovian) Kyrgyzstan Unusually, its teeth were not serrated Fujianvenator 2023 Nanyuan Formation (Late Jurassic, Tithonian) China Possessed proportionally long legs which may be an adaptation to wading Fukuiraptor 2000 Kitadani Formation, Sebayashi Formation? (Early Cretaceous, Barremian to Aptian) Japan Similarly to Megaraptor, it was originally reconstructed as a dromaeosaur with its hand claw on its foot Fukuisaurus 2003 Kitadani Formation (Early Cretaceous, Barremian) Japan The elements of its skull are so strongly fused that it was unable to chew[43] Fukuititan 2010 Kitadani Formation (Early Cretaceous, Barremian to Aptian) Japan The first sauropod named from Japan Fukuivenator 2016 Kitadani Formation (Early Cretaceous, Barremian to Aptian) Japan Possesses traits of various groups of coelurosaurs, though probably a therizinosaur.[44] May have been a herbivore or omnivore due to its heterodont dentition Fulengia 1977 Lufeng Formation (Early Jurassic, Hettangian to Toarcian) China May have been a juvenile Lufengosaurus Fushanosaurus 2019 Shishugou Formation (Late Jurassic, Oxfordian) China Known from a single femur of immense size Fusuisaurus 2006 Xinlong Formation (Early Cretaceous, Aptian to Albian) China A referred humerus may support an extremely large size for this taxon[45] Gallimimus 1972 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Had a relatively long beak with a rounded tip Gannansaurus 2013 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Its vertebrae were more similar to those of Euhelopus than to other sauropods Ganzhousaurus 2013 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Coexisted with at least seven other oviraptorosaurs, which may have niche-partitioned. It was likely primarily herbivorous[46] Garudimimus 1981 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia Was not as well-adapted to running as later ornithomimosaurs Gasosaurus 1985 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China Discovered as a byproduct of construction work Gigantoraptor 2007 Iren Dabasu Formation (Late Cretaceous, Cenomanian) China The largest known oviraptorosaur, comparable in size to Albertosaurus Gigantspinosaurus 1992 Shaximiao Formation (Late Jurassic, Oxfordian) China Possessed broad, greatly enlarged shoulder spines Gilmoreosaurus 1979 Bissekty Formation?, Iren Dabasu Formation, Khodzhakul Formation? (Late Cretaceous, Cenomanian) China
Uzbekistan? Several fossils preserve evidence of cancer-induced tumors[47] Gobihadros 2019 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia Known from multiple specimens representing different growth stages Gobiraptor 2019 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Possessed a deep jaw that may be an adaptation to crushing bivalves or seeds[48] Gobisaurus 2001 Ulansuhai Formation (Late Cretaceous, Turonian) China Had no tail club but already possessed the stiff tail of derived ankylosaurids[49] Gobititan 2003 Xinminbao Group (Early Cretaceous, Aptian) China Retained the fifth digit of the foot, a basal trait Gobivenator 2014 Djadochta Formation (Late Cretaceous, Campanian) Mongolia The most completely known Cretaceous troodontid Gongbusaurus 1983 Shaximiao Formation (Late Jurassic, Oxfordian) China Only known from a pair of teeth. May be an ankylosaurian[50] Gongpoquansaurus 2014 Xinminbao Group (Early Cretaceous, Albian) China Remains originally named as a species of Probactrosaurus Gongxianosaurus 1998 Ziliujing Formation (Early Jurassic, Toarcian) China The only sauropod with ossified distal tarsals, hinting at its basal position Goyocephale 1982 Unnamed formation (Late Cretaceous, Campanian) Mongolia Had a sloping head with a flat skull roof Graciliceratops 2000 Bayan Shireh Formation (Late Cretaceous, Santonian) Mongolia Possessed a short frill with large fenestrae Graciliraptor 2004 Yixian Formation (Early Cretaceous, Barremian) China A close relative of Microraptor with characteristically slender bones Guanlong 2006 Shishugou Formation (Late Jurassic, Oxfordian) China Two specimens have been discovered, one on top of the other Halszkaraptor 2017 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Originally interpreted as a semiaquatic fish hunter similar to a merganser[51] but this hypothesis has been criticized[52] Hamititan 2021 Shengjinkou Formation (Early Cretaceous, Aptian) China Known from seven caudal vertebrae and associated elements Haplocheirus 2010 Shishugou Formation (Late Jurassic, Oxfordian) China Possessed three long fingers with short claws. Originally described as a basal alvarezsauroid but similarities have been noted with other coelurosaurs[14][53] Harpymimus 1984 Khuren Dukh Formation?/Shinekhudag Formation? (Early Cretaceous, Albian) Mongolia Mostly toothless but retains a few teeth in the dentary Haya 2011 Javkhlant Formation (Late Cretaceous, Santonian to Campanian) Mongolia One specimen preserves a large mass of gastroliths Heishansaurus 1953 Minhe Formation (Late Cretaceous, Campanian to Maastrichtian) China May be a junior synonym of Pinacosaurus[54] Helioceratops 2009 Quantou Formation (Early Cretaceous to Late Cretaceous, Aptian to Cenomanian) China Had a distinctively short lower jaw Hexing 2012 Yixian Formation (Early Cretaceous, Valanginian to Barremian) China Three or four teeth are known, but they are not well-preserved Hexinlusaurus 2005 Shaximiao Formation (Middle Jurassic, Bajocian) China Originally named as a species of Yandusaurus Heyuannia 2002 Barun Goyot Formation, Dalangshan Formation (Late Cretaceous, Maastrichtian) China
Mongolia Fossilized pigments in referred eggshells suggest they were blue-green[55] Homalocephale 1974 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Has been suggested to be a juvenile Prenocephale on account of its flat head,[56] but this is no longer thought to be the case[57] Huabeisaurus 2000 Huiquanpu Formation (Late Cretaceous, Cenomanian to Maastrichtian) China May be closely related to Tangvayosaurus[58] Hualianceratops 2015 Shishugou Formation (Late Jurassic, Oxfordian) China Had a series of bumps around the edge of the beak Huanansaurus 2015 Nanxiong Formation (Late Cretaceous, Campanian to Maastrichtian) China Possessed a distinctive short trapezoidal crest Huanghetitan 2006 Haoling Formation, Hekou Group (Early Cretaceous, Aptian to Albian) China Had ribs 3 metres (9.8 ft) long, which supported one of the deepest body cavities of any dinosaur[59] Huangshanlong 2014 Hongqin Formation (Middle Jurassic to Late Jurassic, Aalenian to Oxfordian) China Known from some bones of the right forelimb Huaxiagnathus 2004 Yixian Formation (Early Cretaceous, Aptian) China One of the largest known compsognathids Huayangosaurus 1982 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China Possessed flank osteoderms and a small tail club in addition to plates and spikes Hudiesaurus 1997 Kalaza Formation (Late Jurassic, Tithonian) China Had a butterfly-shaped process on its vertebra Hulsanpes 1982 Barun Goyot Formation (Late Cretaceous, Campanian) Mongolia Closely related to Halszkaraptor but appears to be more cursorial[60] Ichthyovenator 2012 Grès supérieurs Formation (Early Cretaceous, Aptian) Laos One of its sacral vertebrae was greatly reduced, giving the illusion of a break in its sail or of two separate sails Incisivosaurus 2002 Yixian Formation (Early Cretaceous, Barremian) China Two specimens of different ontogenetic stages are known, both with differing types of feathers[61] Irisosaurus 2020 Fengjiahe Formation (Early Jurassic, Hettangian) China Closely related to Mussaurus[62] Isanosaurus 2000 Nam Phong Formation (Late Triassic, Norian to Rhaetian) Thailand May have actually come from the Late Jurassic[63] Ischioceratops 2015 Wangshi Group (Late Cretaceous, Campanian to Maastrichtian) China Noted for its peculiarly-shaped ischium Itemirus 1976 Bissekty Formation (Late Cretaceous, Turonian) Uzbekistan Originally known from a braincase but abundant new remains were described in 2014[64] Jaculinykus 2023 Barun Goyot Formation (Late Cretaceous, Maastrichtian) Mongolia Was didactyl, with a large first finger and a reduced second finger Jaxartosaurus 1937 Dabrazhin Formation (Late Cretaceous, Santonian) Kazakhstan Not known from many remains but they are enough to tell that it was a basal lambeosaurine[65] Jeholosaurus 2000 Yixian Formation (Early Cretaceous, Aptian) China One specimen is preserved in a curled position Jianchangosaurus 2013 Yixian Formation (Early Cretaceous, Barremian) China Several characters of its teeth and jaws are convergently similar to those of ornithischians[66] Jiangjunosaurus 2007 Shishugou Formation (Late Jurassic, Oxfordian) China Had two rows of circular or diamond-shaped plates Jiangshanosaurus 2001 Jinhua Formation (Late Cretaceous, Turonian to Coniacian) China A potential member of the Euhelopodidae[67] Jiangxisaurus 2013 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Overall similar to Heyuannia but with a thinner, frailer mandible Jiangxititan 2023 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Described as one of the few known lognkosaurs from mainland Asia Jianianhualong 2017 Yixian Formation (Early Cretaceous, Aptian) China Possessed a subtriangular tail frond made of asymmetrical feathers, although it was most likely flightless Jinbeisaurus 2019 Huiquanpu Formation (Late Cretaceous, Cenomanian to Maastrichtian) China A medium-sized tyrannosauroid Jinfengopteryx 2005 Huajiying Formation (Early Cretaceous, Barremian) China May have been capable of some sort of flight[68] Jingshanosaurus 1995 Lufeng Formation (Early Jurassic, Hettangian) China One of the latest-surviving non-sauropod sauropodomorphs Jintasaurus 2009 Xinminbao Group (Early Cretaceous, Albian) China Known from only the rear half of a skull, including a complete braincase Jinyunpelta 2018 Liangtoutang Formation (Early Cretaceous to Late Cretaceous, Albian to Cenomanian) China The oldest ankylosaurid known to have a tail club Jinzhousaurus 2001 Yixian Formation (Early Cretaceous, Aptian) China Its holotype is nearly complete, preserved whole on a single slab Jiutaisaurus 2006 Quantou Formation (Early Cretaceous to Late Cretaceous, Barremian to Cenomanian) China Named based on eighteen vertebrae Kaijiangosaurus 1984 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China Potentially synonymous with other medium-sized Shaximiao theropods Kamuysaurus 2019 Hakobuchi Formation (Late Cretaceous, Maastrichtian) Japan Informally referred to as "Mukawaryu" before its formal description Kansaignathus 2021 Ialovachsk Formation (Late Cretaceous, Santonian) Tajikistan The first non-avian dinosaur described from Tajikistan Kazaklambia 2013 Dabrazhin Formation (Late Cretaceous, Santonian) Kazakhstan Morphologically distinct from other Eurasian lambeosaurines[69] Kelmayisaurus 1973 Lianmuqin Formation (Early Cretaceous, Valanginian to Albian) China One popular book mentions a giant species belonging to this genus,[70] but this referral may be incorrect Kerberosaurus 2004 Tsagayan Formation (Late Cretaceous, Maastrichtian) Russia Potentially a close relative of Edmontosaurus[71] Khaan 2001 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Two morphotypes of chevrons are known, which may be a sexually dimorphic trait[72] Khulsanurus 2021 Barun Goyot Formation (Late Cretaceous, Campanian) Mongolia Contemporary with Parvicursor but can be distinguished by characters of its caudal vertebrae[73] Kileskus 2010 Itat Formation (Middle Jurassic, Bathonian) Russia Uncertain if it possesses the head crest as seen in other proceratosaurids Kinnareemimus 2009 Sao Khua Formation (Early Cretaceous, Valanginian to Barremian) Thailand Potentially one of the oldest ornithomimosaurs Klamelisaurus 1993 Shishugou Formation (Middle Jurassic, Callovian) China Close relatives included several referred species of Mamenchisaurus[74] Kol 2009 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Had a "hyperarctometatarsus" more strongly pinched than other arctometatarsalian taxa. Described as an alvarezsaurid[75] but has been suggested to be related to Avimimus[76] Koreaceratops 2011 Sihwa Formation (Early Cretaceous, Albian) South Korea Possessed elongated neural spines on its caudal vertebrae. Its describers suggest that it was used as a swimming organ,[77] but a later study found it to live in a semiarid environment, making this unlikely[78] Koreanosaurus 2011 Seonso Conglomerate (Late Cretaceous, Campanian) South Korea Had short but powerful forelimbs suggesting it may have been a quadruped[79] Koshisaurus 2015 Kitadani Formation (Early Cretaceous, Hauterivian) Japan Distinguished from other hadrosauroids by the presence of an antorbital fossa Kulceratops 1995 Khodzhakul Formation (Early Cretaceous, Albian) Uzbekistan Only known from fragments of a jaw and teeth Kulindadromeus 2014 Ukureyskaya Formation (Middle Jurassic, Bathonian) Russia An ornithischian that preserves evidence of filaments, suggesting that protofeathers were basal to Dinosauria as a whole Kundurosaurus 2012 Udurchukan Formation (Late Cretaceous, Maastrichtian) Russia May be synonymous with Kerberosaurus[80] Kuru 2021 Barun Goyot Formation (Late Cretaceous, Maastrichtian) Mongolia Had been informally referred to as "Airakoraptor" prior to its formal description Laiyangosaurus 2019 Wangshi Group (Late Cretaceous, Maastrichtian) China Some specimens referred to this edmontosaurin actually belong to kritosaurins and lambeosaurines[81] Lanzhousaurus 2005 Hekou Group (Early Cretaceous, Barremian) China Possessed the largest known teeth of any dinosaur Leshansaurus 2009 Shaximiao Formation (Late Jurassic, Oxfordian to Kimmeridgian) China Its braincase is nearly identical to that of Piveteausaurus[82] Levnesovia 2009 Bissekty Formation (Late Cretaceous, Turonian) Uzbekistan One of the smallest known hadrosauroids[42] Liaoceratops 2002 Yixian Formation (Early Cretaceous, Barremian) China One specimen was found without a skull roof, possibly displaced by a predator to eat its brain[83] Liaoningosaurus 2001 Yixian Formation (Early Cretaceous, Barremian to Aptian) China One specimen has been interpreted as possessing fork-like teeth, sharp claws, and stomach contents including fish, which has been claimed to be evidence of a semi-aquatic, turtle-like lifestyle[84] Liaoningotitan 2018 Yixian Formation (Early Cretaceous, Barremian) China The second sauropod named from the Yixian Formation Liaoningvenator 2017 Yixian Formation (Early Cretaceous, Barremian) China Uniquely preserved with the head curving forwards, differing from the classic theropod "death pose" and the sleeping position of other troodontids Limusaurus 2009 Shishugou Formation (Late Jurassic, Oxfordian) China Multiple specimens from different growth stages are known. Juveniles possessed teeth which were lost and replaced with a beak as adults, suggesting a change in diet[85] Lingwulong 2018 Yanan Formation?/Zhiluo Formation? (Middle Jurassic to Late Jurassic, Aalenian to Oxfordian) China The first confirmed diplodocoid from Asia. Originally considered Early Jurassic, making it the oldest known neosauropod, but this age has been disputed[86][87] Lingyuanosaurus 2019 Jiufotang Formation?/Yixian Formation? (Early Cretaceous, Valanginian to Aptian) China Possessed a mix of basal and derived therizinosaurian traits Linhenykus 2011 Bayan Mandahu Formation (Late Cretaceous, Campanian to Maastrichtian) China Completely monodactyl due to lacking the vestigial second and third fingers of other alvarezsaurids Linheraptor 2010 Bayan Mandahu Formation (Late Cretaceous, Campanian) China Potentially a synonym of Tsaagan[88] Linhevenator 2011 Bayan Mandahu Formation (Late Cretaceous, Campanian) China Had a greatly enlarged sickle claw, comparable in size to those of dromaeosaurids Liubangosaurus 2010 Xinlong Formation (Early Cretaceous, Barremian to Aptian) China Described only as a eusauropod[89] but has since been reinterpreted as a somphospondylian[90] Luanchuanraptor 2007 Qiupa Formation (Late Cretaceous, Maastrichtian) China The first Asian dromaeosaurid found outside the Gobi Desert and northeastern China. May have been closely related to Adasaurus[14] Lufengosaurus 1940 Lufeng Formation (Early Jurassic, Hettangian to Sinemurian) China The rib of one specimen preserves the oldest known evidence of collagen proteins[91] Luoyanggia 2009 Haoling Formation (Early Cretaceous, Aptian to Albian) China Originally believed to date from the Late Cretaceous Machairasaurus 2010 Bayan Mandahu Formation (Late Cretaceous, Campanian) China Its hand claws are elongated and blade-like in side view Mahakala 2007 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Possessed basal traits for a dromaeosaurid. May be a close relative of Halszkaraptor[92] Maleevus 1987 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia Its only purportedly distinguishing trait is also shared with Pinacosaurus[36] Mamenchisaurus 1954 Penglaizhen Formation, Shaximiao Formation, Shishugou Formation, Suining Formation (Late Jurassic to Early Cretaceous, Oxfordian to Aptian) China Several species have been named, but most may not belong to this genus[74] Mandschurosaurus 1930 Grès supérieurs Formation?, Yuliangze Formation (Late Cretaceous, Maastrichtian) China
Laos? One of the first non-avian dinosaurs named from Chinese remains Mei 2004 Yixian Formation (Early Cretaceous, Aptian) China Two specimens are preserved in bird-like sleeping positions[93] Microceratus 2008 Ulansuhai Formation (Late Cretaceous, Turonian) China Originally named Microceratops, although that genus name is preoccupied by a wasp Microhadrosaurus 1979 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Reportedly an unusually small hadrosaurid Micropachycephalosaurus 1978 Wangshi Group (Late Cretaceous, Campanian to Maastrichtian) China Once considered to be a pachycephalosaur, although it is now usually considered to be a ceratopsian[94] Microraptor 2000 Jiufotang Formation (Early Cretaceous, Aptian) China Known from over three hundred fossils.[95] Several are well-preserved enough to reveal fine details such as feather covering and an iridescent black coloration[96] Migmanychion 2023 Longjiang Formation (Early Cretaceous, Aptian) China Its hand combines features of multiple groups of coelurosaurs Minimocursor 2023 Phu Kradung Formation (Late Jurassic, Tithonian) Thailand The first basal neornithischian known from southeastern Asia Minotaurasaurus 2009 Djadochta Formation (Late Cretaceous, Campanian) Mongolia The holotype skull was excavated illegally, which obscured its true provenance until recently Mongolosaurus 1933 On Gong Formation (Early Cretaceous, Aptian to Albian) China Known from only scant remains but has been confidently assigned to Somphospondyli in recent years[90] Mongolostegus 2018 Dzunbain Formation (Early Cretaceous, Aptian to Albian) Mongolia Informally assigned to the genus Wuerhosaurus before its formal description Monkonosaurus 1986 Loe-ein Formation?/Lura Formation? (Late Jurassic, Oxfordian to Kimmeridgian?/Early Cretaceous, Albian?) China Poorly known Monolophosaurus 1993 Shishugou Formation (Middle Jurassic, Bathonian to Callovian) China Possessed a short, rectangular crest running along the midline of the skull Mononykus 1993 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Proposed to have an anteater-like lifestyle, using its unique forearms to break into termite mounds[97] Mosaiceratops 2015 Xiaguan Formation (Late Cretaceous, Turonian to Campanian) China Combined features of different groups of basal ceratopsians Nankangia 2013 Nanxiong Formation (Late Cretaceous, Maastrichtian) China May have specialized in soft foods such as leaves and seeds[98] Nanningosaurus 2007 Unnamed formation (Late Cretaceous, Maastrichtian) China Potentially a basal lambeosaurine Nanshiungosaurus 1979 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Originally misidentified as a sauropod on account of its unusual pelvis Nanyangosaurus 2000 Xiaguan Formation (Late Cretaceous, Turonian to Campanian) China Completely lost the first digit of its hands Napaisaurus 2022 Xinlong Formation (Early Cretaceous, Aptian to Albian) China May be closely related to contemporary Thai iguanodonts Natovenator 2022 Barun Goyot Formation (Late Cretaceous, Maastrichtian) Mongolia Possessed a streamlined body and a long, toothed snout, convergently similar to several groups of aquatic vertebrates Nebulasaurus 2015 Zhanghe Formation (Middle Jurassic, Aalenian to Bajocian) China Only known from a single braincase, but it is enough to tell that it was related to Spinophorosaurus Neimongosaurus 2001 Iren Dabasu Formation (Late Cretaceous, Cenomanian) China Could extend its arms considerably forward due to the structure of its shoulder joint[99] Nemegtomaia 2005 Barun Goyot Formation, Nemegt Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia One specimen preserves traces of damage by skin beetles[100] Nemegtonykus 2019 Nemegt Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia The second alvarezsaurid named from the Nemegt Formation Nemegtosaurus 1971 Nemegt Formation, Subashi Formation? (Late Cretaceous, Maastrichtian) China?
Mongolia Had a long, low skull similar in proportions to those of diplodocoids Ningyuansaurus 2012 Yixian Formation (Early Cretaceous, Aptian) China Preserves small oval-shaped structures in its stomach region which may be seeds Nipponosaurus 1936 Yezo Group (Late Cretaceous, Santonian to Campanian) Russia Discovered on the island of Sakhalin, which was owned by Japan in 1936 but later annexed by Russia Oksoko 2020 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Its third finger was so greatly reduced that it was functionally didactyl Olorotitan 2003 Udurchukan Formation (Late Cretaceous, Maastrichtian) Russia Had a broad, hatchet-shaped crest Omeisaurus 1939 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China Members of this genus are characterized by extremely elongated necks Ondogurvel 2022 Barun Goyot Formation (Late Cretaceous, (Campanian) Mongolia Known from well-preserved remains of the hands and feet Opisthocoelicaudia 1977 Nemegt Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia Walked on its metacarpals due to its complete lack of phalanges Oviraptor 1924 Djadochta Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia Originally mistakenly thought to be an egg-eater Pachysuchus 1951 Lufeng Formation (Early Jurassic, Sinemurian to Pliensbachian) China Considered a phytosaur from its original naming until a redescription in 2012[101] Panguraptor 2014 Lufeng Formation (Early Jurassic, Hettangian to Sinemurian) China The first definitive coelophysoid known from Asia Papiliovenator 2021 Bayan Mandahu Formation (Late Cretaceous, Campanian) China Had a short, subtriangular skull similar to those of Early Cretaceous troodontids Paralitherizinosaurus 2022 Yezo Group (Late Cretaceous, Campanian Japan Had stiffened claws that may have been used to pull vegetation to the mouth[102] Parvicursor 1996 Barun Goyot Formation (Late Cretaceous, Campanian) Mongolia Originally believed to represent a diminutive adult dinosaur, although it was recently reinterpreted as a juvenile[103] Pedopenna 2005 Haifanggou Formation (Middle Jurassic, Callovian) China Known from a single leg with the impressions of long, symmetrical feathers Peishansaurus 1953 Minhe Formation (Late Cretaceous, Santonian to Campanian) China Has been compared to thyreophorans and marginocephalians, but it is impossible to determine which assignment is correct Penelopognathus 2005 Bayin-Gobi Formation (Early Cretaceous, Albian) China Named from a single dentary Phaedrolosaurus 1973 Lianmuqin Formation (Early Cretaceous, Valanginian to Albian) China May have been a dromaeosaurid[104] Philovenator 2012 Bayan Mandahu Formation (Late Cretaceous, Campanian) China Closely related to the contemporary Linhevenator[93] but likely represents a separate taxon[105] Phuwiangosaurus 1994 Sao Khua Formation (Early Cretaceous, Valanginian to Hauterivian) Thailand A large member of the Euhelopodidae[90] Phuwiangvenator 2019 Sao Khua Formation (Early Cretaceous, Barremian) Thailand Combines features of both allosauroids and coelurosaurs[106] Pinacosaurus 1933 Bayan Mandahu Formation, Djadochta Formation (Late Cretaceous, Santonian to Campanian) China
Mongolia May have been capable of bird-like vocalizations[107] Plesiohadros 2014 Alagteeg Formation (Late Cretaceous, Campanian) Mongolia The first hadrosauroid known from the Alagteeg Formation Prenocephale 1974 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Had a distinctively conical dome Probactrosaurus 1966 Dashuigou Formation (Early Cretaceous, Albian) China The closest relative to the Hadrosauromorpha based on the definition of the group[108] Prodeinodon 1924 Öösh Formation, Xinlong Formation (Early Cretaceous, Barremian to Aptian) China
Mongolia Potentially a carnosaur[109] Protarchaeopteryx 1997 Yixian Formation (Early Cretaceous, Aptian) China Usually thought to be a basal oviraptorosaur but one study suggests a basal position within Pennaraptora[14] Protoceratops 1923 Bayan Mandahu Formation, Djadochta Formation (Late Cretaceous, Campanian) China
Mongolia Its remains are so abundant that it has been nicknamed the "sheep of the Cretaceous" Protognathosaurus 1991 Shaximiao Formation (Middle Jurassic, Bathonian to Callovian) China Originally named Protognathus, but that is preoccupied by an extinct beetle[110] Psittacosaurus 1923 Andakhuduk Formation, Bayin-Gobi Formation, Ejinhoro Formation, Ilek Formation, Jiufotang Formation, Khok Kruat Formation, Öösh Formation, Qingshan Formation, Tugulu Group, Xinminbao Group, Yixian Formation (Early Cretaceous, Barremian to Albian) China
Mongolia
Russia
Thailand Known from hundreds of specimens, many of them well-preserved. Lived in a broad range Pukyongosaurus 2001 Hasandong Formation (Early Cretaceous, Aptian to Albian) South Korea One of its caudal vertebrae has bite marks caused by theropod teeth Qianlong 2023 Ziliujing Formation (Early Jurassic, Sinemurian) China Associated with fossils of leathery eggs, the oldest of their kind in the world Qianzhousaurus 2014 Nanxiong Formation (Late Cretaceous, Maastrichtian) China Has been nicknamed "Pinocchio rex" on account of its elongated snout Qiaowanlong 2009 Xinminbao Group (Early Cretaceous, Albian) China Originally described as a brachiosaurid[111] but has since been reinterpreted as a euhelopodid[112] Qijianglong 2015 Suining Formation (Early Cretaceous, Aptian) China Once believed to date from the Late Jurassic Qingxiusaurus 2008 Unnamed formation (Late Cretaceous, Maastrichtian) China Known from very limited remains Qinlingosaurus 1996 Hongtuling Formation?/Shanyang Formation? (Late Cretaceous, Maastrichtian) China Potentially a titanosaur given its age, but this cannot be confirmed Qiupalong 2011 Qiupa Formation (Late Cretaceous, Campanian to Maastrichtian) China A referred specimen was found in Canada[113] Qiupanykus 2018 Qiupa Formation (Late Cretaceous, Maastrichtian) China May have used its robust thumb claws to crack open oviraptorid eggshells[114] Quaesitosaurus 1983 Barun Goyot Formation (Late Cretaceous, Maastrichtian) Mongolia Potentially a close relative of Nemegtosaurus Ratchasimasaurus 2011 Khok Kruat Formation (Early Cretaceous, Aptian) Thailand Only known from a single toothless dentary Rhomaleopakhus 2021 Kalaza Formation (Late Jurassic, Tithonian) China Possessed a robust forelimb that may be a locomotory adaptation Rinchenia 1997 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Had a tall, domed crest Ruixinia 2022 Yixian Formation (Early Cretaceous, Barremian) China Its last few caudal vertebrae were fused into a rod-like structure Ruyangosaurus 2009 Haoling Formation (Early Cretaceous, Aptian to Albian) China Only known from scant remains but was one of the largest dinosaurs known from Asia Sahaliyania 2008 Yuliangze Formation (Late Cretaceous, Maastrichtian) China Possibly a synonym of Amurosaurus[115] Saichania 1977 Barun Goyot Formation, Nemegt Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia Possessed complicated nasal passages that may have cooled the air it breathed Sanpasaurus 1944 Ziliujing Formation (Early Jurassic, Toarcian) China Historically conflated with the remains of an ornithischian Sanxiasaurus 2019 Xintiangou Formation (Middle Jurassic, Bajocian) China The oldest neornithischian known from Asia Saurolophus 1912 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Type species was found in Canada. The Asian species is distinguished by its larger size and backwards-pointing diagonal crest Sauroplites 1953 Zhidan Group (Early Cretaceous, Barremian to Aptian) China Preserved lying on its back with parts of its armor in an articulated position Saurornithoides 1924 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Its hindlimbs were well-developed even as juveniles, suggesting it needed little to no parental care Scansoriopteryx 2002 Haifanggou Formation (Middle Jurassic to Late Jurassic, Callovian to Kimmeridgian) China Was well-adapted for climbing due to the structure of its hands and feet Segnosaurus 1979 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Turonian) Mongolia One of the first known therizinosaurs. Its relationships were originally obscure Serikornis 2017 Tiaojishan Formation (Middle Jurassic to Late Jurassic, Callovian to Oxfordian) China Possessed simple, wispy feathers similar to those of a Silkie chicken Shamosaurus 1983 Dzunbain Formation (Early Cretaceous, Aptian to Albian) Mongolia The osteoderms on its head were not separated into obvious tiles as with later ankylosaurs Shanag 2007 Öösh Formation (Early Cretaceous, Berriasian to Barremian) Mongolia Shows a mixture of traits of various paravian groups Shantungosaurus 1973 Wangshi Group (Late Cretaceous, Campanian) China The largest known hadrosaurid Shanxia 1998 Huiquanpu Formation (Late Cretaceous, Cenomanian to Campanian) China May be synonymous with Tianzhenosaurus[116] and/or Saichania[36] Shanyangosaurus 1996 Shanyang Formation (Late Cretaceous, Maastrichtian) China Indeterminate but its hollow bones are a synapomorphy for Coelurosauria. One study suggests an oviraptorosaurian position[14] Shaochilong 2009 Ulansuhai Formation (Late Cretaceous, Cenomanian to Turonian) China Had a relatively short maxilla, suggesting a unique ecological role Shenzhousaurus 2003 Yixian Formation (Early Cretaceous, Aptian) China Preserves pebbles in its thoracic cavity which may be gastroliths Shidaisaurus 2009 Chuanjie Formation (Middle Jurassic, Aalenian) China Potentially one of the oldest known allosauroids Shishugounykus 2019 Shishugou Formation (Late Jurassic, Oxfordian) China Its manus combines features of both alvarezsaurians and more basal coelurosaurs Shixinggia 2005 Pingling Formation (Late Cretaceous, Maastrichtian) China Known from a fair amount of postcranial material Shri 2021 Barun Goyot Formation (Late Cretaceous, Maastrichtian) Mongolia Before its formal description, it was nicknamed "Ichabodcraniosaurus" because its holotype lacked a skull Shuangmiaosaurus 2003 Sunjiawan Formation (Early Cretaceous, Albian) China Only known from some parts of a skull Shunosaurus 1983 Shaximiao Formation (Late Jurassic, Oxfordian to Kimmeridgian) China Possessed a small tail club topped by two short spikes Shuvuuia 1998 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Displays several adaptations that may point to a nocturnal, owl-like lifestyle[117] Siamodon 2011 Khok Kruat Formation (Early Cretaceous, Aptian) Thailand May have been closely related to Probactrosaurus[118] Siamosaurus 1986 Khok Kruat Formation, Sao Khua Formation (Early Cretaceous, Barremian to Aptian) Thailand Only known from teeth. Some spinosaurid postcrania from the same area may be referrable to this genus[119] Siamotyrannus 1996 Sao Khua Formation (Early Cretaceous, Berriasian to Barremian) Thailand Has been recovered in a variety of positions within Avetheropoda Sinankylosaurus 2020 Wangshi Group (Late Cretaceous, Campanian) China Only known from an ilium. Described as an ankylosaur but a recent study doubts this interpretation[120] Sinocalliopteryx 2007 Yixian Formation (Early Cretaceous, Barremian to Aptian) China Stomach contents indicate a possible preference for volant prey such as dromaeosaurids and early birds[121] Sinocephale 2021 Ulansuhai Formation (Late Cretaceous, Turonian) China Originally named as a species of Troodon when that genus was thought to be a pachycephalosaur Sinoceratops 2010 Wangshi Group (Late Cretaceous, Campanian to Maastrichtian) China Possessed forward-curving hornlets and a series of low knobs on the top of the frill Sinocoelurus 1942 Kuangyuan Series (Late Jurassic, Oxfordian to Tithonian China One study considered it to be a potential plesiosaur[122] Sinornithoides 1993 Ejinhoro Formation (Early Cretaceous, Aptian to Albian) China Preserved in a roosting position, its head tucked underneath its left wing Sinornithomimus 2003 Ulansuhai Formation (Late Cretaceous, Turonian) China Formed age-segregated herds as evidenced by a concentration of juvenile skeletons[123] Sinornithosaurus 1999 Yixian Formation (Early Cretaceous, Barremian to Aptian) China One specimen has disloged teeth, leading to suggestions it was venomous[124] Sinosauropteryx 1996 Yixian Formation (Early Cretaceous, Barremian) China The first non-avian dinosaur found with direct evidence of feathers. Analysis of melanosomes suggest it had orange-brown and white countershading with a striped tail and a "bandit mask" around its eyes[125] Sinosaurus 1940 Lufeng Formation (Early Jurassic, Hettangian to Sinemurian) China Had a pair of midline crests similar to Dilophosaurus Sinotyrannus 2009 Jiufotang Formation (Early Cretaceous, Aptian) China One of the earliest known large tyrannosauroids. Closely related to smaller forms such as Proceratosaurus and Guanlong Sinovenator 2002 Yixian Formation (Early Cretaceous, Barremian) China Some specimens are preserved three-dimensionally Sinraptor 1993 Shishugou Formation (Late Jurassic, Oxfordian) China May have used its teeth like blades to inflict deep wounds in prey Sinusonasus 2004 Yixian Formation (Early Cretaceous, Hauterivian) China Had distinctive sinusoid nasal bones Sirindhorna 2015 Khok Kruat Formation (Early Cretaceous, Aptian) Thailand Its fossils were discovered by corn farmers while digging a reservoir Sonidosaurus 2006 Iren Dabasu Formation (Late Cretaceous, Cenomanian to Campanian) China One of the smallest known titanosaurs Stegosaurides 1953 Xinminbao Group (Early Cretaceous, Hauterivian to Albian) China A thyreophoran of uncertain phylogenetic position Suzhousaurus 2007 Xinminbao Group (Early Cretaceous, Barremian to Aptian) China One of the largest Early Cretaceous therizinosaurs Szechuanosaurus 1942 Kuangyuan Series (Late Jurassic, Oxfordian to Tithonian) China Only known from teeth and possibly a very fragmentary skeleton Talarurus 1952 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia Its tail club has been compared to a wicker basket Tambatitanis 2014 Sasayama Group (Early Cretaceous, Albian) Japan Possessed disproportionately large chevrons Tangvayosaurus 1999 Grès supérieurs Formation (Early Cretaceous, Aptian to Albian) Laos Closely related to Phuwiangosaurus Tanius 1929 Wangshi Group (Late Cretaceous, Campanian to Maastrichtian) China Today known from only a few bones; more fossils were once present but were not collected Taohelong 2013 Hekou Group (Early Cretaceous, Albian) China Possessed a sacral shield similar to that of Polacanthus Tarbosaurus 1955 Nemegt Formation, Subashi Formation (Late Cretaceous, Maastrichtian) China
Mongolia An apex predator that hunted large prey. Very similar to Tyrannosaurus Tarchia 1977 Barun Goyot Formation, Nemegt Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia One specimen preserves injuries to its ribs and tail, possibly from a fight with a member of its own kind[126] Tatisaurus 1965 Lufeng Formation (Early Jurassic, Sinemurian) China Potentially a basal thyreophoran Tengrisaurus 2017 Murtoi Formation (Early Cretaceous, Barremian to Aptian) Russia Closely related to South American titanosaurs Therizinosaurus 1954 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Possessed extremely elongated and stiffened hand claws Tianchisaurus 1993 Toutunhe Formation (Late Jurassic, Oxfordian to Kimmeridgian) China Its description uses the spellings Tianchisaurus and Tianchiasaurus interchangeably, but the former is correct[127] Tianyulong 2009 Tiaojishan Formation (Late Jurassic, Oxfordian) China Preserves impressions of long bristles down its back, tail and neck Tianyuraptor 2009 Yixian Formation (Early Cretaceous, Barremian to Aptian) China Combines features of both northern and southern dromaeosaurids. Had unusual proportions Tianzhenosaurus 1998 Huiquanpu Formation (Late Cretaceous, Cenomanian to Campanian) China May be synonymous with Saichania[36] Tienshanosaurus 1937 Shishugou Formation (Late Jurassic, Oxfordian) China Large but basal for a mamenchisaurid[74] Timurlengia 2016 Bissekty Formation (Late Cretaceous, Turonian) Uzbekistan Its inner ear was specialized for detecting low-frequency sounds[128] Tochisaurus 1991 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Known from only a single metatarsus Tonganosaurus 2010 Yimen Formation (Early Jurassic, Pliensbachian) China Potentially the oldest known mamenchisaurid Tongtianlong 2016 Nanxiong Formation (Late Cretaceous, Maastrichtian) China The pose of the holotype suggests it died while trying to free itself from mud Tsaagan 2006 Djadochta Formation (Late Cretaceous, Campanian) Mongolia Very similar to Velociraptor but differs in some features of the skull[129] Tsagantegia 1993 Bayan Shireh Formation (Late Cretaceous, Cenomanian to Santonian) Mongolia Had a long, shovel-shaped snout which may indicate a browsing lifestyle[130] Tsintaosaurus 1958 Wangshi Group (Late Cretaceous, Campanian) China Originally mistakenly believed to have possessed a unicorn horn-like crest Tugulusaurus 1973 Lianmuqin Formation (Early Cretaceous, Barremian to Albian) China Potentially an early, Xiyunykus-grade alvarezsaurian[131] Tuojiangosaurus 1977 Shaximiao Formation (Late Jurassic, Oxfordian to Kimmeridgian) China Possessed two rows of tall, pointed plates, thickened in the center as if they were modified spikes Turanoceratops 1989 Bissekty Formation (Late Cretaceous, Turonian) Uzbekistan Had a pair of brow horns like ceratopsids but was likely not a member of that family Tylocephale 1974 Barun Goyot Formation (Late Cretaceous, Campanian) Mongolia Only known from a partial skull but it is enough to tell that it had a remarkably tall dome Tyrannomimus 2023 Kitadani Formation (Early Cretaceous, Aptian) Japan Its ilium is remarkably similar to that of the supposed tyrannosauroid Aviatyrannis Udanoceratops 1992 Djadochta Formation (Late Cretaceous, Campanian) Mongolia The largest known leptoceratopsid Ultrasaurus 1983 Gugyedong Formation (Early Cretaceous, Aptian to Albian) South Korea Described as very large but this may be due to misidentification of a bone Ulughbegsaurus 2021 Bissekty Formation (Late Cretaceous, Turonian) Uzbekistan Known from only a maxilla. Originally described as a late-surviving carnosaur but may in fact be a large-bodied dromaeosaurid[132] Urbacodon 2007 Bissekty Formation, Dzharakuduk Formation (Late Cretaceous, Cenomanian to Turonian) Uzbekistan The holotype preserves a gap separating the eight rear teeth from the rest of its teeth Vayuraptor 2019 Sao Khua Formation (Early Cretaceous, Barremian) Thailand Potentially ancestral to megaraptorans[133] or an early member of the group[134] Velociraptor 1924 Bayan Mandahu Formation, Djadochta Formation (Late Cretaceous, Campanian) China
Mongolia One potential specimen preserves quill knobs[135] Wakinosaurus 1992 Sengoku Formation (Early Cretaceous, Valanginian to Barremian) Japan May be a close relative of Acrocanthosaurus[109] Wannanosaurus 1977 Xiaoyan Formation (Late Cretaceous, Maastrichtian) China Basal for a pachycephalosaur as indicated by its flat skull with large openings Wuerhosaurus 1973 Ejinhoro Formation, Tugulu Group (Early Cretaceous, Hauterivian) China One of the last and largest known stegosaurs. Preserved with low rectangular plates but these may be broken Wulagasaurus 2008 Yuliangze Formation (Late Cretaceous, Maastrichtian) China A rare hadrosaurid known from far less remains than the contemporary Sahaliyania Wulatelong 2013 Bayan Mandahu Formation (Late Cretaceous, Campanian) China Known from a partial skeleton including some parts of the skull Wulong 2020 Jiufotang Formation (Early Cretaceous, Aptian) China Analysis of preserved melanosomes suggests it was mostly gray with iridescent wings[136] Xianshanosaurus 2009 Haoling Formation (Early Cretaceous, Aptian to Albian) China May have been closely related to Daxiatitan[90] Xiaosaurus 1983 Shaximiao Formation (Middle Jurassic, Bajocian to Callovian) China An ornithischian of uncertain affinities Xiaotingia 2011 Tiaojishan Formation (Middle Jurassic to Late Jurassic, Bathonian to Oxfordian) China Well-preserved but inconsistent in phylogenetic placement. Some studies suggest a position as an early avialan[137] Xingtianosaurus 2019 Yixian Formation (Early Cretaceous, Barremian) China Retained the large third finger that was lost in other caudipterids Xingxiulong 2017 Lufeng Formation (Early Jurassic, Hettangian) China Possessed a robust scapula which increased forelimb mobility for feeding Xinjiangovenator 2005 Lianmuqin Formation (Early Cretaceous, Valanginian to Albian) China Remains originally identified as Phaedrolosaurus Xinjiangtitan 2013 Qiketai Formation (Middle Jurassic, Callovian) China Had an extremely long neck Xiongguanlong 2009 Xinminbao Group, (Early Cretaceous, Aptian) China More robust than other early tyrannosauroids, possibly to support its elongated skull Xixianykus 2010 Majiacun Formation (Late Cretaceous, Santonian to Coniacian) China One of the smallest known non-avian dinosaurs Xixiasaurus 2010 Majiacun Formation (Late Cretaceous, Coniacian to Campanian) China Distinguished from other troodontids by its possession of exactly twenty-two teeth in each maxilla Xixiposaurus 2010 Lufeng Formation (Early Jurassic, Hettangian to Toarcian) China Poorly known Xiyunykus 2018 Tugulu Group (Early Cretaceous, Barremian to Aptian) China Had an unspecialized hand morphology for an alvarezsaur, having three fingers of roughly equal length and construction Xuanhanosaurus 1984 Shaximiao Formation (Middle Jurassic to Late Jurassic, Bathonian) China Originally mistakenly believed to have been capable of quadrupedal locomotion Xuanhuaceratops 2006 Houcheng Formation (Late Jurassic, Tithonian) China Possessed a large premaxillary tooth right behind its beak Xunmenglong 2019 Huajiying Formation (Early Cretaceous, Hauterivian) China The holotype was originally presented as part of a chimera involving three different animals[138] Xuwulong 2011 Xinminbao Group (Early Cretaceous, Aptian to Albian) China The tip of its dentary was V-shaped when viewed from the side Yamaceratops 2006 Javkhlant Formation (Late Cretaceous, Santonian) Mongolia Possessed a short, stubby frill Yamatosaurus 2021 Kita-Ama Formation (Late Cretaceous, Maastrichtian) Japan Basal yet survived late enough to be contemporaneous with more advanced hadrosaurids Yandusaurus 1979 Shaximiao Formation (Middle Jurassic, Bathonian) China Some fossils were destroyed by a composter before they could be studied[139] Yangchuanosaurus 1978 Shaximiao Formation (Middle Jurassic to Late Jurassic, Bathonian to Oxfordian) China The largest theropod known from the Shaximiao Yi 2015 Tiaojishan Formation (Middle Jurassic to Late Jurassic, Callovian to Oxfordian) China Possessed a "styliform element" jutting out from its wrist that supported a bat-like membranous wing Yimenosaurus 1990 Fengjiahe Formation (Early Jurassic, Pliensbachian) China Much of its skeleton is known, including the entirety of the skull Yingshanosaurus 1994 Shaximiao Formation (Late Jurassic, Kimmeridgian) China Possessed greatly enlarged shoulder spines Yinlong 2006 Shishugou Formation (Late Jurassic, Oxfordian) China Its skull displays features of ceratopsians, pachycephalosaurs, and heterodontosaurids Yixianosaurus 2003 Yixian Formation (Early Cretaceous, Aptian) China Inconsistent in phylogenetic placement. Had extremely elongated manual elements Yizhousaurus 2018 Lufeng Formation (Early Jurassic, Sinemurian) China Its skull was very similar to those of sauropods, despite being more primitive Yongjinglong 2014 Hekou Group (Early Cretaceous, Albian) China Possessed an extremely long, broad scapula Yuanmousaurus 2006 Zhanghe Formation (Middle Jurassic, Aalenian to Callovian) China Shares features of its vertebrae with Patagosaurus Yueosaurus 2012 Liangtoutang Formation (Early Cretaceous to Late Cretaceous, Albian to Cenomanian) China Probably closely related to Jeholosaurus[140] Yulong 2013 Qiupa Formation (Late Cretaceous, Maastrichtian) China Known from multiple specimens, most of which are juveniles Yunganglong 2013 Zhumapu Formation (Late Cretaceous, Cenomanian) China Discovered 50 kilometres (31 mi) away from a World Heritage Site Yunmenglong 2013 Haoling Formation (Early Cretaceous, Barremian to Albian) China May have been exceptionally large Yunnanosaurus 1942 Fengjiahe Formation, Lufeng Formation (Early Jurassic, Sinemurian to Pliensbachian) China Its teeth were self-sharpening similar to those of sauropods, likely through convergent evolution[141] Yunyangosaurus 2020 Xintiangou Formation (Middle Jurassic to Late Jurassic, Aalenian to Oxfordian) China Potentially an early megalosauroid Yutyrannus 2012 Yixian Formation (Early Cretaceous, Aptian) China The largest known dinosaur that preserves direct evidence of feathers Yuxisaurus 2022 Fengjiahe Formation (Early Jurassic, Sinemurian to Toarcian) China Had more than one hundred osteoderms Yuzhoulong 2022 Shaximiao Formation (Middle Jurassic, Bathonian) China One of the oldest known macronarians Zanabazar 2009 Nemegt Formation (Late Cretaceous, Maastrichtian) Mongolia Originally named as a species of Saurornithoides. A large troodontid Zaraapelta 2014 Barun Goyot Formation (Late Cretaceous, Campanian to Maastrichtian) Mongolia Had an intricate pattern of osteoderms on its skull Zhanghenglong 2014 Majiacun Formation (Late Cretaceous, Santonian) China Reconstructed by its describers with a straight, rectangular back, although no complete neural spines are known[142] Zhejiangosaurus 2007 Chaochuan Formation (Late Cretaceous, Cenomanian) China Has no diagnostic features[36] Zhenyuanlong 2015 Yixian Formation (Early Cretaceous, Aptian) China Possessed large wings with long feathers, but was most likely flightless Zhongjianosaurus 2017 Yixian Formation (Early Cretaceous, Barremian to Aptian) China Distinguishable by its characteristically elongated legs. Described as a microraptorian[143] but it has been noted that some features of its skeleton are similar to avialans[39] Zhuchengceratops 2010 Wangshi Group (Late Cretaceous, Maastrichtian) China Had a particularly deep mandible Zhuchengtitan 2017 Wangshi Group (Late Cretaceous, Campanian) China The proportions of its humerus suggest a close relationship with Opisthocoelicaudia[144] Zhuchengtyrannus 2011 Wangshi Group (Late Cretaceous, Campanian) China Closely related to Tarbosaurus and Tyrannosaurus Zigongosaurus 1976 Shaximiao Formation (Middle Jurassic to Late Jurassic, Bathonian to Tithonian) China May be a species of Mamenchisaurus[145] Zizhongosaurus 1983 Ziliujing Formation (Early Jurassic, Toarcian) China Poorly known but was most likely basal for a sauropod Zuolong 2010 Shishugou Formation (Late Jurassic, Oxfordian China Known from both cranial and postcranial remains
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https://dokumen.pub/the-princeton-field-guide-to-dinosaurs-course-booknbsped-9781400836154.html
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en
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The Princeton Field Guide to Dinosaurs [Course Book ed.] 9781400836154
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This lavishly illustrated volume is the first authoritative dinosaur book in the style of a field guide. World-renowned...
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dokumen.pub
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https://dokumen.pub/the-princeton-field-guide-to-dinosaurs-course-booknbsped-9781400836154.html
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Table of contents :
CONTENTS
Preface
Introduction
Group and Species Accounts
Dinosaurs
Theropods
Sauropodomorphs
Ornithischians
Additional Reading
Index
Citation preview
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https://en.wikipedia.org/wiki/Ankylosauridae
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en
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Ankylosauridae
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2006-01-25T07:52:55+00:00
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https://en.wikipedia.org/wiki/Ankylosauridae
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Extinct family of dinosaurs
Ankylosaurids
Mounted skeleton of Scolosaurus thronus, Royal Tyrrell Museum of Palaeontology Scientific classification Domain: Eukaryota Kingdom: Animalia Phylum: Chordata Clade: Dinosauria Clade: †Ornithischia Clade: †Thyreophora Clade: †Ankylosauria Clade: †Euankylosauria Family: †Ankylosauridae
Brown, 1908 Type species †Ankylosaurus magniventris
Brown, 1908
Subgroups
†Aletopelta
†Bissektipelta
†Cedarpelta
†Chuanqilong
†Crichtonsaurus
†Liaoningosaurus
†Maleevus?
†Minmi
†Ankylosaurinae
†Shamosaurinae
†Gobisaurus
†Shamosaurus
Synonyms
Syrmosauridae Maleev, 1952
Ankylosauridae ( ) is a family of armored dinosaurs within Ankylosauria, and is the sister group to Nodosauridae. The oldest known Ankylosaurids date to around 122 million years ago and went extinct 66 million years ago during the Cretaceous–Paleogene extinction event.[1] These animals were mainly herbivorous and were obligate quadrupeds, with leaf-shaped teeth and robust, scute-covered bodies. Ankylosaurids possess a distinctly domed and short snout, wedge-shaped osteoderms on their skull, scutes along their torso, and a tail club.[2]
Ankylosauridae is exclusively known from the northern hemisphere, with specimens found in western North America, Europe, and East Asia. The first discoveries within this family were of the genus Ankylosaurus, by Peter Kaiser and Barnum Brown in Montana in 1906.[3] Brown went on to name Ankylosauridae and the subfamily Ankylosaurinae in 1908.
Anatomy
[edit]
Ankylosaurids are stout, solidly built, armoured dinosaurs. They possess accessory ossifications on cranial bones that cover some skull openings and form wedge-shaped, horn-like structures. Along the ankylosaurid torso are scute rows, which are filled in with smaller ossicles to create a fused shield of armour.[2] Only two collars of armour plates can be found on the neck, as opposed to the sister group, nodosaurids, which have three.[1] Nodosauridae and Ankylosauridae also share the unique attribute of abundant structural fibres in both primary and secondary bone.[4] Ankylosaurids also have an S-shaped narial passage.[1]
The most distinguishing feature of ankylosaurids is the presence of a tail club. It is made out of modified interlocking distal caudal vertebrae and enlarged, bulbous osteoderms.[5] The "handle" of the tail club involves the vertebrae, and requires elongated prezygapophyses to overlap at least half of the preceding vertebral centrum length.[5] These distal caudal vertebrae also lack transverse processes and neural spines, and therefore tend to be longer than they are wide; the reverse is true for proximal caudal vertebrae.[5] Derived ankylosaurids possess a fusion of posterior dorsal, sacral, and sometimes anterior caudal vertebrae, which forms a singular structure called a "synsacrum complex". There is a complete fusion between centra, neural arches, zygapophyses, and sometimes neural spines.[6]
In 2017, Victoria M. Arbour and David C. Evans described a new genus of ankylosaurine that preserved extensive soft tissues along the body. This animal, named Zuul after its resemblance to the Ghostbusters monster, is also the first ankylosaur from the Judith River Formation.[7]
History of study
[edit]
Barnum Brown and Peter Kaisen discovered the first ankylosaurid genus, Ankylosaurus, in 1906 in the Hell Creek Beds in Montana.[3] The fossil material they found was a portion of the skull, two teeth, some vertebrae, a distorted scapula, ribs and more than thirty osteoderms.[3] Reconstruction of the specimen was initially met with skepticism by those who believed it to be at least very close to, or completely a part of the genus Stegopelta, and Brown himself placed it within the suborder Stegosauria.[3]
It has previously been interpreted that variation in ankylosaurid tail club shape is due to sexual dimorphism, which assumes that tail club morphology has a sex-linked intraspecific function.[6] This is possible if the tail club was used for agonistic behaviour. However, a sexual dimorphism theory would predict roughly equal numbers of individuals with two distinct sizes of tail clubs. Obvious sexual dimorphism has not been documented, but if the clubs of one sex are much larger, then there would be a bias for preservation and discovery towards that sex.[6][8]
Phylogeny
[edit]
In 1978, W.P. Coombs, Jr. classified almost all valid species of Ankylosauria within either Nodosauridae or Ankylosauridae.[9] This was a pivotal study and described many characters of ankylosaurs in the earliest phylogenetic analyses of the group.
Later in 1998, Paul Sereno formally defined Ankylosauridae as all ankylosaurs more closely related to Ankylosaurus than to Panoplosaurus.[10] Ankylosaurs not known to possess a tail club were included in Kenneth Carpenter's 2001 phylogeny of Ankylosauridae.[11]
In a study done in 2004 by Vickaryous et al., Gargoyleosaurus, Gastonia, and Minmi were recorded as basal ankylosaurids, with the rest of the ankylosaurids filled out with Gobisaurus, Shamosaurus, and ankylosaurines from China, Mongolia, and North America.[12]
In 2012, Thompson et al. undertook an analysis of almost all known valid ankylosaurs and outgroup taxa at the time.[13] They based their resulting phylogeny on characters representing cranial, post-cranial, and osteodermal anatomy, and details of synapomorphies for each recovered clade. This study placed Gargoyleosaurus and Gastonia within basal Nodosauridae, and put Cedarpelta and Liaoningosaurus as basal ankylosaurids.[13]
In 2016, Arbour and Currie have presented a phylogenetic analysis of Ankylosauridae consisting of Gastonia, Cedarpelta, Chuanqilong, other basal ankylosaurids, and a number of derived ankylosaurids. Their phylogeny includes some uncertain phylogenetic relationships, between Ankylosaurus, Anodontosaurus, Scolosaurus, and Ziapelta.[14]
Paleobiology
[edit]
Posture and locomotion
[edit]
Ankylosaurids were likely very slow-moving animals. In all Ankylosauria, the fibula is more slender than the tibia, suggesting that the tibia carried most of the weight of the animal, while the fibula served as an area of muscular attachment.[15] Hindlimb muscles of Euoplocephalus have been restored and the placement of several muscles inserting on the femur have very short moment arms. Muscles inserting on the tibia and fibula have longer moment arms. This pattern of retractor muscles points to an elephantine locomotion, consistent with columnar posture.[15]
Restoration of Euoplocephalus forelimbs demonstrate similarities to crocodilian forelimb musculature.[16] The most well developed muscles in the pectoral region had more of a weight-bearing function than a rotational one. It has also been postulated that the carpals and metacarpals bear resemblance to those of tetrapods with fossorial (burrowing) habits.[16]
Several muscles in the posterior of ankylosaurids (dorsalis caudae, ilio-caudalis, coccygeo-femoralis brevis, coccygeo-femoralis longus, ilio-tibialis, and ischio caudalis) were used for motion of the tail and tail club.[15] Ankylosaurids tend to have horizontal rather than an obliquely vertical orientation of zygapophyseal articulations in the free caudal vertebrae of the tail. This arrangement is most effective for side-to-side rather than vertical mobility.[6] The absence of musculature to elevate the tail, and this orientation of zygapophyses suggest that the tail and its club swept parallel to and slightly above the ground.[6]
Biogeography
[edit]
It is difficult to establish the geographical origin of Ankylosauridae at present. There is a mix of basal ankylosaurids from both North America and Asia, which carries on through accepted cladistic analyses.[17] It appears that in the mid-Cretaceous, Asian nodosaurids were replaced by ankylosaurine ankylosaurids.[14] Some researchers postulate that Ankylosaurines migrated into North America from Asia between the Albian and Campanian, where they diversified into a clade of ankylosaurines characterized by arched snouts and flat cranial bone plates (caputegulae).[14] There is no evidence for ankylosaurids in Gondwana.[14]
Variation
[edit]
Within Ankylosauridae there is much individual and interspecific variation in expression of armour. However, the most researched aspect of ankylosaurid armour is the tail club. There has been substantial ontogenetic and individual variability found in the morphology of this feature. There have been at least 16 caudal vertebrae included in the handle of the tail club of Pinacosaurus grangeri, and Euoplocephalus has an estimated 9 – 11 coossified caudals.[6]
Variations in tail knob shape, thickness, and length are attributed to individual variation, taxonomy, or representation of different growth phases.[6] There are difficulties with this last aspect, however, in that known clubs do not conform to a single growth series, yet some differences must be ontogenetic and allometric.[6][8]
Lifestyle
[edit]
Most ankylosaurid teeth were leaf-shaped, implying a mainly herbivorous diet. Their teeth could be smooth or fluted, or may differ on labial and lingual surfaces.[18] Euoplocephalus tutus possess ridges and grooves on their teeth that have no relation to their marginal cusps.[18] With their downward-facing neck and head, it is plausible for ankylosaurids to feed in a grazing pattern.[1]
Non-herbivorous habits have been implicated for some species, however. Pinacosaurus has been speculated as being an ant-eater-like long tongued insectivore,[19] while Liaoningosaurus has been proposed to be a piscivore. Either would be exceptional evidence of carnivory among ornithischians.
There are a few prevailing theories for ankylosaurid tail club function. The first is agonistic behaviour within a species.[6] In most vertebrates, including dinosaurs, this behaviour is accompanied by structures for display or combat. Some researchers believe this phenomenon would have been implausible considering there is no modern tetrapod analogue that uses the tail for this purpose. These paleontologists instead propose that ankylosaurids made use of their broad, flat skull for head-butting between individuals.[6]
The second theory for tail club function is for defense against predators. It has been postulated that the club would be most effective against the metatarsals of an attacking theropod.[6][15]
The bones that form cranial ornamentation have physiological costs, and so would be inefficient to produce merely for protection against predation. The theory has therefore been posed that these wedge-shaped osteoderms could support a partly sexually selected interpretation.[8]
Timeline of genera
[edit]
See also
[edit]
Dinosaurs portal
Timeline of ankylosaur research
References
[edit]
Dinosaurs and other Prehistoric Creatures, edited by Ingrid Cranfield (2000), Salamander books, pg. 250–257.
Carpenter K (2001). "Phylogenetic analysis of the Ankylosauria". In Carpenter, Kenneth (ed.). The Armored Dinosaurs. Indiana University Press. pp. 455–484. ISBN 978-0-253-33964-5.
Kirkland, J. I. (1996). Biogeography of western North America's mid-Cretaceous faunas - losing European ties and the first great Asian-North American interchange. J. Vertebr. Paleontol. 16 (Suppl. to 3): 45A
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https://www.academia.edu/6019911/The_Basal_Nodosaurid_Ankylosaur_Europelta_carbonensis_n_gen_n_sp_from_the_Lower_Cretaceous_Lower_Albian_Escucha_Formation_of_Northeastern_Spain
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The Basal Nodosaurid Ankylosaur Europelta carbonensis n. gen., n. sp. from the Lower Cretaceous (Lower Albian) Escucha Formation of Northeastern Spain
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Nodosaurids are poorly known from the Lower Cretaceous of Europe. Two associated ankylosaur skeletons excavated from the lower Albian carbonaceous member of the Escucha Formation near Ariño in northeastern Teruel, Spain reveal nearly all the
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https://www.academia.edu/6019911/The_Basal_Nodosaurid_Ankylosaur_Europelta_carbonensis_n_gen_n_sp_from_the_Lower_Cretaceous_Lower_Albian_Escucha_Formation_of_Northeastern_Spain
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The Zorralbo locality of the eastern Cameros Basin, near Soria, Spain, has produced a diverse dinosaur assemblage from the Lower Cretaceous Golmayo Formation. Ankylosaurs are represented by dorsal vertebrae and ribs, a fragmentary sacrum and ilium, and several types of dermal armour. Most, if not all, of the material probably belongs to a single medium to large-sized adult individual. The Soria remains are referred to Polacanthus on the basis of the presence of conical, ungrooved presacral spines, a sacropelvic shield composed of irregularly arranged bosses and small tubercles, large spined plates, and hollow-based triangular caudal plates with an extended posterior basal edge and a pointed spine. Polacanthus is well known from the Wealden Group (Barremian-Aptian) of the Isle of Wight and from the Weald Clay Group of West Sussex (England). In addition, isolated remains have been reported from the penicontemporaneous formations of the Iberian Peninsula. The Soria outcrop is currently the most productive Polacanthus site outside England. Moreover, it has yielded the oldest record (late Hauterivian to basal Barremian according to charophyte association) of this ankylosaur known to date in Europe. Minor anatomic differences between the Soria material and the taxa P. foxii (type-species) and P. rudgwickensis suggest the presence of a third species of Polacanthus in the Iberian Peninsula, but additional material is needed to confirm this interpretation.
Nodosaurid ankylosaur remains from the Upper Cretaceous of Mexico are summarized. The specimens are from the El Gallo Formation of Baja California, and the Pen and Aguja Formations of northwestern Coahuila, Mexico. These specimens show significant differences from other known nodosaurids, including ulna with very well developed olecranon and prominent humeral notch, the distal end of the femur not flaring to the extent seen in other nodosaurids, and a horn-like spine with vascular grooves on one side. The specimens are important because they are the southern-most occurrences in North America, and provide an important biogeographical link between nodosaurids of the United States and Canada on the one hand, and Argentina and Antarctica on the other.
Sources of morphological variation (dimorphism, individual variability, ontogeny, and pathology) can obfuscate taxonomic variation. This is true for Late Cretaceous nodosaurid ankylosaurs, for which several complete, well-preserved skulls and postcrania are known. Although a generalized taxonomy has been accepted for about 25 years, new specimens show mixtures of features considered diagnostic for more than one taxon. Because these taxa are well-accepted and overlap temporally and geographically, they are good candidates for testing intraspecific variation in dinosaurs. This was done by quantitatively testing taxonomic characters a priori with bivariate and clustering analyses. A character-specimen matrix was coded for a parsimony analysis to aid in taxonomic referrals. Because many phylogenetic characters are based on relative proportions or shapes, taphonomic distortion is problematic for this group. Nevertheless, four taxa are valid: Edmontonia longiceps, E. rugosidens, Panoplosaurus mirus, and Denversaurus schlessmani, the latter two more derived. Compared to contemporaneous North American ankylosaurids, nodosaurid taxa do not correlate as well with their stratigraphic distribution. A posteriori character analysis reveals that Panoplosaurus has a shortened skull and rounded cervical/pectoral osteoderms. Denversaurus and Panoplosaurus share inflated cranial sculpturing with visible sulci between individual elements. Denversaurus has a relatively wider anterior snout. The clade shows some overall evolutionary trends: doming of the skull over the orbits, thickening of the vomer and closure of prevomer foramen, encroachment of sculpturing over the anterior temporal bar, shortening and widening of the snout, etc. Use of proportional character data is common in dinosaur systematics, often subjectively. This study demonstrates that quantitatively testing such characters increases their repeatability and clarifies taxonomic decisions.
New stegosaurian remains have been recently recovered from the Jurassic-Cretaceous transition sandstones of the Villar del Arzobispo Formation (Tithonian-Berriasian) in the Valencia province, eastern Spain. Specimens consist of two partially articulated (or closely associated) postcranial skeletons. The Baldovar specimen is composed of appendicular bones (scapula, femur) and two pairs of dermal tail spines, two of them articulated with two distal caudal vertebrae. The second specimen, unearthed in the vicinity of La Yesa village, consists of dorsal vertebrae and ribs, fragments of caudal centra and an incomplete femur. The new specimens are tentatively referred to the clade Dacentrurinae and may belong to Dacentrurus on the basis of features observed on the dorsal vertebrae and caudal dermal spines. Stegosaurs are represented so far in the Jurassic-Cretaceous transition of Spain by Dacentrurus. The presence of other taxa (Stegosaurus, Miragaia) in Spain, recently documented in the Late Jurassic of Portugal, cannot be attested on the basis of the currently recorded material.
We present new nodosaurid teeth from the Valanginian of Bexhill, Sussex and the Barremian of the Isle of Wight, the first from the Lower Cretaceous of the United Kingdom. Teeth found during the mid-1800s from the Valanginian and ascribed to the nodosaurid Hylaeosaurus are probably from sauropod dinosaurs. The Isle of Wight tooth could possibly be referred to Polacanthus foxii, the teeth of which are unknown. These new English nodosaurid teeth are similar to those of North American and European Jurassic to Late Cretaceous nodosaurids, especially the American Gastonia, Texasetes,
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The Theropod Database Blog: Raven et al. 2023 on ankylosaur phylogeny missed the shortest trees and should not have dropped Nodosauridae
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A new paper came out analyzing thyreophoran phylogenetics - Raven et al. (2023). The abstract states "This dataset was analysed using equal...
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http://theropoddatabase.blogspot.com/2023/05/raven-et-al-2023-on-ankylosaur.html
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A new paper came out analyzing thyreophoran phylogenetics - Raven et al. (2023). The abstract states "This dataset was analysed using equal- and implied-weights parsimony and Bayesian inference, and further explored using constraint trees and partitioned datasets. Stratigraphical congruence was used to identify a 'preferred tree' and these analyses reveal a novel hypothesis for thyreophoran relationships. The traditional ankylosaurian dichotomy is not supported: instead, four distinct ankylosaur clades are identified, with the long-standing 'traditional' clade Nodosauridae rendered paraphyletic. Ankylosauridae, Panoplosauridae, Polacanthidae and Struthiosauridae have distinct morphotypes..."
Four distinct ankylosaur clades with distinct morphotypes?
It sounds intriguing, but then Polacanthidae as a separate clade or grade has been a viable hypothesis since 1998, so I guess it's the paraphyly of Panoplosaurus and Struthiosaurus versus Ankylosaurus that is new? Looking back, that was ambiguous in Vickaryous et al. (2001), Osi (2005) and Parsons and Parsons (2009) but rejected by Osi and Makadi (2009), Thompson et al. (2012) and Arbour et al. (2016), thus it would be surprising to someone like myself who doesn't specialize in ornithischians so is not all that familiar with character support and such.
So let's see what Raven et al. found. They did five basic analyses (A-D with [edited thanks to David Marjanovic's comment] progressively more different assumed weighting- unweighted, then k = 3, 8 and 12; E as a Bayesian attempt) then some constraint analyses and ones using only certain parts of the skeleton. I'm going to ignore the latter as I don't think anyone considers "cranial only", "postcranial only" or "armor only" to give better results in Mesozoic dinosaur phylogenetics. In any case, the first analysis (Analysis A) was equal weights, which is the standard for Mesozoic dinosaur analyses and how basically every prior ankylosaur analysis was run. As an aside, statements like "The strict consensus tree (Supplementary material, Fig. S75) shows a lack of resolution in Stegosauria" .... "and Ankylosauria is found in an unresolved polytomy with most stegosaur taxa" just means you haven't pruned enough taxa a posteriori to see the underlying structure. It's annoying the authors never look into what that structure is and leave the tree looking artificially uncertain. But the main result in Ankylosauria is... "There are two clades within Ankylosauria (Ankylosauridae + Nodosauridae)"! So the expected usual result. Hmm. But what about their novel hypothesis of three nodosaur clades at least?
They state "Within Nodosauridae, there are three groupings of taxa: 'polacanthid' ankylosaurs, but excluding Polacanthus; a 'panoplosaurid' group typified by Edmontonia and Panoplosaurus; and a 'struthiosaurid' group typified by Struthiosaurus and Hungarosaurus." But are there? Just look at the cladogram above (which is only in the supp info). 'Polacanthidae excluding Polacanthus' is Texasetes, the then-unnamed Patagopelta, Sauropelta, Hylaeosaurus, Hoplitosaurus, Tatankacephalus, an undescribed Wessex specimen, Zhejiangosaurus, Antarctopelta and Dongyangopelta. Besides Hoplitosaurus and Hylaeosaurus (which I don't know has ever been recovered as a polacanthid, just guessed to be there), I don't think any of these have been associated with Polacanthidae/inae before. The 'core' polacanthids besides Polacanthus itself are Gastonia, Mymoorapelta and maybe Gargoyleosaurus, so I wouldn't say this clade is reflective of Polacanthidae in any sense. But okay you say, maybe it's not Polacanthidae, but surely the analyses (PLURAL as in the abstract) revealed a 'distinct morphotype' for this Sauropelta-Hylaeosaurus clade of nodosaurs? Looking at their "preferred tree" (Analysis B, k=3) only Patagopelta, Texasetes, Hoplitosaurus and Hylaeosaurus are shared between 'polacanthid' clades, with five new taxa in there compared to Analysis A. And in Analysis C (k=8) only Hoplitosaurus and Texasetes are shared (with seven new taxa), while by Analysis D (k=12) all these taxa are scattered to the wind and there's no equivalent at all. And Analysis E (Bayesian) is one huge polytomy for ankylosaurs. So I would suggest their published results do not support any clade like this that is robust when analyzed under different criteria.
But what about Panoplosauridae and Struthiosauridae? First of all, just as there are nine 'nodosaurs' that fall outside the Nodosauridae+ Ankylosauridae split, there are one to five nodosaurids that don't fall into either the panoplosaur or struthiosaur clades. We can't tell how many because Raven et al. do not prune a posteriori to try to resolve any polytomies. But the panoplosaur group is Dracopelta, Aletopelta, both Edmontonia species, Denversaurus and of course Panoplosaurus. The latter three have always been grouped together, but the first two would be interesting if they actually were panoplosaurs. The 'preferred tree' (k=3) takes away Aletopelta and adds Nodosaurus, Anoplosaurus and Tianchisaurus; Analysis C (k=8) has the core three genera plus Dracopelta and Anoplosaurus, while Analysis D (k=12) has everything in A plus Anoplosaurus and Nodosaurus. So Dracopelta is always a panoplosaur and Anoplosaurus is with any unequal weighting, which are the first surprising, new and widely supported nodosaur placements in this study. Yet neither is touted in the text or written up with character support, and honestly the idea of Albian-Cenomanian English Anoplosaurus and especially Jurassic European Dracopelta breaking up not just Campanian-Maastrichtian North American Panoplosaurus+Edmontonia, but the genus Edmontonia itself(!) just seems unlikely. And indeed, a fairly complete Dracopelta specimen (Russo and Mateus, 2023) was recently discovered, and the authors found "D. zbyszewskii [was] consistently recovered as sister taxa of G[argoyleosaurus] parkpinorum, from the Upper Jurassic of Morrison Formation, USA, in a basal ankylosaur group that also includes the other Morrison Formation ankylosaur, M[ymoorapelta] maysi," which matches temporally so much better.
As for Struthiosauridae, the unweighted tree would have this include Borealopelta, Minmi, Niobrarasaurus, Polacanthus, Europelta, Liaoningosaurus, Stegopelta, the Paw Paw Formation juvenile (* see below), Hungarosaurus and all three Struthiosaurus species (not monophyletic, at least Tianchisaurus is closer to S. transylvanicus than to S. austriacus or S. languedocensis). The 'preferred tree' (k=3) keeps only Struthiosaurus, Hungarosaurus, the Paw Paw juvenile and Europelta, and adds Silvisaurus and Taohelong. Analysis C (k=8) drops the latter two but adds Tianchisaurus back, and Analysis D (k=12) keeps the two and everything from the unweighted tree plus adds Invictarx and Hoplitosaurus. So far from dividing the 'distinct morphotypes' of Polacanthidae and Struthiosauridae, two of five trees have Polacanthus as a struthiosaur. Here the results are besides the Santonian-Maastrichtian Central European Struthiosaurus/Hungarosaurus, the struthiosaur clade includes Europelta (as guessed by its describers), the Early Cretaceous American Paw Paw juvenile and usually the Jurassic Chinese Tianchisaurus. Yet the authors never mention the Paw Paw juvenile ever falling out here, and have Tianchisaurus as a panoplosaurid because Analysis B is the one time it wasn't the sister to Struthiosaurus transylvanicus. In fact, the authors falsely* state the Paw Paw juvenile is usually a basal ankylosaur and Figure 2 incorrectly* shows it being one in a supposed "Agreement subtree of the three implied weighing analyses (analyses B–D)." How did this happen?! ...
(*) I figured it out- the trees in the supp info switched Pawpawsaurus and the Paw Paw juvenile, while figures 1-3 in the paper are correct. So actually their trees have Pawpawsaurus as the struthiosaur sister to Hungarosaurus, which has the same chronostratigraphic issues as the juvenile from the same formation.
If we go back to the big picture in Ankylosauria, the unweighted Analysis A gave us a large Nodosauridae with some mostly Jurassic taxa basal to the Nodo-Ankylo split, and a few 'nodosaurs' (Kunbarrasaurus, Peloroplites, Liaoningosaurus) as basal ankylosaurids. 'Preferred' Analysis B (k=3) has polacanthids basal to the split, then panoplosaurs and struthiosaurs sister to each other, so again is pretty standard. Analysis C (k=8) has struthiosaurs, panoplosaurs and polacanthids successively closer to ankylosaurids, so at least that has the paraphyletic nodosaurs the article touts. Finally, Analysis D (k=12) has struthiosaurs further from ankylosaurids than panoplosaurs while polacanthids cease to really exist (their internal specifier Gastonia [see below] is an ankylosaurid but Polacanthus and Hoplitosaurus are struthiosaurs).
So I guess if I had a takeaway from their published results (**, see below), it would be that Dracopelta (probably incorrectly) and usually Anoplosaurus are panoplosaurs; Europelta, Pawpawsaurus and usually Tianchisaurus (not even supported by the authors) are struthiosaurs; polacanthids are not strongly supported in any form; and nodosaurs become increasingly paraphyletic with more weighting greater values of k (corrected again thanks to David Marjanovic's comment), although you need to get to k=8 for anything really novel. And it's the weighting that is one of my major issues with this paper, because why is k=3 the preferred tree? Because "The stratigraphically most congruent topology, as identified by the four stratigraphical congruence metrics (SCI, RCI, MSM and GER), was Analysis B, and so this was selected as the 'preferred tree'." But if you look at their Table 2 (below), the Bayesian analysis destroys the others at SCI (0.929 vs. 0.438-0.500), but we never get to know what those results are since the authors just leave it as a huge polytomy without further analysis. And in the other three, Analysis B is 16.954, .023 and .005 better respectively, which seems increasingly less important. I have no idea how any of these measures work, but it seems incredibly arbitrary to say whichever k value is best in a majority of four methods wins, ignoring anything quantitative.
(**) Raven et al. didn't find the shortest trees
But now we get to the part where I reveal nothing I said above matters, because Raven et al. didn't get the shortest trees. EDIT BELOW Not even close. Instead of producing "eight MPTs with lengths of 1508 steps", their unweighted matrix Analysis A results in >99999 MPTs of 1464 steps. Here's the real strict consensus with 13 taxa pruned a posteriori for resolution-
As you can see, it's not the same as Raven et al.'s Figure S75. For one, stegosaurs resolve, although weirdly with Toujiangosaurus+Paranthodon as ankylosaurs. For two it's WAY less resolved in Ankylosauria. All those taxa from Zhejiangosaurus through Ahshishlepelta never form a consistent clade with each other or the four ankylosaur clades, and if you prune all nine genera, struthiosaurs, ankylosaurids and 'panoplosaurs'+polacanthids are still a trichotomy. So their matrix doesn't actually show how these taxa relate (besides polacanthids being sister to 'panoplosaurs'). Isn't it ironic though that we do get Raven et al.'s three nodosaur clades including a classic Polacanthinae that includes Gastonia, Gargoyleosaurus and Mymoorapelta in addition to Jurassic Sarcolestes (and basally some parankylosaurs, but the paper was too late to include Stegouros)? Struthiosauridae is only Struthiosaurus spp. plus Hungarosaurus and Tianchisaurus, which again is funny because the latter was not included in the clade by the authors due only to their Analysis B. 'Panoplosauridae' includes a lot more taxa, and yes those basal ones don't clade with each other or successively to the core group no matter how many are pruned, so that's another real polytomy. As for the pruned taxa-
Mongolostegus can at least be sister to Chungkingosaurus or a struthiosaurine.
Adratiklit is part of the Dacentrurus+Stegosaurus clade.
Anodontosaurus and Scolosaurus are part of the Euoplocephalus+Pinacosaurus clade.
Tarchia kielanae is part of the Ankylosaurus+Euoplocephalus clade.
Acantholipan can be a 'panoplosaur', a struthiosaurine or outside (by which I mean a taxon closer to Ankylosaurus than Toujiangosaurus but not part of the struthiosaur, ankylosaurid or polacanthid+'panoplosaur' clades shown, though it could be e.g. sister to any of these clades and thus fall under their definitions).
Borealopelta can be a struthiosaurine, a 'panoplosaur' or outside(?).
Europelta can be a struthiosaurine or outside.
Invictarx can be a struthiosaurine, a 'panoplosaur' or outside.
Nodosaurus is always closer to Panoplosaurus than Silvisaurus.
Patagopelta can be a basal 'panoplosaur' or outside.
Pawpawsaurus can be a struthiosaurine or 'panoplosaur'.
Stegopelta can be a struthiosaurine or outside.
EDIT ADDED 5-21: Thanks to Andrea Cau in the comments for pointing out Raven et al. didn't include their character ordering settings in their txt file. I wrongly assumed it would have a ccode line or a ctype line below the matrix but never scrolled all the way down. I should have been suspicious when I had to manually choose the outgroup instead of them just making Lesothosaurus the first taxon in the matrix. On the one hand, my bad. On the other hand, it's surely best practice to not force your readers to modify the settings of the file you provided to the journal.
In any case, it still doesn't matter because Raven et al. STILL didn't find the shortest trees. Instead of producing "eight MPTs with lengths of 1508 steps", their unweighted matrix Analysis A results in >99999 MPTs of 1506 steps. Here's the real REAL strict consensus with 13 taxa pruned a posteriori for resolution-
Two steps doesn't sound like much, but it's enough to make 'polacanthids' a grade of basal ankylosaurids, make struthiosaurines nodosaurids as in traditional phylogenies and kick Dracopelta and Anoplosaurus out of panoplosaurs. And yes, Nodosaurus is still closer to Panoplosaurus than Silvisaurus or Struthiosaurus. The toplogy still has Tuojiangosaurus+Paranthodon as ankylosaurs and a struthiosaur Tianchisaurus too.
What are these clades named?
Raven et al. propose new definitions for their three nodosaur families-
"Panoplosauridae
All ankylosaurs more closely related to Panoplosaurus than to Ankylosaurus, Struthiosaurus austriacus or Gastonia burgei
Polacanthidae
All ankylosaurs more closely related to Gastonia burgei than to Ankylosaurus, Panoplosaurus or Struthiosaurus austriacus
Struthiosauridae
All ankylosaurs more closely related to Struthiosaurus austriacus than to Ankylosaurus, Panoplosaurus or Gastonia burgei"
Tim Williams has already rightfully complained on the DML that their Polacanthidae definition needs to use Polacanthus foxii. Why is this so hard in 2023?! Do Arbour and the "three anonymous referees" not know the basics of phylogenetic nomenclature? PhyloCode Article 11.10 states "when a clade name is converted from a preexisting name that is typified under a rank-based code or is a new or converted name derived from the stem of a typified name, the definition of the clade name must use the type species of that preexisting typified name or of the genus name from which it is derived (or the type specimen of that species) as an internal specifier." We've been complaining about it since Sereno 24 years ago, surely every dinosaur worker knows by now.
Another obvious issue is that Nodosauridae has priority over Panoplosauridae, Struthiosauridae and Polacanthidae, so where's Nodosauridae? Raven et al. explain "Nodosaurus is recovered outside of
Panoplosauridae in Analyses A and C, further suggesting that application of the name Nodosauridae would add confusion." But as noted above, Nodosaurus is actually always a 'panoplosaur' when Analysis A is run correctly, and the fact it's supposedly an ankylosaurid sister to Dyoplosaurus when k=8 (fig. S77) should be close to worthless in my view. Unless one of the authors wants to claim whatever character is being weighed eight times more than others could realistically mean this Cenomanian taxon is really within a Campanian species complex that are so similar they were all placed under Euoplocephalus tutus until recently? Also, Polacanthus doesn't fall in their definition of Polacanthidae in two of their four trees, so why is that still allowed as a family name?
In any case, the clades have already been officially defined with PhyloCode registrations, by Madzia et al. (2021). Raven et al. was "Received 7 February 2022", so I don't know why Arbour (a coauthor on Madzia et al.!) or the reviewers would let that stay in the paper. Reading through, Raven et al. actually cite Madzia et al. and state "the underlying philosophy of the latter study is based on the PhyloCode (de Queiroz & Cantino, 2020) and offers an alternative hypothesis to our study, which is framed by the traditional principles of the International Commission on Zoological Nomenclature (1999), and so is not discussed further." Hahahaha I hate to tell you guys, but defining clades based on phylogenetic relationships has nothing to do with the ICZN. And if you were following the "traditional principles" of the 1999 ICZN, you couldn't just throw Nodosauridae away while stating "In the 'preferred' tree Panoplosauridae consists of ... Nodosaurus" and "A clade of generally Late Cretaceous North American taxa is also recovered here and named Panoplosauridae. As well as Denversaurus, Edmontonia spp., Nodosaurus and Panoplosaurus..." Instead you would follow ICZN Article 65.2.3 - "by the discovery that the type genus was, when established, based on a type species then misidentified, the author may fix as the type species a nominal species as prescribed in Article 70.3. If the threat cannot be overcome by the fixation of a type species under the provisions of Article 70.3 the case is to be referred to the Commission for a ruling." But that's not happening because I bet they think it's unlikely Nodosaurus is outside Nodosauridae. "... is not discussed further" is short here for "... we know it makes no sense but we don't want to address it."
And Madzia et al. do a good job because they actually follow the rules. Except in Raven et al.'s topology Stegosauridae ends up being the Chungkingosaurus+Eurypoda clade due to Huayangosaurus' weird position outside Eurypoda (which supposedly happens in the Bayesian analysis too), which could be saved by adding Ankylosaurus magniventris as an external specifier. Similarly, Struthiosaurini might benefit from an Ankylosaurus magniventris external specifier due to the polytomy. So here's the actual unweighted results with official clade names-
1. Ankylosauria; 2. Ankylosauridae; 3. Ankylosaurinae; 4. Ankylosaurini; 5. Eurypoda; 6. Huayangosauridae; 7. Nodosauridae; 8. Nodosaurinae; 9. Polacanthinae; 10. Shamosaurinae; 11. Stegosauria; 12. Struthiosaurini. Note due to Nodosaurus' possible positions, Panoplosaurini cannot be placed precisely in this tree.
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Alvarezsaurs in the Lori matrix
This time our topology is-
I provide a new definition for Alvarezsauroidea that adds Therizinosaurus as an external specifier since I find it most parsimonious for Therizinosauria to be its sister group, and uses Alvarezsaurus as the internal specifier unlike Sereno's that uses Shuvuuia- (Alvarezsaurus calvoi< - Ornithomimus velox, Therizinosaurus cheloniformis, Passer domesticus) . Alvarezsauroids have had a controversial phylogenetic placement, with the Lori matrix recovering them as basal maniraptorans sister to therizinosaurs. Yet they can be outside therizinosaurs plus pennaraptorans in 3 steps, become avemetatarsalians in 4 steps (can bring therizinosaurs or not), non-maniraptoriforms in 6 steps (they bring therizinosaurs), closer to pennaraptorans than therizinosaurs in 6 steps, paravians in 11 steps (therizinosaurs move with), closer to Compsognathus than to birds in 15 steps, closer to birds than deinonychosaurs in 27 steps, and closer to Archaeopteryx and other birds than to dromaeosaurids and troodontids in 30 steps.
Fukuivenator is an odd taxon, recovered here as the basalmost alvarezsauroid. But it can be a therizinosaurian in only two steps, and outside Maniraptoriformes in 4 steps (it emerges in Coeluridae). One thing I don't think it is is a dromaeosaurid, as that takes 27 more steps, and getting it into Paraves or Pennaraptora requires 11 and 7 steps respectively. Still, I wouldn't be surprised to see this taxon work its way around the base of Maniraptoriformes once an osteology comes out.
Shuvuuia deserti IGM 100/975 axial elements in ventral view and pelvis in dorsal view (courtesy AMNH).
Nqwebasaurus was recently redescribed by Sereno (2017), which I incorporated into its scorings. Choiniere et al. (2012) recovered it in Ornithomimosauria, but note most of the characters they list to support that are also said to be present in alvarezsauroids. Even they could place it in Alvarezsauroidea with only 4 steps. The Lori matrix needs 6 steps to place it in Ornithomimosauria, which I think is higher partially due to it finding Pelecanimimus to be an alvarezsauroid too. So similarities between the two like their teeth being in a common groove and maxillary teeth being confined to the anterior third of the bone are no longer ornithomimosaur-like. As recently noted by Cerroni et al. (2019), this makes more sense biogeographically as well. Oh, and note that the Lori matrix found Afromimus to be a ceratosaur as in that paper. In any case, Nqwebasaurus takes 10 steps to move to Compsognathidae, and 7 steps to move sister to Pennaraptora.
As for Pelecanimimus itself, it seems plausibly alvarezsauroid if you think about it. The skull is famously similar to Shuvuuia, the posterior tympanic recess is in the otic recess, ossified sterna are otherwise unknown for ornithomimosaurs, the long manual digit I was always out of place compared to Harpymimus, and Europe makes more sense for otherwise Gondwanan clades in the Cretaceous. Now if only someone would release Perez-Moreno's thesis describing it in detail...
Shuvuuia deserti IGM 100/975 pectoral and forelimb elements. Note the tiny phalanx from digit II or III at the bottom (courtesy AMNH).
Patagonykus and Bonapartenykus are usually closer to parvicursorines than Alvarezsaurus and Achillesaurus, but the Lori matrix found them just outside Alvarezsauridae instead. Interestingly, Xu et al. (2018) recovered the same results. It takes 3 steps to move Patagonykus closer to parvicursorines, and 4 steps to join Alvarezsaurus and Patagonykus to the exclusion of parvicursorines as in Alifanov and Barsbold (2009). Xu et al. recover these in 5 and 7 steps respectively, and the most recent version of Longrich and Currie's alvarezsaurid matrix (Lu et al., 2018) recovers a basal Patagonykus and a basal Parvicursorinae in 3 steps each.
One odd result is that the newly described Xiyunykus and Bannykus fall in Patagonykinae too. Yet only 2 steps move them outside the Patagonykus plus Parvicursorinae clade, where they form a clade. Another step breaks that up to place Xiyunykus more basal as in Xu et al.. Them being basal certainly fits better stratigraphically, and Xu et al. use several characters designed for alvarezsauroids that the Lori matrix didn't include yet. Hopefully full osteologies will be published as well.
Mononykus olecranus cast YPM 56693 (of holotype) pes in plantar view (courtesy of Senter).
A patagonykine Achillesaurus as suggested by Agnolin et al. (2012) takes 7 additional steps in the Lori matrix where it instead emerges just closer to parvicursorines than Alvarezsaurus. On the other hand, only a single step joins it with Alvarezsaurus as in Longrich and Currie (2009) and only 2 steps makes it just further from parvicursorines than Alvarezsaurus as in Xu et al. (2018).
Alnashetri is known from type hindlimb material, but now also from MPCA 377, a nearly complete specimen with interesting characters like flat and unfused sternal plates. Makovicky et al. (2016) used this data to recover it as the sister group to Alvarezsauridae, and while the few published details left it more derived in the Lori tree, it can go to a more basal position with only two steps. It should be interesting to compare to e.g. Bannykus once it is published.
Mononykus olecranus cast YPM 56693 (of holotype) (courtesy of Senter).
The arctometatarsal clade has a unique topology, but no other analysis has included nearly as many characters or all of these taxa, with Lu et al. omitting Albinykus and Ceratonykus among non-fragmentary specimens, and Xu et al. omitting the more recently described Qiupanykus. Enforcing the Lori topology in Lu et al.'s matrix is only 5 steps longer, and doing so in Xu et al.'s matrix is only 6 steps longer. On the other hand, Xu et al.'s topology is so unresolved at this level, the only difference in mine is placing the Albinykus plus Xixianykus clade basally near Albertonykus, which takes 5 steps to do in the Lori matrix.
It should be noted that Lu et al.'s illustrated topology (their Figure 3) is not their matrix's real result, as they did not fully analyze tree space. Instead of 20 trees, there are 214 trees. These differ in that Albertonykus, YPM 1049 and undescribed 41HIII-0104 can fall out anywhere more derived than Patagonykus, and that Parvicursor, the Tugriken Shireh taxon, Shuvuuia and Mononykus form an unresolved polytomy. This leaves Linhenykus, Qiupanykus and Xixianykus unresolved between that polytomy and Patagonykus, which is perfectly compatible with the Lori topology. This may also show that the small alvarezsauroid-specific matrix of Longrich and Currie is insufficient given all the new taxa described since 2009. YPM 1049 was far too fragmentary to include (distal metatarsal III) but I tried testing undescribed Quipa specimen 41HIII-0104. Didn't make it into the publication, but here's its scorings-
'41HIII0104' ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ???????-?? ?????????1 ?10??????? ?????????? ?????????? ???0?????? ????1????? ???1?????{01} ?????????? ?????????? ?????????? ????????0? ????????3? ?????????? ?????????? ?????????? ?????????? ?????????1 ?????????1 1????????? 1????????? ?????????? ?????????? ?????????? ???1?????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ????????0? ?????????? ?????????? ?????????? ?????????? ?????????? ???1?????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????{123}???? ?????????? ?????????? ?????????? ?????????? ?????????? ??1???0???
Tugriken Shireh parvicursorine (IGM 100/99) vertebrae and ilia in ventral view, forelimb and fibula in lower right (courtesy AMNH).
Interestingly, Agnolin et al., Xu et al. and the Lori analysis all recovered Albinykus sister to Xixianykus outside Parvicursorinae. Wonder if that's a real signal? Unfortunately, the only attempt to name this clade was Agnolin et al. who also recovered Ceratonykus in there and called it Ceratonykini. Xu et al. place Ceratonykus closer to parvicursorines, while I found it more basal than either, sister to Qiupanykus which neither of the other studies used. Forcing Ceratonykus sister to Albonykus plus Xixianykus takes 3 more steps in the Lori matrix. Forcing Ceratonykus sister to Mononykus as in its original description (with or without Qiupanykus) takes 5 more steps. As stated in the paper, we were the first analysis to include Hateg tibiotarsi Bradycneme and Heptasteornis. While the former can fall into many positions in Maniraptora, the latter was resolved as an alvarezsaurid as proposed by Naish and Dyke (2004). Note this used only the tibiotarsus and not alvarezsaurid-like distal femur FGGUB R.1957. A single step moves Heptasteornis to Troodontidae.
We also provide an updated definition for Parvicursorinae (Mononykus olecranus + Parvicursor remotus), like Choiniere et al.'s (2010) but using species. One accident of our definitional and discovery history is that all these newer arctometatarsal alvarezsaurids (Xixianykus, Albertonykus, Albinykus, Linhenykus, Qiupanykus, Ceratonykus, etc.) emerge outside the originally discovered and defined Parvicursorinae. We could really use some clade defining taxa closer to Mononykus than Patagonykus, Alvarezsaurus or Achillesaurus. In any case, I got a lot of experience with parvicursorine specimens, examining Shuvuuia and the Tugriken Shireh specimen IGM 100/99 in person, and having photos of high quality casts of Mononykus thanks to Senter. I found the Tugriken Shireh taxon closer to Shuvuuia, but moving it closer to Parvicursor as in Longrich and Currie is just 1 step longer.
Tugriken Shireh parvicursorine (IGM 100/99) vertebrae and ilia in dorsal view, forelimb and fibula in lower right (courtesy AMNH).
Next time, therizinosaurs...
References- Naish and Dyke, 2004. Heptasteornis was no ornithomimid, troodontid, dromaeosaurid or owl: The first alvarezsaurid (Dinosauria: Theropoda) from Europe. Neus Jahrbuch für Geologie und Paläontologie. 7, 385-401.
Alifanov and Barsbold, 2009. Ceratonykus oculatus gen. et sp. nov., a new dinosaur (?Theropoda, Alvarezsauria) from the Late Cretaceous of Mongolia. Paleontological Journal. 43(1), 94-106.
Longrich and Currie, 2009. Albertonykus borealis, a new alvarezsaur (Dinosauria: Theropoda) from the Early Maastrichtian of Alberta, Canada: Implications for the systematics and ecology of the Alvarezsauridae. Cretaceous Research. 30(1), 239-252.
Choiniere, Xu, Clark, Forster, Guo and Han, 2010. A basal alvarezsauroid theropod from the early Late Jurassic of Xinjiang, China. Science. 327, 571-574.
Agnolin, Powell, Novas and Kundrat, 2012. New alvarezsaurid (Dinosauria, Theropoda) from uppermost Cretaceous of north-western Patagonia with associated eggs. Cretaceous Research. 35, 33-56.
Makovicky, Apesteguia and Gianechini, 2016. A new, almost complete specimen of Alnashetri cerropoliciensis(Dinosauria: Theropoda) impacts our understanding of alvarezsauroid evolution. XXX Jornadas Argentinas de Paleontologia de Vertebrados. Libro de resumenes, 74.
Sereno, 2017. Early Cretaceous ornithomimosaurs (Dinosauria: Coelurosauria) from Africa. Ameghiniana. 54, 576-616.
Lu, Xu, Chang, Jia, Zhang, Gao, Zhang, Zhang and Ding, 2018. A new alvarezsaurid dinosaur from the Late Cretaceous Qiupa Formation of Luanchuan, Henan Province, central China. China Geology. 1, 28-35.
Xu, Choiniere, Tan, Benson, Clark, Sullivan, Zhao, Han, Ma, He, Wang, Xing and Tan, 2018. Two Early Cretaceous fossils document transitional stages in alvarezsaurian dinosaur evolution. Current Biology. 28, 1-8. DOI: 10.1016/j.cub.2018.07.057
Cerroni, Agnolin, Egli and Novas, 2019. The phylogenetic position of Afromimus tenerensis Sereno, 2017 and its paleobiogeographical implications. Journal of African Earth Sciences. DOI: 10.1016/j.jafrearsci.2019.103572
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
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Therizinosaurs in the Lori matrix
Next up are therizinosaurs. These are one of the best analyzed clades because I incorporated all of Zanno's (2010) characters, which is by far the largest and most recent analysis of the group until the Lori paper was published. The topology is-
Falcarius is the most basal taxon shown of course, but Martharaptor was pruned a posteriori and can fall out anywhere in Therizinosauria outside the Alxasaurus plus Segnosaurus clade. I tried including Thecocoelurus, but the Lori matrix is pretty terrible when it comes to scoring single vertebrae-
Thecocoelurus ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ???????(01)0? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ???1?????? ?????????? ?????2???? ?????????? ?????????? ?????????? ?????????? ???????0?? ?????????? ?????????? ?????????? ?????1???? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ??0??????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ??????????
Jianchangosaurus fell out in the same place as its original description, with Cau's (2018) placement in Alvarezsauroidea taking 18 more steps so is very unlikely. Mine is the only published matrix besides Senter (2011) and its derivatives to use information from the second Beipaiosaurus specimen, and incorporated photos from Zanno and the new paper on the holotype skull elements too.
Beipiaosaurus inexpectus holotype (IVPP V11559) cervical vertebra in dorsal view (courtesy of Zanno).
Zanno also provided photos of Alxasaurus and Enigmosaurus, and its depressing how much of the former is lost. Enigmosaurus rather famously was shown by Zanno to not resemble Barsbold's original illustration that was the only reference picture known for over two decades. Placing Enigmosaurus closer to Segnosaurus than Neimongosaurus or Erliansaurus as in Zanno's tree takes 4 more steps. Forcing Enigmosaurus and Erlikosaurus to be sister taxa to simulate the synonymy mentioned by Barsbold (1983) takes 4 steps, so seems unlikely. The duo moves between Nanshiungosaurus and the Segnosaurus plus Nothronychus clade. We were the first analysis to include "Chilantaisaurus" zheziangensis, which emerged in a polytomy with Alxasaurus, Enigmosaurus and therizinosaurids.
Alxasaurus elesitaiensis holotype (IVPP V88402a) chevrons in right lateral view (natural order reversed) (courtesy of Zanno).
As was the case with Archaeornithomimus? bissektensis, we didn't include the possible chimaera of Bissekty Therizinosauria as an OTU, unlike Sues and Averianov (2015). But if you do want to experiment with it, here's the scorings. It emerges in a polytomy in the Suzhousaurus plus Therizinosaurus clade of therizinosaurids. Btw, Archaeornithomimus? bissektensis does fall out most parsimoniously sister to A. asiaticus when all Bissekty material is used.
'Bissekty-Therizinosauroidea' ????1??0?? ????1????? ?????1???? ?1???????? ?????{01}1000 ?01??????? 00???????? 2??21?0??? ???????00{123} {12}00(01)2011?? ??0???0(01)00 ??11????{12}? ?{01}0?100??? ?????????? ?1{01}?0????? ??0?0???{012}0 000(01)?010?? ?????????? ?????????? ?????0{12}?00 1?0???0??0 ??0??{01}0??? 0?1??????? ????0?1?0? {01}0?1???{01}?? ??0??????? ?????????? ?????????0 ???001??00 ??00?01?1? 10???????? ?????????1 ???0?????? ???1?????? ??????0??1 0???1(12)???? ????0???1? ????0?011? ???0????0? ???0{12}0???? ????????0? 0?0????1?? ?????????? ??010101?1 ??0?0(01)???? ?????{01}010? ???100???? ????????11 1010?????? ??0??0???? ?????????? ?????????? ??????0??? ??00?????? ?00??????? ?????????? ???????0-- -??00??010 ?????????? ??????-??? ???-010??1 0100????00 10?1?00??? ???0?00??? ?????????? ????0????? ???000???? ?0???-???? ?????????? ?????001??
Enigmosaurus mongoliensis holotype (IGM 100/84) synsacrum and ilium in ventral view (courtesy of Zanno).
Next is Therizinosauridae itself, which we refined Zhang et al.'s (2001) definition of to include type species. Therizinosaurids first split into a clade of Erliansaurus, Neimongosaurus, Suzhousaurus and Therizinosaurus. Forcing the former two to be outside a clade of Suzhousaurus, Therizinosaurus and the taxa below, as in Zanno's tree, takes 5 more steps. Notably, we did not include the hindlimb IGM 100/45 in the Therizinosaurus OTU since there's no overlap and its not even particularly large. But here's the Therizinosaurus OTU including the hindlimb. Using this version of Therizinosaurus leaves the tree basically the same but destabilizes it somewhat in that Therizinosaurus and Erlikosaurus can now go in multiple positions within Therizinosauridae, and the Nanchao embryos are in a trichotomy with the Suzhousaurus clade and the Nothronychus clade. Is this an indication the hindlimb produces homoplasy and so might not belong to Therizinosaurus?
Therizinosaurus ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ????????01 0110021000 01?0001100 00001110?? ?????????? ?????????? ?????????? ???0???010 1?0000000? 0?1??1???? ?????????? ?????????? 000??0???? ??????3??1 ?????????0 ???00????? 0?1?0?1??? ?????0???? ?????????1 ?????????? ?????0???? ?????????? ?????????? 1011010101 100100??1? ????{12}?00?? 00?12110?? 101??????? 0????0?111 00?010???? ????????1? ?????????? ????100111 1101110??? ?????????? ????????1? ?1???????? ?????????1 ???0?????? ?000?????? ?????0???1 ?????????? ?????????? ?????????? ?????????? ?????0???? ??????-00? ?00-000001 0?0100??0? 10000???0{01} ?{01}?0000??0 {01}0???????? ????????00 ?00?????0- -??11-???? ?????????? ???1?0???0
Segnosaurus galbinensis paratype (IGM 100/83) cervical neural arch in right lateral view (courtesy of Zanno).
Now comes the Nanchao therizinosaur embryos, those described by Kundrat et al. inside dendroolithid eggs. While including such young specimens might be seen as risky, my ontogenetically conservative scoring method with state N seems to have worked fine here. They fall out where you'd expect a Santonian-Campanian therizinosaur to do so. Following that is Nanshiungosaurus brevispinus, which Senter et al. (2012) recovered as the next most derived therizinosaur after Alxasaurus. Forcing it into this basal position takes 4 steps. Nanshiungosaurus? bohlini was included but pruned a posteriori since it can go anywhere in the Segnosaurus plus Nothronychus clade. Forcing Nanshiungosaurus monophyly is just a single step longer though, while forcing bohlini to be sister to the contemporaneous Suzhousaurus takes 2 steps.
Segnosaurus itself (which Zanno also provided photos of) pairs with ex-Alectrosaurus forelimb AMNH 6368, which has only previously been analyzed by Zanno (2006) where it pairs with Erliansaurus. Forcing that here compared to other taxa she included results in trees 3 steps longer. Erlikosaurus groups with the Nothronychus species in a trichotomy where it can be sister to either species. Forcing Nothronychus monophyly takes only a single step, but note that no proposed Nothronychus characters involve elements that can be compared to Erlikosaurus (humerus and pes). Forcing Erlikosaurus to group with Therizinosaurus as in Senter et al. requires only a single step, with Erlikosaurus moving to the Therizinosaurus clade.
Next time, oviraptorosaurs...
References- Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Transactions of the Joint Soviet-Mongolian Palaeontological Expedition. 19, 117 pp.
Zhang, Xu, Sereno, Kwang and Tan, 2001. A long-necked therizinosauroid dinosaur from the Upper Cretaceous Iren Dabasu Formation of Nei Mongol, People’s Republic of China. Vertebrata PalAsiatica. 39(4), 282-290.
Zanno, 2006. The pectoral girle and forelimb of the primitive therizinosauroid Falcarius utahensis (Theropoda, Maniraptora): Analyzing evolutionary trends within Therizinosauroidea. Journal of Vertebrate Paleontology. 26(3), 636-650.
Zanno, 2010. A taxonomic and phylogenetic re-evaluation of Therizinosauria (Dinosauria: Maniraptora). Journal of Systematic Palaeontology. 8(4), 503-543.
Senter, 2011. Using creation science to demonstrate evolution 2: Morphological continuity within Dinosauria. Journal of Evolutionary Biology. 24(10), 2197-2216.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid tail. PLoS ONE. 7(5), e36790.
Sues and Averianov, 2015. Therizinosauroidea (Dinosauria: Theropoda) from the Upper Cretaceous of Uzbekistan. Cretaceous Research. 59, 155-178.
Cau, 2018. The assembly of the avian body plan: A 160-million-year long process. Bollettino della Società Paleontologica Italiana. 57(1), 1-25.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
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Happy New Year 2020
Hi all. A Theropod Database update is online, with the main additions being troodontid information and info from the Hayashibara Museum of Natural Sciences Research Bulletins 1-3. I love these publications and wish more like them existed for other collections. They detail the expeditions into Mongolia with exact discovery dates and field numbers for taxa like Nomingia, Elsornis and Aepyornithomimus, and tons of still undescribed specimens. It's amazing just how many ornithomimosaurs are known from the Bayanshiree Formation for instance, when only the Garudimimus holotype has been described. There are over twenty more including the sort-of-described "Gallimimus""mongoliensis" specimen IGM 100/14. So often for new taxa, especially those from the Jehol biota, no information is provided in the description as to when the specimen was discovered. I get that many are found by non-professionals and given to museums, but at least say "the specimen was given to the museum on x-x-xx by someone who said it was excavated around year y." Next up, halszkaraptorine and dromaeosaurid updates...
undescribed ?Gobivenator skull (HMNS coll.; field number 940801 TS-I WTB) (after Tsogtbaatar and Chinzorig, 2010).
Reference- Tsogtbaatar and Chinzorig, 2010. Fossil specimens prepared in Mongolian Paleontological Center: 2002–2008. Hayashibara Museum of Natural Sciences Research Bulletin. 3, 155-166.
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Details on Teinurosaurus and random musings
Hi all. When updating The Theropod Database I noticed my entry for Teinurosaurus is pathetically bad- wrong authors, wrong age, wrong size, and generally missing the complicated history of this innocuous vertebra. How embarrassing! So here's the revised version that will be uploaded-
Teinurosaurus Nopcsa, 1928
= Saurornithoides Nopsca, 1928 (preoccupied Osborn, 1924)
= Caudocoelus Huene, 1932
T. sauvagei (Huene, 1932) Olshevsky, 1978
= Caudocoelus sauvagei Huene, 1932
Tithonian, Late Jurassic
Mont-Lambert Formation, Hauts-de-France, France
Holotype- (BHN2R 240; = Boulogne Museum 500) incomplete distal caudal vertebra (75 mm)
Diagnosis- Provisionally indeterminate relative to Kaijiangosaurus, Tanycolagreus and Ornitholestes.
Other diagnoses- (after Huene, 1932; compared to Elaphrosaurus) centrum wider; narrower ventral surface; ventral median groove wider; transversely narrower prezygapophyses.
While Huene attmpted to distinguish Teinurosaurus from Elaphrosaurus, only the wider median ventral groove is apparent in existing photos of the former. This is compared to the one distal caudal of the latter figured in ventral view, but as Kobayashi reports grooves become distally narrower in Harpymimus while Ostrom reports they become distally wider in Deinonychus, groove width is not considered taxonomically distinctive at our current level of understanding. Indeed, this lack of data is most relevent to both diagnosing and identifying Teinurosaurus. Very few taxa have detailed descriptions of distal caudal vertebrae or more than lateral views figured, let alone indications of variation within the distal caudal series. So the facts that Fukuiraptor and Deinonychus share ventrally concave central articulations with Teinurosaurus in their single anteriorly/posteriorly figured distal caudal vertebra, or that Afromimus, "Grusimimus" and Falcariusalso have have wide ventral grooves in their few ventrally figured distal caudals, are not considered taxonomically important.
Comments- Sauvage (1897-1898; in a section written in January 1898) first mentioned a distal caudal vertebra he referred to the ornithischian Iguanodon prestwichii (now recognized as the basal styracosternan Cumnoria prestwichii) - "We are disposed to regard as belonging to the same species the caudal vertebra of a remote region, the part which we figure under n ° 7, 8" [translated]. Note Galton (1982) was incorrect in claiming Sauvage reported on this specimen in his 1897 paper (written December 6), which includes a section on prestwichiinearly identical to the 1897-1898 one but which lacks the paragraph describing this vertebra. This could provide a specific date of December 1897 to January 1898 for the discovery and/or recognition of the specimen. Huene (1932) correctly noted Sauvage mislabeled plate VII figure 8 as dorsal view, when it is in ventral view as understood by the text. Compared to Cumnoria, the caudal is more elongate (length 3.93 times posterior height compared to 2.54 times at most), has a ventral median groove instead of a keel, and the prezygapophyseal base in 71% of the anterior central height compared to ~30-40%, all typical of avepods. Nopcsa (1928) recognized its theropod nature and in his list of reptile genera meant to use a footnote to propose Teinurosaurusas a "new name for the piece described and figured by Sauvage (Direct. Traveaux Geol. Portugal Lisbonne 1897-1898, plate VII, Fig. 7-10) as late caudal of Iguanodon Prestwichi." Teinurosaurusis listed as an aublysodontine megalosaurid (not as an ornithomimine, contra Galton), roughly equivalent to modern Eutyrannosauria. However due to a typographical error, the footnote's superscript 1 was placed after Saurornithoides instead of Teinurosaurus. Sauvage (1929) corrected this in an addendum- "footnote 1 does not refer to Saurornithoides (line 19 from below) but to Teinurosaurus(last line of text)." Unfortunately, Huene missed the addendum, and thus wrote "Nopcsa recognized in 1927 (43, p. 183) that this was a coelurosaur and intended to give it a name, but used one already used by Osborn, namely "Saurornithoides" (91, 1924, p. 3- 7). For this reason, a new name had to be given here" [translated]. Huene's proposed new name was Caudocoelus sauvagei, placed in Coeluridae and "somewhat reminiscent of Elaphrosaurus." Huene is also perhaps the first of several authors to place the specimen in the Kimmeridgian, when it is actually from the Tithonian (Buffetaut and Martin, 1993; as Portlandian). Galton wrote "Lapparent and Lavocat (1955: 801) gave a line drawing of the vertebra after Sauavage (1898) and included it in the section on Elaphrosaurus" and that the specimen "was referred to Elaphrosaurusby Lapparent and Lavocat (1955)." This was perhaps done because Huene explicitly compared the two, ironically making it the only taxon distinguished from Teinurosaurus at the time. Most of Huene's characters cannot be checked in the few published photos of Teinurosaurus, but the ventral median sulcus is indeed much wider than Elaphrosaurus. Ostrom (1969) was the first author to detail Nopcsa's (1929) addendum, stating "Nopcsa's name Teinurosaurus has clear piority over Huene's Caudocoelus, but since Nopcsa failed to provbide a specific name, Teinurosaurus is not valid." Olshevsky (1978) solved this by writing "Teinurosaurus has clear priority over Caudocoelus, as noted in Ostrom 1969, and it is certainly a valid generic name. The species Caudocoelus sauvagei is proposed here as the type species of the genus Teinurosaurus, resulting in the new combination Teinurosaurus sauvagei(von Huene 1932) as the proper name of the type specimen." He also claimed "the specimen itself, unfortunately, was destroyed during World War II and thus must remain a nomen dubium." This was repeated by Galton, but as Buffetaut et al. (1991) wrote- "Contrary to a widespread opinion (expressed, for instance, by Lapparent, 1967), the vertebra in question has survived two world wars and years of neglect, like a large part of the other fossil reptile remains in the collections of the Boulogne Natural History Museum (see Vadet and Rose, 1986)." Olshevsky noted Steel misunderstood Nopsca in a different way, believing Teinurosaurus instead of Aublysodon was a "name, proposed by Cope in 1869 ... used instead of Deinodon", as stated under superscript 2. Galton did have the first modern opinion on Teinurosaurus' affinities, stating "In addition to Elaphrosaurus, elongate prezygapophyses occur in the allosaurid Allosaurus and the dromaeosaurid Deinonychus, so this caudal vertebra can only be identified as theropod, family incertae sedis." Buffetaut and Martin (1993) agreed, saying "no really distinctive characters that would allow a familial assignment can be observed." Ford (2005 online) gave the type repository as "Dortigen Museum", but this is a misunderstanding based on Huene's "Boulogne-sur-mer (Nr. 500 im dortigen Museum)", which roughly translated is "Boulogne-sur-mer region (No. 500 in the museum there)", referring to the Boulogne Museum where it has always been held. It was originally number 500, but was recatalogued at some point.
Sauvage lists the vertebra's length as 75 mm and his plate at natural size would have it be 79 mm, Huene lists it as 11 cm (110 mm) and his figure at 1:2 size would have it be 152 mm. Galton's drawing with supposed 5 cm scale would have it be 235 mm, while Buffetaut and Martin's plate with scale would leave it at 74 mm. As Huene's and Galton's figures are taken from Sauvage's original plate and the newest and unique photo matches Sauvage's reported length almost exactly, 75 mm is taken as the correct length.
Relationships- While prior authors haven't specified Teinurosaurus' relationships past Theropoda (besides Lapparent and Lavocat's apparent synonymy with Elaphrosaurus), there are several ways to narrow down its identity. Only neotheropods are known from the Late Jurassic onward, so coelophysoid-grade taxa are excluded. Some theropod clades were too small to have a 75 mm caudal, including most non-tyrannosauroid coelurosaurs besides ornithomimosaurs, therizinosaurs and eudromaeosaurs. The former two are unknown from the Jurassic, and additionally paravians like eudromaeosaurs lack any neural spine by the time the centrum gets as elongate as Teinurosaurus (e.g. by caudal 12 in Deinonychus at elongation index of 2.4). Teinurosaurus has an elongation index (centrum length/height) of 3.9, which also excludes Ceratosauridae, Beipiaosaurus+ therizinosauroids and oviraptorosaurs. Prezygapophyses basal depth is significantly less in ceratosaurids, megalosaurids, carnosaurs except Neovenator, compsognathids, Fukuivenator and Falcarius. Remaining taxa are elaphrosaur-grade ceratosaurs, piatnitzkysaurids, Neovenator and basal tyrannosauroids.
References- Sauvage, 1897. Notes sur les Reptiles Fossiles (1). Bulletin de la Société géologique de France. 3(25), 864-875.
Sauvage, 1897-1898. Vertebres Fossiles du Portugual, Contributions a l'etude des poissions et des reptiles du Jurassique et du Cretaceous. Direction des Travaux Geologiques Portugal. 1-46.
Osborn, 1924. Three new Theropoda, Protoceratops zone, central Mongolia. American Museum Novitates. 144, 1-12.
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Nopcsa, 1929. Addendum "The genera of reptiles". Palaeobiologica. 2, 201.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1), 361 pp.
Lapparent and Lavocat, 1955. Dinosauriens. In Piveteau (ed.). Traite de Paleontologie. Masson et Cie. 5, 785-962.
Lapparent, 1967. Les dinosaures de France. Sciences. 51, 4-19.
Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Peabody Museum of Natural History Bulletin. 30, 1-165.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia of Paleoherpetology. Gustav Fischer Verlag. 87 pp.
Olshevsky, 1978. The archosaurian taxa (excluding the Crocodylia). Mesozoic Meanderings. 1, 50 pp.
Galton, 1982. Elaphrosaurus, an ornithomimid dinosaur from the Upper Jurassic of North America and Africa. Paläontologische Zeitschrift. 56, 265-275.
Vadet and Rose, 1986. Catalogue commente des types et figures de dinosauriens, ichthyosauriens, sauropterygiens, pterosauriens et cheloninens du Musée d'Histoire Naturelle de Boulogne-sur-Mer. In E. Buffetaut, Rose and Vadet (eds.). Vértébrés Fossiles du Boulonnais. Mémoires de la Société Académique du Boulonnais. 1(2), 85-97.
Rose, 1987. Redecouverte d'une vertebre caudale reptilienne (Archosauriens) de status controverse et provenant des terrains jurassiques superieurs du Boulonnais. Bulletin de la Société académique du Boulonnais. 1(5), 150-153.
Buffetaut, Cuny and le Loeuff, 1991. French Dinosaurs: The best record in Europe? Modern Geology. 16(1-2), 17-42.
Buffetaut and Martin, 1993. Late Jurassic dinosaurs from the Boulonnais (northern France): A review. Revue de Paléobiologie. 7(vol. spéc.), 17-28.
Ford, 2005 online. http://www.paleofile.com/Dinosaurs/Theropods/Teinurosaurus.asp
And before we go, here are a couple more tidbits I've noticed in the upcoming update...
- That theropod tail preserved in Burmese amber (DIP-V-15103) described by Xing et al. (2016) was only placed as specifically as a non-pygostylian maniraptoriform. But as the deposits are Gondwanan (e.g. Poinar, 2018), the range of potential Cenomanian theropods is better understood. And only one group has caudal centra over three times longer than tall- unenlagiines. I bet DIP-V-15103 is our first sample of preserved plumage in an unenlagiine, which makes you wonder if the weird alternating barb placement was a feature that evolved on Gondwana, and if so did Rahonavis' remiges exhibit it too?
- Does anyone realize both "Tralkasaurus" (Cerroni et al., 2019) and "Thanos" (Delcourt and Iori, 2018) are nomina nuda? Neither are in an official volume yet, though "Tralkasaurus" is scheduled for March and "Thanos" will probably make it this year if the average papers per volume of Historical Biology holds up. "Tralkasaurus" has an empty space in its "Zoobank registration:" section, while the "Thanos" paper doesn't mention ZooBank at all, and neither show up in ZooBank searches. Also, one of "Thanos"' supposed autapomorphies is a deep prezygapophyseal spinodiapophyseal fossa, which does not exist in abelisaurs as it would require a spinodiapophyseal lamina. The labeled structure seems internal, probably the centroprezygapophyseal fossa or prezygapophyseal centrodiapophyseal fossa based on CT-scanned noasaurid cervical DGM929-R. That leaves axial pleurocoel size and distance from each other, and ventral keel strength as suggested characters. Which can only be compared to Carnotaurus among brachyrostrans. Hmmm...
References- Xing, McKellar, Xu, Li, Bai, Persons IV, Miyashita, Benton, Zhang, Wolfe, Yi, Tseng, Ran and Currie, 2016. A feathered dinosaur tail with primitive plumage trapped in Mid-Cretaceous amber. Current Biology. 26(24), 3352-3360.
Delcourt and Iori, 2018. A new Abelisauridae (Dinosauria: Theropoda) from São José do Rio Preto Formation, Upper Cretaceous of Brazil and comments on the Bauru Group fauna. Historical Biology. DOI: 10.1080/08912963.2018.1546700
Poinar, 2018. Burmese amber: Evidence of Gondwanan origin and Cretaceous dispersion. Historical Biology. DOI: 10.1080/08912963.2018.1446531
Cerroni, Motta, Agnolín, Aranciaga Rolando, Brissón Egli and Novas, 2019. A new abelisaurid from the Huincul Formation (Cenomanian-Turonian; Upper Cretaceous) of Río Negro province, Argentina. Journal of South American Earth Sciences. 98, 102445.
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Oculudentavis is not a theropod
Hi all. This week we got the announcement of a tiny theropod skull in Myanmar amber, which was bound to happen eventually as amazing finds from that deposit keep being published. Alas, whatever Oculudentavis is, it's not a theropod.
Oculudentavis skull (after Xing et al., 2020).
Just look at it. No antorbital fenestra, incomplete ventral bar to the laterotemporal fenestra, huge posttemporal fenestrae, teeth that extend posteriorly far under the orbit...
All of which might be coincidental, but then look at the mandible.
Oculudentavis mandible (after Xing et al., 2020).
That spike-like coronoid process is classic lepidosaur, plus the dentary is way too long compared to the post-dentary elements, then the description says "The tooth geometry appears to be acrodont to pleurodont; no grooves or sockets are discernable." And of course "the scleral ring is very large and is formed by elongated spoon-shaped ossicles; a morphology similar to this is otherwise known only in lizards (for example, Lacerta viridis)."
Add to this the size of this partially fused specimen being smaller than any extant bird (14 mm), and no feather remains, and why is this a theropod again? The endocast is big, but why not a clade of brainier lizards or late surviving megalancosaurs by the Cenomanian?
The authors add it to Jingmai's bird analysis where it ends in a huge polytomy closer to Aves than Archaeopteryx, but outside fake Ornithuromorpha. That's often what happens when a taxon is wrongly placed in a clade. Note the figured placement between Archaeopteryx and Jeholornis is only found using implied weights. At least add it to e.g. Nesbitt's or Ezcurra's archosauromorph analyses, or Cau's theropod analyses before assuming it's a bird.
Thanks to Ruben Molina Perez for suggesting this issue in the first place.
Reference- Xing, O'Connor, Schmitz, Chiappe, McKellar, Yi and Li, 2020. Hummingbird-sized dinosaur from the Cretaceous period of Myanmar. Nature. 579, 245-249.
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What is Oculudentavis if it's not a theropod?
In my last post, I argued the recently described Oculudentavis (Xing et al., 2020) is not a theropod. So what is it? To answer that question, I entered it into Simoes et al.'s (2018) sauropsid analysis which emphasizes basal lepidosauromorphs and comes out with basal gekkos and nested iguanians even using just morphological characters. To test Jingmai's avialan hypothesis, I also added Archaeopteryx to the matrix. The result is 384 MPTs of 2337 steps each.
Strict consensus of 384 MPTs of Simoes et al.'s (2018) analysis after adding Oculudentavis and Archaeopteryx. Compare to Extended Data Figure 3 of Simoes et al..
As you can see, Oculudentavis resolves as a stem-squamate in a trichotomy with Huehuecuetzpalli and squamates, while Archaeopteryx is an archosauromorph sister to Erythrosuchus. And this matrix didn't score for scleral ossicle shape, posttemporal fenestra size or maxillary tooth row length. After scoring Oculudentavis, its teeth are clearly not acrodont, it seems to have a ventral parietal fossa and lacks an ossified laterosphenoid. The authors could have made it easier to evaluate by separating the cranial elements in the 3D pdf file. As it is, a lot of palatal and braincase info is uncertain. But Huehuecuetzpalli is Albian compared to Oculudentavis' Cenomanian, and has a skull length of 32 mm (19 mm in the juvenile) versus 14 mm in Oculudentavis.
Huehuecuetzpalli skull (top; after Reynoso, 1998), Oculudentavis skull and separate mandible (middle; after Xing et al., 2020), and Archaeopteryx skull (after Rauhut, 2014).
References- Reynoso, 1998. Huehuecuetzpalli mixtecus gen. et sp. nov: A basal squamate (Reptilia) from the Early Cretaceous of Tepexi de Rodriguez, central Mexico. Philosophical Transactions of the Royal Society B: Biological Sciences. 353, 477-500.
Rauhut, 2014. New observations on the skull of Archaeopteryx. Paläontologische Zeitschrift. 88(2), 211-221.
Simōes, Caldwell, Talanda, Bernardi, Palci, Vernygora, Bernardini, Mancini and Nydam, 2018. The origin of squamates revealed by a Middle Triassic lizard from the Italian Alps. Nature. 557(7707), 706-709.
Xing, O'Connor, Schmitz, Chiappe, McKellar, Yi and Li, 2020. Hummingbird-sized dinosaur from the Cretaceous period of Myanmar. Nature. 579, 245-249.
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It's finally January 1, 200n and Phylonyms is published!
Ah the PhyloCode, the so-called future of biological nomenclature whose release has always kept on slipping ever more distantly into the future. After 20 years of waiting, we now have Phylonyms: A Companion to the PhyloCode, by de Queiroz et al. (2020), "a turning point in the history of phylogenetic nomenclature" according to its introduction. As the book states "Phylonyms serves as the starting point for phylogenetic nomenclature governed by the PhyloCode. According to the preamble, “This code will take effect on the publication of Phylonyms: A Companion to the PhyloCode, and it is not retroactive.” Thus, names and definitions published here have precedence over any competing names and definitions published either before (or after) the publication of Phylonyms." So for anyone invested in standardized phylogenetic nomenclature, this is it. Nothing better is coming down the pipeline in our lifetimes, so let's see what we're stuck with.
First of all, it's expensive. You can get an ebook for $222 on Amazon or a hardcover sometime after June 9th for $234. I found an electronic version for $169 plus tax on VitalSource, but you have to use their reader. It's 1323 pages though, so isn't a bad deal. That's less than five $37.95 Cretaceous Research pdfs, and I figure this is one of those historical volumes it's good to have, like Sibley and Ahlquist's bird phylogeny book.
The format is an encyclopedia-style list of clades in phylogenetic order with Registration Number, Definition, Etymology, Reference Phylogeny, Composition, Apomorphies, Synonyms, Comments and Literature Cited. Rather like my Theropod Database, so no complaints there. Well, one complaint is really more to do with the PhyloCode itself where they decided to abbreviate definitions with the non-standard del/nabla triangle symbol ∇. If you want people to start using your format, you might want to choose symbols that exist on a standard keyboard. Alt+2207 is supposed to generate it in Windows, but results in ƒ here in Blogger. Anyone know the correct Unicode numbers?
On to the substance, where Phylonyms covers all life. Dinosaurs are the last section of the book, and non-avian dinosaurs get all of four definitions-
Dinosauria R. Owen 1842 [M. C. Langer, F. E. Novas, J. S. Bittencourt, M. D. Ezcurra, and J. A. Gauthier], converted clade name
Registration Number: 194
Definition: The smallest clade containing Iguanodon bernissartensis Boulenger in Beneden 1881 (Ornithischia/Euornithopoda) Megalosaurus bucklandii Mantell 1827 (Theropoda/Megalosauroidea) and Cetiosaurus oxoniensis Phillips 1871 (Sauropodomorpha).
I'm glad we've standardized which theropod, ornithischian and sauropodomorph are used (or so I thought, see below), but otherwise there's not much to say. The caveats around which apomorphies are also found in Nyasasaurus and at least some silesaurs illustrate why apomorphy-based definitions are bad. The reference phylogeny for this and Saurischia is Lloyd et al.'s (2008) supertree, which is quite outdated and has a lot of artifacts from being a supertree.
Saurischia H. G. Seeley 1888 [J. A. Gauthier, M. C. Langer, F. E. Novas, J. Bittencourt, and M. D. Ezcurra], converted clade name
Registration Number: 195
Definition: The largest clade containing Allosaurus fragilis Marsh 1877 (Theropoda/Carnosauria) and Camarasaurus supremus Cope 1877 (Sauropodomorpha), but not Stegosaurus stenops Marsh 1887 (Ornithischia/Stegosauridae).
It's rather odd the same authors didn't choose the same specifiers for each dinosaurian clade as they did in the previous definition, leaving us without a neat node-stem triplet. Instead they went with the Kischlatian approach of using taxa"mentioned and figured as examples of their respective groups by Seeley (1888)." This is funny because I don't think this rationale is ever suggested in the PhyloCode, whereas Dinosauria and Saurischia are actually the official examples used for Recommendation 11F encouraging node-stem triplets ("If it is important to establish two names as applying to sister clades regardless of the phylogeny, reciprocal maximum-clade definitions should be used in which the single internal specifier of one is the single external specifier of the other, and vice versa"). Specifically- "If one wishes to define the names Saurischia and Ornithischia such that they will always refer to sister clades, Saurischia might be defined as the largest clade containing Megalosaurus bucklandiiMantell 1827 but not Iguanodon bernissartensisBoulenger in Beneden 1881, and Ornithischia would be defined as the largest clade containing Iguanodon bernissartensis but not Megalosaurus bucklandii. To stabilize the name Dinosauria as referring to the clade comprising Saurischia and Ornithischia, Dinosauria should be defined as the smallest clade containing Megalosaurus bucklandii and Iguanodon bernissartensis."
Ornithoscelida and its consequences are mentioned, but I'm glad more time is not taken up with it as I expect the hypothesis to fall away as Baron's phylogenetic mistakes are not followed by future authors.
Sauropodomorpha F. R. von Huene 1932 [M. Fabbri, E. Tschopp, B. McPhee, S. Nesbitt, D. Pol, and M. Langer], converted clade name
Registration Number: 295
Definition: The largest clade containing Saltasaurus loricatus Bonaparte and Powell 1980 (Sauropodomorpha) but not Allosaurus fragilis Marsh 1877 (Theropoda) and Iguanodon bernissartensis Boulenger in Beneden 1881 (Ornithischia).
I dislike the use of Saltasaurus as the internal specifier, which is a holdover of Sereno's weird use of deeply nested OTUs when others would be more historically relevant and/or eponymous. Fabbri et al. defend the choice because "Fossil specimens referred to Saltasaurus loricatus are abundant, the species is well known, and its phylogenetic position is consistent among phylogenetic analyses", but this would be even more true for e.g. Camarasaurus supremus used in Saurischia's definition. The other specifiers are a mix of those in Dinosauria's and Saurischia's definition, so there's absolutely no consistency. The reference phylogeny is Otero et al.'s (2015) Sefapanosaurus description using Yates' matrix, so is fine.
There's a rare error in the comments for this entry. Fabbri et al. state "Segnosaurus galbinensis from the Cretaceous was briefly thought to be a relatively early diverging sauropodomorph (Paul, 1984; Gauthier, 1986; Olshevsky, 1991). More material referable to that species and the discovery of closely related taxa later showed that Segnosaurus galbinensis is part of the Therizinosauria", but material of S. galbinensis besides that initially recovered in the 1970s is not known.
Theropoda O. C. Marsh 1881 [D. Naish, A. Cau, T. R. Holtz, Jr., M. Fabbri, and J. A. Gauthier], converted clade name
Registration Number: 216
Definition: The largest clade containing Allosaurus fragilis Marsh 1877 (Theropoda) but neither Plateosaurus engelhardti Meyer 1837 (Sauropodomorpha) nor Heterodontosaurus tucki Crompton and Charig 1962 (Ornithischia).
Here we've chosen two completely different specifiers for Sauropodomorpha and Ornithischia, so again we have no consistency. The reference phylogeny is Cau (2018), which is ideal.
What about the rest? It's a HUGE volume, and obviously most of Pan-Biota is outside my area of expertise. One obvious issue is the wildly varying coverage of different clades. Apparently nobody could be bothered with the vast majority of vertebrates (euteleosts) or animals (insects, except one definition for Trichoptera), and Molluska doesn't even get a definition. But we do get several entries for edrioasterid taxa down to subfamily-level, generally obscure Paleozoic echinoderms. Closer to dinosaurs, there's nothing at all for pan-crocs, but we get an entry for Pterosauromorpha for which only Scleromochlus is given as a plausible non-pterosaurian example (perhaps wrongly- Bennett, 2020).
Then there are the apomorphy-based definitions which will cause headaches in the future. Look at Apo-Chiroptera- "Definition: The clade for which the unique modifications of the hand, forearm, humerus, scapula, hip, and ankle (see Diagnostic Apomorphies) associated with flapping flight, as inherited by Vespertilio murinus Linnaeus 1758, are apomorphies." Then you go down to the nine listed sets of Diagnostic Apomorphies like "Modification of the scapula: Scapular spine originates at the posterior edge of the glenoid fossa. Long axis of scapular spine offset 20–30 degrees from axis of rotation of the humeral head. Scapular spine reduced in height—acromion process appears more strongly arched and less well supported than in other mammals. Presence of at least two facets in infraspinous fossa." These are all going spread out as more stem bats are discovered, and indeed the authors already note "Simmons and Geisler (1998) included the absence of claws on wing digits III-V with this suite of modifications; however, the presence of claws on all the wing digits of Onychonycteris suggests that claws were present primitively in Apo-Chiroptera."
Ungulata is defined by Archibald as "The least inclusive crown clade containing Bos primigenius Bojanus 1827 (= Bos taurus Linnaeus 1758) (Artiodactyla) and Equus ferus Boddaert 1785 (= Equus caballus Linnaeus 1758) (Perissodactyla), provided that this clade does not include Felis silvestris Schreber 1777 (= Felis catus Linnaeus 1758) (Carnivora), Manis pentadactyla Linnaeus 1758 (Pholidota), Vespertilio murinus Linnaeus 1758 (Chiroptera), or Erinaceus europaeus Linnaeus 1758 (Lipotyphla)." But this doesn't exist in molecular studies, including those of ultraconserved elements, which consistently place carnivorans, pangolins and bats closer to perrisodactyls. So this is likely to be a historical footnote, as well established molecular relationships end up trumping morphological relationships in every example I know of.
Finally, we get Pan-Lepidosauria for the total group of lepidosaurs, which has been Lepidosauromorpha for over thirty years. Yet Archosauromorpha is retained as "The least inclusive clade containing Gallus (originally Phasianus) gallus (Aves) (Linnaeus 1758), Alligator (originally Crocdilus) mississippiensis (Daudin 1802) (Crocodylia), Mesosuchus browni Watson 1912 (Rhynchosauria), Trilophosaurus buettneri Case 1928 (Trilophosauridae), Prolacerta broomi Parrington 1935 (Prolacertiformes), and Protorosaurus speneri von Meyer 1830 (Protorosauria)" even though Pan-Archosauria is also used for the total group of archosaurs, traditionally the definition of Archosauromorpha. I agree our new Archosauromorpha deserved a name for being a generally recognized group, whereas whether e.g. choristoderes or sauropterygians fell out closer to lizards or birds is highly unstable. But I would have rather kept the tradition of -omorpha for the stem clades and gave this a new name.
Overall, I'm not very impressed for something 20 years in the making that intends to be so important. How do you contradict your own example for choosing specifiers in four papers, where two share the same author list, the other two share another author (Fabbri), and each of those shares an author with both of the first two (Langer and Gauthier)? And one of those is an editor for the volume. Nothing could be negotiated in over two decades? But it's what we have to work with now, and in the name of consistancy I'll adopt the definitions proposed. Now to see what happens when RegNum goes online.
References- Lloyd, Davis, Pisani, Tarver, Ruta, Sakamoto, Hone, Jennings and Benton, 2008. Dinosaurs and the Cretaceous terrestrial revolution. Proceedings of the Royal Society B. 275, 2483-2490.
Otero, Krupandan, Pol, Chinsamy and Choiniere, 2015. A new basal sauropodiform from South Africa and the phylogenetic relationships of basal sauropodomorphs. Zoological Journal of the Linnean Society. 174, 589-634.
Cau, 2018. The assembly of the avian body plan: A 160-million-year long process. Bollettino della Società Paleontologica Italiana. 57(1), 1-25.
Bennett, 2020. Reassessment of the Triassic archosauriform Scleromochlus taylori: Neither runner nor biped, but hopper. PeerJ. 8:e8418.
de Queiroz, Cantiono and Gauthier, 2020. Phylonyms: A Companion to the PhyloCode, 1st Edition. Taylor & Francis Group. 1323 pp.
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The Unecessary Death of Steneosaurus
Not a dinosaur, but a new paper on the classic crocodylomorph Steneosaurus exemplifies a troubling trend in recent vertebrate taxonomy. Johnson et al. (2020) reexamine the original material of Steneosaurus, an aquatic croc from the Jurassic of France. It hadn't been seriously looked at since the 1860s, so this is one of my favorite kinds of paleontology papers- restudying a fragmentary old specimen in a modern light. What do they find?
We first get a detailed recount of its history, with two decades as Cuvier's "tête à museau plus allongé" (= head with a more elongated snout; I have to praise the authors for translating all the French to English, even in our spoiled era of Google Translate it saves time), before it was named Steneosaurus rostro-major by Geoffroy Saint-Hilaire in 1825. Eudes Deslongchamps and son tackled it in the 1860s, where they viewed the specimen as too poorly preserved and so "stated that the taxon to represent the genus Steneosaurus should be either ‘Steneosaurus’ megistorhynchus Eudes-Deslongchamps, 1866, or ‘Steneosaurus’ edwardsi Eudes-Deslongchamps, 1868c." Ha! You don't get to just take somebody's genus and affix your new species as its type. They were the last to examine the specimen in detail however, making that a pretty bad note to end on.
Johnson et al. then reexamine the type snout of Steneosaurus, correcting the species name by eliminating the hyphen, officially making it the lectotype, noting Steel had determined the posterior skull to be Metriorhynchus, and illustrating and redescribing the specimen. Excellent work and very well done. After eliminating Mycterosuchus nasutus, 'Steneosaurus' leedsi, 'S.' heberti and Lemmysuchus and other machimosaurins based on numerous dissimilar characters, the authors come to the contemporaneous 'Steneosaurus' edwardsi.
"As mentioned before, this was a second species that Eudes-Deslongchamps (1867–69) considered identical to S. rostromajor. These two taxa share a combination of features including:
1. A subcircular, moderately interdigitating premaxilla-maxilla suture.
2. Maxillae ornamented with irregular grooves.
3. A shallower mediolateral compression of the posterior maxillae, as opposed to ‘S.’ heberti (MNHN.F 1890-13).
4. Horizontally flat posterior premaxilla in lateral view.
5. Deep anterior and mid-maxillary reception pits that gradually become shallower towards the posterior maxilla.
6. Subcircular to circular alveoli that remain relatively the same size throughout the maxilla.
7. Teeth with well-pronounced enamel ridges at the base."
Well how cool is that? They put in the hard work, found the matching more complete specimens, and now we have Steneosaurus edwardsi as a junior synonym of S. rostromajor, giving us a good look at what Steneosaurus really was after two hundred years.
Lectotype of Steneosaurus rostromajor (MNHN.RJN 134c-d) in dorsal (A, B) and ventral (C, D) views. (after Johnson et al., 2020).
But no.
Johnson et al. immediately say "it is important to note that many of these characters may, in fact, be related to sexual dimorphism, ontogeny and intraspecific variation." True, but that could be said for basically every character supposed to diagnose Mesozoic croc genera, or theropod genera, pterosaur genera, etc.. Unless you have some specific example like 'enamel ridges have been shown to develop with age and both S. rostromajor and S. edwardsi are larger than S? leedsi or S? heberti with weak ridges', then it's just hand-waving. And no, Johnson et al. never develop such an argument for one of those characters, let alone all seven.
Next, we get "In addition to the sexual dimorphism/ontogeny problem, one of the critical issues about MNHN.RJN 134c-d is that it is poorly preserved." Sure, but you were still able to perform many comparisons. Again, the authors never say any of their seven characters are taphonomic, so it's another objection without substance.
Yet the worst rationale for rejecting Steneosaurus is "in reality, the name Steneosaurus is extremely impractical. It was used for many metriorhynchid specimens (e.g. ‘Steneosaurus’ gracilis, ‘Steneosaurus’ palpebrosus and ‘Steneosaurus’ manselii) during much of the 19th century, largely in part due to Cuvier’s metriorhynchid skull region (MNHN. RJN 134a-b) being attributed to the teleosauroid rostral section (MNHN.RJN 134c-d). Indeed, the concise, classical definition of ‘Steneosaurus’ as we interpret it today was not given until the work of both Eudes-Deslongchampses (1868c, 1867–69)"
Substitute Megalosaurus in there to see how ridiculous it is. That has had over 45 species assigned to it, and was named in the 1820s but didn't have a modern concept associated with it until the 1980s. When Johnson et al. lament that for Steneosaurus "rather than comparing characters outright, comparison is by process of elimination (or the question of ‘what features does this specimen lack?’)", that perfectly describes the Megalosaurus paralectotype dentary.
"After the Eudes-Deslongchampses’ treatment, what was left was an undiagnostic, chimeric type specimen for S. rostromajor (MNHN.RJN 134) and the genus Steneosaurus was redefined using a new type species that was not accepted by some researchers. In addition, since the Eudes-Deslongchampses, there has been no attempt to rectify this taxonomic nightmare;"
You just showed it was diagnostic, Steel long ago got rid of the chimaeric portion, Eudes-Deslongchamps' stupid attempts to name new type species have no relevance, and you have done the work to finally rectify this taxonomic nightmare.
"Due to these three significant factors (uncertainty of variable characters, poor preservation and
unreasonable name), we have concluded that S. rostromajor, and therefore ‘Steneosaurus’ (MNHN.RJN 134c-d), cannot be confidently assigned to an existing teleosauroid species."
Nope, you just showed it can be assigned to the same species as S. edwardsi.
Actually, I correct myself. THIS is the worst rationale for rejecting Steneosaurus- "In addition, MNHN.RJN 134c-d was initially diagnosed based on significant orbital and temporal characteristics (from the metriorhynchid MNHN.RJN 134a-b), along with generic rostral ones. Because the skull material is now known to be from a metriorhynchid, this ‘hybrid type specimen’ factor adds to the doubtful validity of Steneosaurus. According to Article 23.8 of the ICZN Code, ‘a species-group name established for an animal later found to be a hybrid (Art. 17) must not be used as the valid name for either of the parental species (even if it is older than all other available names for them)’ (this also signifies that the species name rostromajor is itself invalid). As such, MNHN.RJN 134c-d serves as an undiagnostic specimen; we, therefore, consider MNHN.RJN 134c-d to be a nomen dubium and, as such, Steneosaurus is treated as an undiagnostic genus."
If the term "parental species" didn't tip you off, Article 23.8 applies to hybrid individuals (those resulting from different species interbreeding), not type specimens chimaerically combined from multiple species. The Article doesn't even say what Johnson et al. think- it says a name for a hybrid can't be used for either of the species that bred to make it, so that e.g. even if a mule's scientific name was erected prior to that of horse's or ass's, it can't be the name for horse or ass. And indeed even the cited Article 17 says that hybrids and chimaeras can be the basis of valid names- "The availability of a name is not affected even if 17.1. it is found that the original description or name-bearing type specimen(s) relates to more than one taxon, or to parts of animals belonging to more than one taxon; or 17.2. it is applied to a taxon known, or later found, to be of hybrid origin..."
If Johnson et al.'s interpretation were right, there goes Gojirasaurus, Protoavis, Chuandongocoelurus, Chilantaisaurus, Fukuiraptor, Coelurus, Alectrosaurus, Dakotaraptor, etc..
Before the big reveal, we have in the Conclusion what can only be described as a lie- "Through character comparison-and-elimination, the only taxon with which MHNH.RJN 134c-d could hypothetically be referred to is ‘S.’ edwardsi, but the two do not share any clear autapomorphic characters or a unique combination of characters."
What are your seven listed characters if not "a unique combination of characters"? Does any other teleosaurid have them? If not, they are unique. In any case, we get the motivation for dumping Steneosaurus twice at the end of the paper-
"We believe that establishing teleosauroid taxonomy from the beginning with a series of ‘clean’ type species/specimens, with every nomenclatural act correctly formulated, is the best course of action, which we will highlight in a forthcoming paper (Johnson, 2019)."
"We believe that establishing teleosauroid taxonomy from the beginning with a series of ‘clean’ type species/specimens, with every nomenclatural act correctly formulated, is the best course of action. This will necessitate a revised teleosauroid taxonomy, in which species previously referred to the genus Steneosaurus are given new generic names. This work will be published by us in a separate contribution, based on the comprehensive teleosauroid phylogenetic analysis in Johnson’s PhD thesis (2019)."
Basically everything I hate about a current trend in vertebrate paleontology- just throw out old specimens and dishonor their authors who correctly reported what was new at the time to come up with your own names. At least dumping Stegosaurus armatus or designating a neotype for Allosaurus fragilis could be claimed to save time and effort actually analysing the types, if you don't want to do the science to figure out if armatus is actually different from stenops or if fragilis can be distinguished from Saurophaganax. But Johnson et al. already did all the hard work and found Steneosaurus edwardsi was S. rostromajor, they would just rather use Johnson's new genus name for the taxon.
And their reasons are just grasping at straws. 'Sure we identified these seven charactesrs uniquely shared by Steneosaurus rostromajor and S. edwardsi, but uhh.. could be sexually dimorphic? Or anything could be individual variation. Or ontogenetic? Lots of things turn out to be ontogenetic. Plus it's broken. Sooo broken. Sure we could evaluate characters, but who wants a taxon whose holotype isn't pristine? Plus a lot of people had stupid ideas about Steneosaurus over the past two hundred years. What do us scientists do when we have a complicated situation to resolve that was only partially understood historically? Trash their names and give yourselves credit for new genera.' Thus Steneosaurus gets the eternal identity of "all evidence points to it being Johnsonosaurus edwardsi, but ehhh... we just sort of ignore it now as Teleosauridae indet. and it's forgotten."
To conclude, Steneosaurus is really outside my wheelhouse. But if Johnson et al.'s philosophy spreads, we're in danger of losing a lot of historical taxonomy and deserved credit to lazy or selfish authors. Just look at Microraptor for example, whose holotype of M. zhaoianus lacks a decent skull. Some decades down the line, what if cranial differences support various Jiufotang species and someone's like 'the postcranial proportions are unique between the M. zhaoianus type and M. hanqingi, but I want a complete type specimen, so Microraptor is an invalid undiagnostic nomen dubium, and instead I propose Mybetterraptorgenus hanqingi and M. gui.' Just hope they don't pull a Wilson and Upchurch and claim 'Microraptor is invalid and co-ordinate suprageneric Linnean taxa must likewise be abandoned' and replace Microraptorinae with Mybetterraptorgenusinae.
References- Johnson, 2019. The taxonomy, systematics and ecomorphological diversity of Teleosauroidea (Crocodylomorpha, Thalattosuchia), and the evaluation of the genus 'Steneosaurus'. PhD Thesis, University of Edinburgh. 1062 pp.
Johnson, Young and Brusatte, 2020. Emptying the wastebasket: A historical and taxonomic revision of the Jurassic crocodylomorph Steneosaurus. Zoological Journal of the Linnean Society. 189(2), 428-448.
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The Arguable Identity of Paraxenisaurus
Hi everyone. In light of Cau's recent post on the supposed new Mexican deinocheirid "Paraxenisaurus normalensis" (Serrano-Brañas et al., 2020), I figured I'd check the taxon out to see what I thought.
The first thing you might note are the quotation marks surrounding its name, as this is yet another example of authors not including an lsid or reference to ZooBank in their electronic descriptions. ICZN Article 8.5.3. states names published electronically must "be registered in the Official Register of Zoological Nomenclature (ZooBank) (see Article 78.2.4) and contain evidence in the work itself that such registration has occurred", and the pre-print is said to be in preparation for Volume 101 of Journal of South American Earth Sciences, cited as August 2020. Thus it gets to join the ranks of "Thanos" and "Trierarchuncus"as theropods that will eventually be validly named this year. But at least it's not stuck in the purgatory of twelve Scientific Reports Mesozoic theropods, which will never be physically published and thus will remain invalid unless outside action is taken.
One of the big takeaways from Cau's blogpost is that "I am doubtful about the possibility of referring these elements [the paratypes] to the same species of the holotype, since there are very few superimposable elements among the three specimens. Therefore, there is a risk that Paraxenisaurus , - understood as the sum of all three specimens - is a chimera." After reading the paper, Andrea REALLY undersold this critique. Here are the specimen materials lists, with the overlapping elements highlighted in matching colors-
(BENC 2/2-001; proposed holotype) proximal manual phalanx II-2 or III-3, partial astragalocalcaneum, partial metatarsal II, phalanx II-1 (115 mm), proximal phalanx II-2, partial metatarsal III, proximal phalanx III-3, distal metatarsal IV, phalanx IV-1 (104 mm), phalanx IV-3 (67 mm), phalanx IV-4 (45 mm), partial pedal ungual IV
(BENC 1/2-0054) distal metacarpal I, proximal phalanx I-1, partial manual ungual I, distal metacarpal II, distal phalanx II-2
(BENC 1/2-0091) several proximal caudal central fragments (66, 75, 76 mm), proximal metacarpal II, partial metacarpal III, distal femur (155 mm trans), distal metatarsal IV
(BENC 1/2-0092) several distal caudal vertebrae (70, 71 mm)
(BENC 30/2-001) pedal ungual II, pedal ungual III
As you can see, there's only one strict overlap, with BENC 1/2-0091 sharing a distal metatarsal IV with the proposed holotype, found ~14 kilometers away. The paper lists no proposed apomorphies or unique combination of characters for distal metatarsal IV, and indeed the description states they preserve largely non-overlapping portions-
"In the holotype, the distal articular surface is fragmented (Figures 11a1 and 11a2); but in the referred specimen (BENC ½-0091), this surface is nicely preserved and has a non-ginglymoid outline (Figures 11b1 and 11b2). The medial condyle is mostly preserved in the holotype (Figure 11a3), but in the referred specimen it is completely broken (Figure 11b3). Conversely, the lateral condyle is broken in the holotype (Figure 11a4), but is well preserved in the referred specimen (Figure 11b4). Collateral ligament fossae are well developed on both condyles and have approximately the same size and depth (Figures 11a3 and 11b4). In cross-section, the shaft of metatarsal IV near the distal end is thicker dorsoventrally than wide."
Needless to say, metatarsal IV has a shaft which is deeper than wide in all ornithomimosaurs, and the preserved ligament fossae are on opposite sides in each specimen (medial in proposed holotype, lateral in 1/2-0091). Below is a figure comparing the two Mexican specimens with Ornithomimus velox, with 1/2-0091 flipped so that all are comparable as left elements. I don't see anything the "Paraxenisaurus" specimens have in common that could diagnose a taxon.
Left distal metatarsal IV of (left to right) intended "Paraxenisaurus normalensis" holotype BENC 2/2-001, intended "Paraxenisaurus normalensis" paratype BENC 1/2-0091 (right element flipped), and Ornithomomus velox holotype YPM 542 in (top to bottom) dorsal, lateral, ventral and medial views ("Paraxenisaurus" after Serrano-Brañas et al., 2020; Ornithomimus after Claessens and Loewen, 2016).
While no other elements are exactly matched, referred specimen BENC 30/2-001 does include pedal unguals II and III, while the intended holotype has pedal ungual IV. These are again from different localities, although closer this time (~2.8 km), and this time we have characters listed in the diagnosis-
"(9) distinctively broad and ventrally curved pedal unguals that angled downward with respect to the proximal articular surface and depending on the digit, the proximodorsal process becomes slightly enlarge and changes its position from nearly horizontal to mostly vertical, adopting a lipshaped appearance; and (10) pedal unguals with a rounded, large foramen on the medial side* and a deep ventral fossa that surrounds a strongly developed, ridge-like flexor tubercle."
Pedal unguals of (left to right) intended "Paraxenisaurus normalensis" paratype BENC30/2-001 right digit II, right digit III and intended "Paraxenisaurus normalensis" holotype BENC 2/2-001 left digit IV in (top to bottom) right, left, proximal and dorsal views (after Serrano-Brañas et al., 2020). Green lines point to supposedly natural median foramen
Ventral curvature is plesiomorphic, the unguals of BENC 30/2-001 are not broader than other ornithomimosaurs', and ventral angling with the proximal end held vertically is common in theropods and present in e.g. Garudimimus and Beishanlong. The proximodorsal process "changing its position" is using a difference between 30/2-001's mostly horizontal processes and the intended holotype's more vertical process as character, which in itself presupposes they are the same taxon. The ventral fossa surrounding a ridge-like flexor tubercle is also present in Harpymimus, Garudimimus, Beishanlong and large Dinosaur Park unguals (NMC 1349, RTMP 1967.19.145) and is not shown in the intended holotype but is claimed to be "partially broken." This leaves the medial foramen, which might be a valid character in unguals III and IV (II is damaged in that area), but might also be taphonomic, as there are many other small circular areas of damage (e.g. center of proximal surface of ungual IV). While the two unguals in 30/2-001 are similar to each other, that of the intended holotype is more strongly curved, has that smaller more dorsally angled proximodorsal process, is wider in proximal view, and lacks the expanded ventral half characteristic of ornithomimosaurs that is present in the other specimen. But even if these two pedal unguals are correctly referred, they are all that's present in specimen BENC 30/2-001. So they get us nowhere in determining caudal, manual (besides proximal manual phalanx II-2 or III-3) or femoral morphology.
The final issue I noticed was the emphasis on "Paraxenisaurus" having a first pedal digit. This would ironically be unlike Deinocheirus, but plesiomorphically shared with Nedcolbertia, "Grusimimus", Garudimimus, Beishanlong, Archaeornithomimus and Sinornithomimus. The character state is based on metatarsal II, where "a facet on the posterior surface of the distal quarter of this shaft, indicates the presence of an articulation area for metatarsal I." The figure shows a longitudinal groove extending down the posterior center of distal metatarsal II, which as anyone who has scored taxa for Clarke's bird matrix could tell you, is not how non-birds attach their hallux to the metatarsus. Hattori (2016) for instance writes in Allosaurus "there is no attachment scar corresponding to the metatarsal I fossa on either medial or plantar aspect of MT II" and in Citipati "there is no obvious attachment scar of MT I on either medial or plantar aspect of MT II." Serrano-Brañas et al. state "in Garudimimus brevipes ... the attachment site is also placed in the same area as in Paraxenisaurus normalensis", but the feature in Garudimimus is a raised scar with sharp medial demarcation from the shaft. As Middleton (2003) recognized, this scar is for the m. gastrocnemius, specifically the m. gastrocnemius pars medialis (Carrano and Hutchinson, 2002), and I'll note it's present even in Gallimimus which lacks pedal digit I (Osmolska et al., 1972: Plate XLIX Fig. 1b). "Paraxenisaurus"'s groove is then more likely to be the m. flexor digitorum longus II tendon, which "passed through the ventral groove in its respective metatarsal to insert serially on each of the pedal phalanges" in e.g. Tyrannosaurus (Carrano and Hutchinson, 2002).
Left metatarsal II in ventral view of (left to right) intended "Paraxenisaurus normalensis" holotype BENC 2/2-001 (after Serrano-Brañas et al., 2020; yellow arrow points to supposed articulation for metatarsal I), Garudimimus brevipes holotype IGM 100/13 (after Kobayashi and Barsbold, 2005; line points to supposed articulation for metatarsal I), Gallimimus bullatus ZPAL MgD-I/94 (after Osmolska et al., 1972), and Tyrannosaurus rex FMNH PR2081 (after Brochu, 2003).
What exactly is "Paraxenisaurus"? Comparison is hindered by the specimens being figured mixed together, and the figures are not in numerical order in the preprint, being shown in the order of- 1, 10-19, 2, 20-23, 3-9. In addition, the scale bars vary within the same figure (e.g. phalanx IV-1 is proximally ~61 mm wide in figure 14a but ~93 mm wide in figure 14e) and the listed measurements are different yet (e.g. IV-1 is listed as 83 mm wide). Thus any composite reconstruction is necessarily approximate. The supposed manual element is too fragmentary to give much information, but it is of the appropriate size and shape to be a proximal pedal phalanx I-1. This would make more sense preservationally since the other material preserved in the specimen is all from the tarsus and pes. It's a shame the astragalocalcaneum is not described better or figured in more views, as the dorsal (= proximal?) perspective has many broken surfaces and edges, so that e.g. the small calcaneum might be preservational. The fused proximal tarsals are like ceratosaurs, deinocheirids (Deinocheirus plus Hexing), alvarezsaurids and caenagnathids. Having any sense of the ascending process morphology could tell us much. Metatarsal II is not obviously deeper than wide, unlike ornithomimosaurs (except Harpymimus; unreported in deinocheirids), but like carcharodontosaurids, therizinosauroids, some oviraptorids and velociraptorines. The proximal outline of metatarsal III would at first glance appear to be the strangest thing about this material, being reconstructed as strictly dorsoventrally oval unlike all(?) other theropods. Tilting it and adding a posterior tapered tip results in a close match to Majungasaurus however (see figure below). If it is an unreduced proximal metatarsal III, tyrannosauroids, most ornithomimosaurs, alvarezsauroids and pennaraptorans would be excluded. Proximal phalanx II-2 lacks a proximoventral heel, so is not from a deinonychosaur. The pedal phalanges are too elongate to be therizinosauroid, and the pedal ungual is too broad. Phalanges are not as dorsoventrally compressed as Mapusaurus, and as noted above they lack the ventrolateral shelves found in ornithomimosaurs. Abelisaurid phalanges seem similar however. I wonder if we have a case like Camarillasaurus or probably Dandakosaurus involving misidentified elements making the specimen seem stranger than it really was, with so many edges of supposed metatarsal III dotted to indicate incompleteness that it could actually be metatarsal II or IV. Certainly nothing connects this specimen with Deinocheirus. As per the numerous errors illustrated by Hartman et al. (2019) nobody should trust Choiniere et al.'s scorings in any case. The Lori matrix recovers "Paraxenisaurus" as a ceratosaur closest to Aucasaurus as far as taxa with well preserved feet are concerned, but also doesn't include characters particular to ceratosaurs and isn't great with pedal characters in general. So I would place the specimen as Neotheropoda incertae sedis (or even indet.) pending a better description of the tarsus and of the real bone surfaces on supposed proximal metatarsal III.
Pes of "Paraxenisaurus normalensis" holotype (center) in dorsal view compared to Majungasaurus crenatissimus composite (left) and Deinocheirus mirificus referred specimen IGM 100/127 (right). Colored proximal view of "Paraxenisaurus" is after Serrano-Brañas et al., with reoriented metatarsal III as per my interpretation shown above that. Note "Paraxenisaurus" elements were scaled using their scale bars, whereas scaling to listed measurements results in different proportions, so those should be seen as approximate. "Paraxenisaurus" after Serrano-Brañas et al. (2020), Majungasaurus after Carrano (2007) and Deinocheirus after Lee et al. (2014).
References- Osmólska, Roniewicz and Barsbold, 1972. A new dinosaur, Gallimimus bullatus n. gen., n. sp. (Ornithomimidae) from the Upper Cretaceous of Mongolia. Palaeontologica Polonica. 27, 103-143.
Carrano and Hutchinson, 2002. Pelvic and hindlimb musculature of Tyrannosaurus rex (Dinosauria: Theropoda). Journal of Morphology. 253, 207-228.
Brochu, 2003. Osteology of Tyrannosaurus rex: Insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Society of Vertebrate Paleontology Memoir. 7, 138 pp.
Middleton, 2003. Morphology, evolution, and function of the avian hallux. PhD thesis, Brown University. 147 pp.
Carrano, 2007. The appendicular skeleton of Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. In Sampson and Krause (eds.). Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. SVP Memoir 8, 164-179.
Lee, Barsbold, Currie, Kobayashi, Lee, Godefroit, Escuillie and Tsogtbaatar, 2014. Resolving the long-standing enigmas of a giant ornithomimosaur Deinocheirus mirificus. Nature. 515, 257-260.
Kobayashi and Barsbold, 2005. Reexamination of a primitive ornithomimosaur, Garudimimus brevipes Barsbold, 1981 (Dinosauria: Theropoda), from the Late Cretaceous of Mongolia. Canadian Journal of Earth Sciences. 42(9), 1501-1521.
Claessens and Loewen, 2016 (online 2015). A redescription of Ornithomimus velox Marsh, 1890 (Dinosauria, Theropoda). Journal of Vertebrate Paleontology. 36(1), e1034593.
Hattori, 2016. Evolution of the hallux in non-avian theropod dinosaurs. Journal of Vertebrate Paleontology. 36(4), e1116995.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247.
Cau, 2020 online. http://theropoda.blogspot.com/2020/04/paraxenisaurus-un-deinocheiride.html
Serrano-Brañas, Espinosa-Chávez, Maccracken, Gutiérrez-Blando, de León-Dávila and Ventura, 2020. Paraxenisaurus normalensis, a large deinocheirid ornithomimosaur from the Cerro del Pueblo Formation (Upper Cretaceous), Coahuila, Mexico. Journal of South American Earth Sciences. 101, 102610.
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Is Falcatakely a bird?
So this week we got the description of the new Maevarano skull Falcatakely forsterae (O'Connor et al., 2020). It's a pseudo-toucan, with a long, tall snout formed mostly by the maxilla unlike modern birds. O'Connor et al. recover it as an enantiornithine using Brusatte et al.'s TWiG analysis and O'Connor's bird analysis. Our problem is that we only have the anteroventral skull preserved, so no braincase, mandible or postcrania. And being a Maastrichtian deposit in Africa, we don't have the most detailed coelurosaur record. Add in the fact beaks are known to evolve fast on islands (as Madagascar was even back then) and we have a potential problem on our hands.
Follow your nose to the exciting topic of African coelurosaur diversity. Reconstructed holotype skull of Falcatakely forsterae (UA 10015) (after O'Connor et al., 2020).
The Lori analysis places Falcatakely in two potential positions- a therizinosaurid or an omnivoropterygid. The former doesn't make much sense biostratigraphically, but there is a Maevarano synsacrum of the correct size (FMNH PA 741) that was claimed to share characters with Sapeornis by O'Connor and Forster (2010). So there's a possibility. Forcing Falcatakely to be an enantiornithine as in O'Connor et al.'s analyses requires six more steps. Most of the discordant characters relate to the beak, but the wide laterotemporal fenestra would be odd in an enantiornithine. While I was writing this, Andrea Cau published a post on this topic and reported that he recovered Falcatakely as a noasaurid, which would be quite the phylogenetic jump, but it only takes three more steps in the Lori matrix, so is more parsimonious than the enantiornithine option. It falls out as an elaphrosaurine, so could relate to e.g. Afromimus. The non-beak contradictory characters here include a lack of antorbital fossa lacrimal foramen, long posterior lacrimal process and triradiate palatine, which seem more convincing to me. Additional evidence against these latter two positions is the absence of small ceratosaur (Masiaksaurus is twice as big) or large enantiornithine (Maevarano elements are much smaller) postcrania. Andrea reported (translated) "It takes 6 further steps to place it in Coelurosauria, and in that case it is a basal dromaeosaurid: interesting in that regard to note that Rahonavis , known from the same Formation, has also been hypothesized to be a basal dromaeosaurid. Can we rule out that Falcatakely is the (still unknown) skull of Rahonavis? The estimated dimensions of the two animals coincide." Forcing Falcatakely to be Rahonavis only requires one more step, which is pretty impressive. In their (amazing) osteology, Forster et al. (2020) refer an isolated dentary "found near the Rahonavis holotype (its precise location was not recorded during excavation)" which does not match Falcatakely's upper jaw, being upcurved and extensively toothed. But it is similar to other unenlagiines like Buitreraptor and Austroraptor. So much as we have two synsacrum types at this size, unenlagiine-like Rahonavis and Sapeornis-like, we have two cranial types, unenlagiine-like and Falcatakely which is Sapeornis-like in the combination of reduced maxillary dentition, triradiate palatine, modified/reduced antorbital fossa, anteriorly limited naris and strong postorbital-jugal articulation.
Referred dentary of Rahonavis ostromi (FMNH PA 740) as a transparent CT reconstruction (after Forster et al., 2020).
Thus my best guess is that Falcatakely is a basal avialan belonging to the same taxon as FMNH PA 741. But this comes with a huge chunk of salt as it is so far removed temporally and geographically from potential comparable sister taxa. Which is actually a common problem with this part of the tree, as shown by Balaur (= Elopteryx?), Hesperonychus, Imperobator and even Rahonavis itself. We compare these Late Cretaceous taxa to our far more complete Early Cretaceous Jehol record and say Hesperonychus is sorta like Microraptor, Falcatakely is kind of like Sapeornis and Balaur is Jeholornis-grade, but North America, Africa and Europe had their own avialan fauna for 70 million years before them that we're basically unaware of. If the only alvarezsauroid we had was Mononykus' holotype, could we place it correctly as a basal maniraptoran? If the only oviraptorosaur we had was Citipati's skull, would we recover that correctly as the sister taxon of Paraves? I think that's the position we find ourselves in with Falcatakely, and that future discoveries of African small theropods will lead to new interpretations.
References- O'Connor and Forster, 2010. A Late Cretaceous (Maastrichtian) avifauna from the Maevarano Formation, Madagascar. Journal of Vertebrate Paleontology. 30(4), 1178-1201.
Cau, 2020 online. theropoda.blogspot.com/2020/11/falcatakely-eterodossia-e-pluralismo.html
Forster, O'Connor, Chiappe and Turner, 2020. The osteology of the Late Cretaceous paravian Rahonavis ostromi from Madagascar. Palaeontologia Electronica. 23(2):a31.
O'Connor, Turner, Groenke, Felice, Rogers, Krause and Rahantarisoa, 2020. Late Cretaceous bird from Madagascar reveals unique development of beaks. Nature. DOI: 10.1038/s41586-020-2945-x
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Antarctic Ichthyornis solved
So I've been doing some major updates to the Database for what will probably be a New Years upload, including the ornithuromorph section. One rather sad entry as it currently stands is the Antarctic Ichthyornis-
I? sp. (Zinsmeister, 1985)
Late Cretaceous
Seymour Island, Antarctica
Reference- Zinsmeister, 1985. 1985 Seymour Island expedition. Antarctic Journal of U.S. 20, 41-42.
Now with Googling I found the original paper online, which allowed only a bit of improvement-
I? sp. (Zinsmeister, 1985)
Late Maastrictian, Late Cretaceous
Lopez de Bertodano Formation, Seymour Island, Antarctica
Material- several elements
Comments- Zinsmeister (1985) states "several small bones tentatively identified as belonging to the Cretaceous bird Ichthyornis were discovered in the upper Cretaceous Lopez de Bertodano formation."
Reference- Zinsmeister, 1985. 1985 Seymour Island expedition. Antarctic Journal of U.S. 20, 41-42.
So I saw that Zinsmeister worked with Chatterjee in the 80s, who found the Polarornis holotype in the same place two years before that. I emailed Chatterjee about it, who replied-
"It was misidentified in the field. These were some shark teeth."
Mystery solved! But can we do better? Here's an Ichthyornis tooth-
Right eleventh dentary tooth of Ichthyornis dispar (YPM 1450) (after Field et al., 2018).
And here's the array of shark teeth from the Lopez de Bertodano Formation of Seymour Island (from a January 2011 expedition). Can we find any easily confusable matches?
Chondrichthyan teeth from the Lopez de Bertodano Formation (scale 10 mm) (after Otero et al., 2014).
I think the circled 16 and 17 are pretty decent matches for a field identification, though much larger if compared directly. Figures 6-17 are all identified as Odontaspidae indet., which covers any morphology similar to Ichthyornis. Add in the fact that they were by far the most abundant teeth recovered (8 samples versus 1-3 for the other taxa), and I think we have a nice solution on our hands.
I wonder how many other weird records are out there that are based on initial misidentification but stay in the literature because nobody ever publishes a correction?
References- Otero, Gutstein, Vargas, Rubilar-Rogers, Yury-Yañez, Bastías and Ramírez, 2014. New chondrichthyans from the Upper Cretaceous (Campanian-Maastrichtian) of Seymour and James Ross islands, Antarctica. Journal of Paleontology. 88(3), 411-420.
Field, Hanson, Burnham, Wilson, Super, Ehret, Ebersole and Bhullar, 2018. Complete Ichthyornis skull illuminates mosaic assembly of the avian head. Nature. 557, 96-100.
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"Megalosaurus" cloacinus and more - September 2021 Database Update
Hi everyone. I realize it's been ten months since the last post, and that's because I've been prioritizing updating the Database over writing blogs. As a compromise of sorts and to not force people to constantly check the Database updates page, I decided to try out posting when I update including features that could have made it into their own blog post.
One thing I've been doing is working my way through Skawiński et al.'s (2017) paper on Polish Triassic dinosaur reports, which in addition to unnamed fragments, also led to the creation of entries for two supposed Megalosaurus species. silesiacus is a generic carnivorous archosauriform tooth too early to be dinosaurian, while cloacinus has been used for basically every carnivorous archosaur tooth from Rhaetian beds of Germany. The interesting thing about the latter is that workers apparently forgot that it was based on lost teeth described by Quenstedt, not the SMNS tooth figured 47 years later by Huene.
"Zanclodon" silesiacus Jaekel, 1910
= Megalosaurus silesiacus (Jaekel, 1910) Kuhn, 1965
Early Anisian, Middle Triassic
Lower Gogolin Formation, Lower Muschelkalk, Poland
Holotype- (University of Griefswalden/Göttinger coll.; lost?) tooth (24x12x5 mm)
Referred- ?(Geological Museum of the Polish Geological Institute-National Research Institute coll.) tooth (Skawiński, Ziegler, Czepiński, Szermański, Tałanda, Surmik and Niedźwiedzki, 2017)
?(Silesian University of Technology, Faculty of Mining and Geology coll.) tooth (37 mm) (Surmik and Brachaniec, 2013)
Comments- Jaekel (1910) noted (translated) "a dinosaur tooth from the lower shell limestone of Upper Silesia, which would probably be the oldest known dinosaur tooth to date. It comes from the Chorzov strata of the lower shell limestone of Gogolin, Upper Silesia, and came to me through the kindness of engineer Fedder in Opole. The crown shown is 24 mm high, 12 mm wide and 5 mm thick, so it is quite strongly compressed and slightly curved backwards. Its edge is extremely finely serrated (Fig. 16 A). I call the form, which for the time being cannot be specified generically, Zanclodon silesiacus. The only difference between [phytosaur Mesorhinosuchus] and this tooth form lies in the fact that the former is somewhat thicker, somewhat less bent back, and that no notch can be detected on the edge." He referred it to Megalosauridae, and Kuhn (1965) later referred it to the genus Megalosaurus. Carrano et al. (2012) correctly noted "could be considered as Theropoda indet., but we cannot rule out the possibility that it represents a 'rauisuchian' archosaur." Surmik and Brachaniec (2013) describe a tooth from Gogolin Quarry in which "a poor state of preservation makes it impossible to identification of the presence of edge serration, however it still shows a slightly curvature and specific both sides flattening" and identify it as seemingly archosaurian. Skawiński et al. (2017) listed this and another tooth labeled as Megalosaurus silesiacus as other material of Zanclodon silesiacus. The latter tooth is stated to be serrated mesially and distally with a density of 12 per 5 mm. They describe the holotype tooth as "Probably lost" and "lost", and place all three teeth as Archosauromorpha indet.. They are more specifically referred to the Teyujagua plus archosauriform clade here given the recurvature and small serrations, as authors from Kuhn onward have noted plesiomorphic theropod teeth are difficult to distinguish from several clades of archosauriforms (e.g. erythrosuchids, euparkeriids) known from the Anisian. The age is far too early for Megalosaurus or another neotheropod, and the presence of serrations is unlike Zanclodon, so neither genus is appropriate. It should also be noted the three Gogolin teeth differ in shape with the Silesian University specimen less recurved and less tapered than the other two, while the Polish Geological Institute specimen is shorter than the holotype and less concave distally. This could be positional variation, but given the lack of proposed synapomorphies could easily represent multiple taxa.
References- Jaekel, 1910. Ueber einen neuen Belodonten aus dem Buntsandstein von Bernburg. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin. 5, 197-229.
Kuhn, 1965. Fossilium Catalogus 1: Animalia. Pars 109: Saurischia. Ysel Press. 94 pp.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Surmik and Brachaniec, 2013. The large superpredators' teeth from Middle Triassic of Poland. Contemporary Trends in Geoscience. 2, 91-94.
Skawiński, Ziegler, Czepiński, Szermański, Tałanda, Surmik and Niedźwiedzki, 2017 (online 2016). A re-evaluation of the historical 'dinosaur' remains from the Middle-Upper Triassic of Poland. Historical Biology. 29(4), 442-472.
Holotype tooth of "Zanclodon" silesiacus (University of Griefswalden/Göttinger coll.; lost?) in labial (A), basal section (B) and more apical section (C) (after Jaekel, 1910).
"Megalosaurus" cloacinus Quenstedt, 1858
= Plateosaurus cloacinus (Quenstedt, 1858) Huene, 1905
= Gresslyosaurus cloacinus (Quenstedt, 1858) Huene, 1932
= Pachysaurus cloacinus (Quenstedt, 1858) Huene, 1932
Rhaetian, Late Triassic
Exter Formation, Germany
Syntypes- (lost) two teeth
Referred- ?(GPIT and SMNS coll.) many teeth (Huene, 1905)
?(SMNS 52457) tooth (~25x11x? mm) (Huene, 1905)
?(SMNS coll.) teeth (Roemer, 1870)
? seven teeth (Miller Endlich, 1870)
Norian-Rhaetian?, Late Triassic
'Lisów Breccia', Poland
?(University of Wroclaw coll.; lost) two teeth (Roemer, 1870)
Early Hettangian, Early Jurassic
Calcaire de Valognes, Manche, France
?(University of Caen coll.; destroyed) tooth (Rioult, 1978)
Comments- Quenstedt (1858) originally described (translated) "barb-shaped teeth, which are sharp and finely serrated on the concave side, but rounded and smooth on the convex side" with a large mesioapically placed wear facet that makes that edge look straight in side view. He also figures a smaller tooth which has mesial serrations apically that transition to a rounded edge basally. These teeth do not share any obvious synapomorphies and differ in elongation (height/FABL ~300% vs. 138%) and transverse thickness (42% vs. 75% of FABL), so may not belong to the same taxon. Miller Endlich (1870) figured seven teeth from the type locality, stating (translated) they "are mostly flat teeth, slightly curved on one side, with fine serrations on the sharp inner edge. The convex side, the back, does not seem to be serrated, but it is not certain." The figured teeth show a wide range of variation, with figure 13 in particular being stout and unrecurved with large serrations, similar to the Lucianosaurus paratype and similarly referrable to Archosauromorpha incertae sedis. The other teeth have small serrations, with 14 and 18 being straight and 15-17 and 19 being recurved, with 14, 18 and 15 being progressively more transversely compressed. As with the syntypes, these exhibit variation which could be positional or interspecific, and share no obvious characters that connect them to each other or the syntypes. Roemer (1870) wrote (translated) "In the Stuttgart Museum I saw teeth from the bone breccia of Bebenhausen near Tubingen, which show the same fine serration of the side edges as the teeth described by Quenstedt, but are not curved in a sickle shape, but are straight. It is very likely that these latter teeth belong to the same dinosaur as the crooked teeth. With these straight teeth from Bebenhausen, the tooth shown in FIGS. 4 and 5 from the Lisów Breccia from Lubsza near Woźniki completely coincides. The double-edged tooth, which is very delicately and regularly notched at the edges, shows a more strongly curved (outer) and a less curved (inner) surface, both of which are smooth except for a very fine, irregular wrinkle. There is also a much smaller tooth of the same type from the same location." The straight Bebenhausen teeth sound similar to Miller Endlich's figures 14 and 18, although the illustrated straight tooth from Lubsza differs from these in having an increased amount of mesiodistal expansion basally. The Lubsza tooth also has this marked basal expansion labiolingually, and both types of root expansion are atypical of dinosaurs, suggesting this is some other type of vertebrate. Dzik and Sulej (2007) suggested it "may have belonged to a phytosaur" without evidence but Skawiński et al. (2017) stated "phytosaur fossils have not been found in the upper Keuper strata in Silesia" and instead placed it in Archosauromorpha indet.. While this could merely mean phytosaurs were rare in that strata, phytosaur teeth don't seem to have expanded roots either (e.g. Nicrosaurus), and it could even be a fish tooth which often have these types of root expansion. Huene (1905) listed the species as "Plateosaurus" cloacinus within Theropoda, stating it includes Rhaetian dentary "Zanclodon cambrensis". In 1908 he places it in Plateosauridae within Theropoda and states (translated) "The originals can no longer be found. The Tübingen collection still has several teeth from Bebenhausen and Schloßlesmuehle, which can be reconciled well with [Quenstedt's] fig. 12 (l. c.), but are larger. The serrations are coarse and short, the mesial carina does not extend all the way to the base." He illustrated a tooth in figure 274 as "From the Rhaetian Bonebed of Bebenhausen near Tübingen. Tooth in nat. Size. The tip is missing. Original in the natural history cabinet in Stuttgart." Regarding cambrensis, Huene states "The teeth have the greatest resemblance to Plateosaurus cloacinus both in the whole shape and in the serrations. Whether it is really the same or just a very similar species, of course, cannot be decided with certainty given the scanty material", which is not explicit enough to evaluate given published details. Huene later (1932) assigns cloacinus to Teratosauridae within Carnosauria, listed as both Pachysaurus cloacinus (pg. 6) and Gresslyosaurus cloacinus (pg. 72, 114). Steel (1970) calls it Gresslyosaurus cloacinus within Plateosauridae. Buffetaut et al. (1991) mentions "A tooth referred to Megalosaurus cloacinusQuenstedt, from the Lower Hettangian of the Calcaire de Valognes at Valognes (Manche), [which] has been mentioned by Rioult (1978a) as having been destroyed by an air raid on the University of Caen in 1944." Without additional details, it can only be said that the timing suggests a neotheropod. Carrano et al. (2012) incorrectly claimed SMNS 52457, apparently the tooth in Huene's (1908) figure 274, is "the holotype and only specimen" of cloacinus, when Huene stated it was only one of "Many teeth ... in the stone quarries of the Schoenbuch (e.g. Bebenhausen, Schloesslesmuehle), Wuerttemberg; in the university collection in Tubingen and in the natural history cabinet in Stuttgart", and that Quenstedt's originals were lost. SMNS 52457 could be made into a neotype, but this must be done explicitly (ICZN Article 75.3) and so has not been
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Hyracotherium :: ARK: Survival Evolved General Discussions
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Hyracotherium is the early horse dated to have been alive after the extinction of the dinosaurs. This small creature is the direct descendant of modern horses today. I think it would be cool if this creature was implimented into the game don't you think? Here is some info I found on them: Name: Hyracotherium (Greek for "hyrax-like mammal"); pronounced HIGH-rack-oh-THEE-ree-um; also known as Eohippus (Greek for "dawn horse") Habitat: Woodlands of North America and Western Europe Historical...
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https://m.famousfix.com/list/lishui
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FamousFix.com
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https://www.famousfix.com/list/lishui
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1.
Lishui Airport
Airport
Overview: Lishui Airport is a civil and military dual-use airport being built to serve the city of Lishui in Zhejiang Province, China. The airport project began in 2009, and it received approval from the national ...
0 0
2.
Fengyangshan–Baishanzu National Nature Reserve
Nature reserve in China
Overview: Fengyangshan-Baishanzu National Nature Reserve is a nature reserve in southwest Zhejiang province of eastern China. The reserve lies within Longquan and Qingyuan counties. The highest peak of Zhejiang ...
0 0
3.
Qingtian County
County in Zhejiang, People's Republic of China
Overview: Qingtian (Chinese: 青田; pinyin: Qīngtián; Wade–Giles: Ch'ing-t'ien; literally: 'azure field'), is a county in southeastern Zhejiang Province, on the middle-lower reaches of the Ou River ...
0 0
4.
Longquan
County-level city in Zhejiang, People's Republic of China
Overview: Longquan (simplified Chinese: 龙泉; traditional Chinese: 龍泉; pinyin: Lóngquán; literally: 'dragon spring') is a county-level city and former county under the administration of the prefec ...
0 0
5.
Jingning She Autonomous County
Autonomous county in Zhejiang, People's Republic of China
Overview: Jingning She Autonomous County (simplified Chinese: 景宁畲族自治县; traditional Chinese: 景寧畬族自治縣; pinyin: Jǐngníng Shēzú Zìzhìxiàn) is an autonomous county for the She people, under the jurisdiction ...
0 0
6.
Lishui
Prefecture-level city in Zhejiang, People's Republic of China
Overview: Lishui (simplified Chinese: 丽水; traditional Chinese: 麗水; pinyin: Líshuǐ; Wu: Lih-sy) is a prefecture-level city in the southwest of Zhejiang province, People's Republic of China. It borders ...
0 0
7.
Liandu District
District in Zhejiang, People's Republic of China
Overview: Liandu District (simplified Chinese: 莲都区; traditional Chinese: 蓮都區; pinyin: Liándū Qū) is the central urban district of the prefecture-level city of Lishui in Zhejiang Province, China. It was ...
0 0
8.
Songyang County
County in Zhejiang, People's Republic of China
Overview: Songyang County (simplified Chinese: 松阳县; traditional Chinese: 松陽縣; pinyin: Sōngyáng xiàn) is a county in the southwest of Zhejiang province, China. It is under the administration of the Lishui ...
0 0
9.
Yunhe County
County in Zhejiang, People's Republic of China
Overview: Yunhe County (simplified Chinese: 云和县; traditional Chinese: 雲和縣; pinyin: Yúnhé Xiàn) is a county in the southwest of Zhejiang province, China. It is under the administration of the Lishui city ...
0 0
10.
Jinyun County
County in Zhejiang, People's Republic of China
Overview: Jinyun County (simplified Chinese: 缙云县; traditional Chinese: 縉雲縣; pinyin: Jìnyún Xiàn) is a county of south-central Zhejiang province, China. It is under the administration of the Lishui City ...
0 0
11.
Qingyuan County, Zhejiang
County in Zhejiang, People's Republic of China
Overview: Qingyuan County (simplified Chinese: 庆元县; traditional Chinese: 慶元縣; pinyin: Qìngyuán Xiàn) is a county in Lishui, in the south of Zhejiang province, China, bordering Fujian province to the ...
0 0
12.
Huangmaojian
Mountain in Zhejiang, China
Overview: Huangmaojian (Chinese: 黄茅尖; pinyin: Huángmáojiān; literally: 'Black Speargrass Peak') is a 1,930-meter (6,330 ft) mountain in Longquan County in southwest of Zhejiang province in eastern ...
0 0
13.
Suichang County
County in Zhejiang, China
Overview: Suichang County (Chinese: 遂昌县; pinyin: Suíchāng Xiàn) is a county under the jurisdiction of Lishui City, in the southwest of Zhejiang Province, China, bordering Fujian province to the southwest ...
0 0
14.
Yaxi, Zhejiang
Town in Zhejiang, People's Republic of China
Overview: Yaxi (Chinese: 雅溪; pinyin: Yǎxī) is a town in Liandu District, Lishui, Zhejiang province, China. As of 2020, it administers the following 19 villages:
0 0
15.
Zhejiangosaurus
Extinct genus of dinosaurs
Overview: Zhejiangosaurus (meaning "Zhejiang lizard") is an extinct genus of nodosaurid dinosaur from the Upper Cretaceous (Cenomanian stage) of Zhejiang, eastern China. It was first named by a group of Chinese ...
0 0
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https://cyberleninka.ru/article/n/the-braincase-of-bissektipelta-archibaldi-new-insights-into-endocranial-osteology-vasculature-and-paleoneurobiology-of
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THE BRAINCASE OF BISSEKTIPELTA ARCHIBALDI - NEW INSIGHTS INTO ENDOCRANIAL OSTEOLOGY, VASCULATURE, AND PALEONEUROBIOLOGY OF ANKYLOSAURIAN DINOSAURS Текст научной статьи по специальности « Биологические
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https://cyberleninka.ru/article/n/the-braincase-of-bissektipelta-archibaldi-new-insights-into-endocranial-osteology-vasculature-and-paleoneurobiology-of/og
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https://cyberleninka.ru/article/n/the-braincase-of-bissektipelta-archibaldi-new-insights-into-endocranial-osteology-vasculature-and-paleoneurobiology-of/og
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[
"научная статья бесплатно на тему THE BRAINCASE OF BISSEKTIPELTA ARCHIBALDI - NEW INSIGHTS INTO ENDOCRANIAL OSTEOLOGY",
"VASCULATURE",
"AND PALEONEUROBIOLOGY OF ANKYLOSAURIAN DINOSAURS текст научной работы по биологическим наукам из научного журнала Biological Communications. DINOSAURIA",
"ANKYLOSAURIA",
"ENDOCAST",
"BLOOD VESSELS",
"PALEOBIOLOGY",
"LATE CRETACEOUS",
"UZBEKISTAN"
] | null |
[
"Kuzmin Ivan",
"Petrov Ivan",
"Averianov Alexander",
"Boitsova Elizaveta",
"Skutschas Pavel",
"Sues Hans-Dieter"
] |
2020-08-10T00:00:00
|
We describe in detail three braincases of the ankylosaur Bissektipelta archibaldi from the Late Cretaceous (Turonian) of Uzbekistan with the aid of computed tomography, segmentation, and 3D modeling. Bissektipelta archibaldi is confirmed as a valid taxon and attributed to Ankylosaurinae based on the results of a phylogenetic analysis. The topographic relationships between the elements forming the braincase are determined using a newly referred specimen with preserved sutures, which is an exceedingly rare condition for ankylosaurs. The mesethmoid appears to be a separate ossification in the newly referred specimen ZIN PH 281/16. We revise and discuss features of the neurocranial osteology in Ankylosauria and propose new diagnostic characters for a number of its subclades. We present a 3D model of the braincase vasculature of Bissektipelta and comment on vascular patterns of armored dinosaurs. A complex vascular network piercing the skull roof and the wall of the braincase is reported for ankylosaurs for the first time. We imply the presence of a lepidosaur-like dorsal head vein and the venous parietal sinus in the adductor cavity of Bissektipelta. We suggest that the presence of the dorsal head vein in dinosaurs is a plesiomorphic diapsid trait, and extant archosaur groups independently lost the vessel. A study of two complete endocranial casts of Bissektipelta allowed us to compare endocranial anatomy within Ankylosauria and infer an extremely developed sense of smell, a keen sense of hearing at lower frequencies (100-3000 Hz), and the presence of physiological mechanisms for precise temperature control of neurosensory tissues at least in derived ankylosaurids.
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КиберЛенинка
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https://cyberleninka.ru/article/n/the-braincase-of-bissektipelta-archibaldi-new-insights-into-endocranial-osteology-vasculature-and-paleoneurobiology-of
|
FULL COMMUNICATIONS
PALAEONTOLOGY
The braincase of Bissektipelta archibaldi — new insights into endocranial osteology, vasculature, and paleoneurobiology of ankylosaurian dinosaurs
Ivan Kuzmin1, Ivan Petrov2, Alexander Averianov3, Elizaveta Boitsova1, Pavel Skutschas1, and Hans-Dieter Sues4
1Department of Vertebrate Zoology, Faculty of Biology, Saint Petersburg State University, Universitetskaya nab., 7-9, Saint Petersburg, 199034, Russian Federation; 2Saint Petersburg City Palace of Youth Creativity, Nevsky pr., 39A, Saint Petersburg, 191011, Russian Federation;
3Zoological Institute, Russian Academy of Sciences, Universitetskaya nab., 1, Saint Petersburg, 199034, Russian Federation;
4Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, MRC 121, P.O. Box 37012, Washington, DC 20013-7012, USA
Address correspondence and requests for materials to Ivan Kuzmin, kuzminit@mail.ru
Abstract
Citation: Kuzmin, I., Petrov, I., Averianov, A., Boitsova, E., Skutschas, P., and Sues, H.-D. 2020. The braincase of Bissektipelta archibaldi — new insights into endocranial osteology, vasculature, and paleoneurobiology of ankylosaurian dinosaurs. Bio. Comm. 65(2): 85-156. https://doi.org/10.21638/spbu03.2020.201
Authors' information: Ivan Kuzmin, Master of Sci. in Biology, PhD student, Junior Researcher, orcid.org/0000-0003-3086-2237; Ivan Petrov, School student, orcid.org/0000-0003-3617-2317; Alexander Averianov, Dr. of Sci. in Biology, Head of Laboratory, orcid.org/0000-0001-5948-0799; Elizaveta Boitsova, Master of Sci. in Biology, orcid.org/0000-0001-8590-9835; Pavel Skutschas, Dr. of Sci. in Biology, Associate Professor, orcid.org/0000-0001-8093-2905; Hans-Dieter Sues, PhD, Senior Scientist, orcid.org/0000-0002-9911-7254
Manuscript Editor: Nikita Zelenkov, Cabineth of Palaeoornithology, Borissiak Palaeontological Institute, Russian Academy of Sciences, Moscow, Russia
Received: November 21, 2019;
Revised: February 25, 2020;
Accepted: March 10, 2020.
Copyright: © 2020 Kuzmin et al. This is an open-access article distributed under the terms of the License Agreement with Saint Petersburg State University, which permits to the authors unrestricted distribution, and self-archiving free of charge.
Funding: The field work in 1997-2006 was funded by the National Science Foundation (EAR-9804771 and EAR-0207004 to J. D.Archibald and H.-D. Sues), the National Geographic Society (5901-97 and 628198 to J. D.Archibald and H.-D. Sues), and the Navoi Mining and Metallurgy Combinat. The laboratory research received support from the Russian Science Foundation (19-1400020). AA was supported by the Zoological Institute, Russian Academy of Sciences (project AAAA-A19-119032590102-7).
Competing interests: The authors have declared that no competing interests exist.
We describe in detail three braincases of the ankylosaur Bissektipelta archibaldi from the Late Cretaceous (Turonian) of Uzbekistan with the aid of computed tomography, segmentation, and 3D modeling. Bissektipelta archibaldi is confirmed as a valid taxon and attributed to Ankylosaurinae based on the results of a phylogenetic analysis. The topographic relationships between the elements forming the braincase are determined using a newly referred specimen with preserved sutures, which is an exceedingly rare condition for ankylosaurs. The mesethmoid appears to be a separate ossification in the newly referred specimen ZIN PH 281/16. We revise and discuss features of the neurocranial osteology in Ankylosauria and propose new diagnostic characters for a number of its subclades. We present a 3D model of the braincase vasculature of Bissektipelta and comment on vascular patterns of armored dinosaurs. A complex vascular network piercing the skull roof and the wall of the braincase is reported for ankylosaurs for the first time. We imply the presence of a lepidosaur-like dorsal head vein and the venous parietal sinus in the adductor cavity of Bissektipelta. We suggest that the presence of the dorsal head vein in dinosaurs is a ple-siomorphic diapsid trait, and extant archosaur groups independently lost the vessel. A study of two complete endocranial casts of Bissektipelta allowed us to compare endocranial anatomy within Ankylosauria and infer an extremely developed sense of smell, a keen sense of hearing at lower frequencies (1003000 Hz), and the presence of physiological mechanisms for precise temperature control of neurosensory tissues at least in derived ankylosaurids. Keywords: Dinosauria, Ankylosauria, endocast, blood vessels, paleobiology, Late Cretaceous, Uzbekistan.
Introduction
Ankylosaurs constitute a clade of quadrupedal heavily armored ornithischian dinosaurs. Their remains are known from the Jurassic to the Late Cretaceous from every continent except Africa (Tumanova, 1987; Vickaryous et al., 2004). Aspects of ankylosaurian anatomy, phylogeny, and paleobiogeography have been thoroughly studied in the last few decades (e.g., Maryanska, 1977; Tumanova, 1987, 2012; Coombs and Maryanska, 1990; Carpenter, 2001; Vickaryous et al., 2004; Thompson et al., 2012; Arbour and Currie, 2016). Despite this progress, our knowledge of the neurocranial osteology and endocranial morphology within
Table 1. Measurements of the studied braincases of Bissektipelta archibaldi. All linear measurements in millimeters
Parameter ZIN PH 1/16 ZIN PH 281/16 ZIN PH 2329/16
Length from the anterior margin of the sphenethmoidal complex to the posterior tip of occipital condyle 89.2 82.7 84
Depth from the dorsal tip of the laterosphenoid capitate process to the ventral margin of the parabasisphenoid 60.4 58.5 -
Dorsoventral depth of the cranial nerve II foramen 8 6.8 -
Paroccipital process, dorsoventral depth at the mid-section 23.5 22.5 -
Occipital condyle, dorsoventral depth 23.8 21.5 21
Occipital condyle, transversal breadth 36 31.4 42.6
Basioccipital, transversal breadth at the basioccipital-parabasisphenoid contact 52 42 46
Basioccipital, length from the posterior tip of the condyle to the basioccipital-parabasisphenoid contact, in sagittal plane 36 30 35
Foramen magnum, transversal breadth 22 18 18
Foramen magnum, dorsoventral height 19 20 19
Parabasisphenoid, transversal breadth between basipterygoid processes 33 23.8 34
the clade is comparatively poor (see the recent review by Paulina-Carabajal et al. [2018]).
A number of isolated specimens belonging to An-kylosauria are known from the Late Cretaceous of Central Asia (Averianov, 2009). Bissektipelta archibaldi is the only valid ankylosaur species from the territory of the former USSR reported to date. It was initially described as "Amtosaurus" archibaldi based upon a single braincase incorporating the skull roof from the Late Cretaceous Bissekty Formation of Uzbekistan (Averianov, 2002). Later, it was re-assigned to a new genus (Bissektipelta) by Parish and Barrett (2004) as these authors concluded the type species of "Amtosaurus" "A. magnus" is nondiagnostic and should be considered a nomen dubium. Since the initial description, the affinities and phyloge-netic position of Bissektipelta have been debated (Averianov, 2002; Parish and Barrett, 2004; Tumanova, 2012; Arbour and Currie, 2016; see "Phylogenetic analysis" below) but have never been formally assessed. Recently, Alifanov and Saveliev (2019) described a high-quality synthetic endocast made from the holotype of Bissektipelta archibaldi. However, many of their anatomical interpretations and biological inferences appear to be controversial and in need of further review.
Here, we redescribe in detail the holotype of Bissektipelta archibaldi (ZIN PH 1/16) with the aid of CT scanning. Additionally, two new ankylosaur braincases from the Bissekty Formation are described and assigned to the same species. One of these (ZIN PH 281/16) preserves clear sutures between the elements forming the brain-case, which is exceedingly rare for ankylosaurs. Endo-casts for two studied specimens have been made, which is the largest sample for a single species of ankylosaurs to date. A thorough review of the literature and com-
parison between the described taxa allowed us to propose new and revise previously known braincase characters from the most current taxon-character matrices of ankylosaurs (Thompson et al., 2012; Arbour and Currie, 2016; Arbour and Evans, 2017; Zheng et al., 2018) and subsequently test the phylogenetic relationships of Bissektipelta. Based on a solid phylogenetic framework and detailed digital endocranial casts, we discuss aspects of cranial vasculature and inferences concerning the paleobiology of ankylosaurs.
Material and methods
Institutional abbreviations. OUVC, Ohio University Vertebrate Collection, USA; ZIN PH, Paleoherpetologi-cal Collection, Zoological Institute, Russian Academy of Sciences, Saint Petersburg, Russia.
Material. The studied material comprises three braincases: the holotype of Bissektipelta archibaldi (ZIN PH 1/16) and two newly described specimens (ZIN PH 281/16 and ZIN PH 2329/16). The material came from the Late Cretaceous (Turonian) Bissekty Formation at the Dzharakuduk locality in the Central Kyzylkum Desert, Uzbekistan. The measurements for the specimens are provided in Table 1.
The holotype of Bissektipelta archibaldi ZIN PH 1/16 is a well-preserved, fully ossified braincase with a partial skull roof. This specimen was the only known cranial material of the Bissekty ankylosaur and constituted the basis of the original description of "Amtosaurus" ar-chibaldi (Averianov, 2002) and subsequent taxonomic reappraisal of this taxon as Bissektipelta archibaldi (Parish and Barrett, 2004). The newly described specimens include ZIN PH 281/16, a partial braincase of slightly
smaller size with open sutures between some bones, and ZIN PH 2329/16, which is similar in size to the holotype of Bissektipelta archibaldi (Table 1). ZIN PH 2329/16 preserves most of the braincase and part of the skull roof. The sutures cannot be traced in ZIN PH 2329/16 because it is damaged and partially covered with matrix.
Computed tomography. The holotype ZIN PH 1/16 and the referred specimen ZIN PH 281/16 were X-ray CT scanned using a Toshiba Aquilon 64 medical tomographer at 0.5 mm slice thickness, 120 kV, and 300 mA. The resulting stacks compile 334 images (512x512x334 resolution) in DICOM format for ZIN PH 1/16 and 149 images (512x512x149 resolution) for ZIN PH 281/16. Data acquired from CT scans were imported into the visualization software Amira 6.3.0 (FEI-VSG Company) and manually segmented. The resulting 3D models have the voxel size of 0.313x0.313x0.3 for ZIN PH 1/16 and 0.625x0.625x0.8 for ZIN PH 281/16. Measurements on the 3D models were performed using Amira 6.3.0 and MeshLab (Cignoni et al., 2008). The CT scan data and 3D models are available upon request from the first author.
Description of the holotype of
Bissektipelta archibaldi ZIN PH 1/16 (Figs. 1-9)
General comments. The braincase of Bissektipelta is highly ossified, and the bones of the skull roof are completely fused to it. Most sutures were obliterated. We do not support previous assumptions about incompletely ossified portions of some elements in the holotype (e.g., basal tubera, right basipterygoid process, occipital condyle, and the distal tip of the paroccipital process; Averianov, 2002) and regard those as preservational artifacts. These structures are variably preserved in the three studied braincases (notably, also in the smaller specimen ZIN PH 281/16) and are frequently broken off. The braincase is non-pneumatic. CT scans show that no internal pneumatic structures are present. Externally, there is neither the medial pharyngeal recess on the ventral surface of the basicranium nor a well-defined anterior/lateral pneumatic recess on the lateral surface of the parabasisphenoid.
Skull roof. The preserved skull roof has a relatively flat dorsal surface (Fig. 1A, B). Sutures are completely obliterated and are not evident either on the specimen's surface or in the CT images. General observations suggest that ZIN PH 1/16 preserves the posterior portion of the skull roof that corresponds to the frontoparietal region of taxa with known sutural relationships (e.g., Pinacosaurus, Maryañska, 1977; Godefroit et al., 1999; Hill et al., 2003; Kunbarrasaurus, Leahey et al., 2015; Ce-darpelta, Carpenter et al., 2001; "ZhongyuansaurusXu et al., 2007, = Gobisaurus in Arbour and Currie, 2016: Fig. 6D). A truncated Y-shaped groove that separates
three polygonal areas of remodeled bone (= caputegu-lae; Blows, 2001; Arbour and Currie, 2013a) is present. The resulting areas are identified here as the paired pos-terolateral nuchal caputegulae (nuca, Fig. 1B) and central parietal caputegulum (paca, Fig. 1B) using the terminology of Arbour and Currie (2013a). Each groove terminates in a pronounced pit; a small offshoot of the left groove is present and is directed anteromedially from the corresponding pit. The CT data for ZIN PH 1/16 shows that these grooves, paired pits, and the skull roof surface are pierced by numerous vascular foramina that connect through canals with the endocranial cavity and the lateral braincase wall. The left branch of the Y-shaped groove interrupts its course for one millimeter, and there is a short contact between the left nuchal and the central parietal caputegulae. The skull roof surface of ZIN PH 1/16 was remodeled, but it is uncertain if osteo-dermal ossifications were involved in that process. According to the hypothesis of Vickaryous et al. (2001a), "the superficial furrows that divide the cranium.. .represent the areas of coosification between adjacent cephalic osteoderms". The presence of the Y-shaped groove thus implies that the osteoderms are preserved and co-ossified with the skull roof in ZIN PH 1/16. There is no frontoparietal depression. The posterior edge of the skull roof is broken off. The broken lateral edges of the skull roof overhang the adductor cavities, and there are no traces of the supratemporal fenestrae.
The boundaries between the skull roof and brain-case are partly recognized on the preserved right par-occipital process in the occipital view (Fig. 1E), and are inferred on the lateral surface of the specimen based on the position of small vascular foramina that frequently lie near the border between the skull roof and brain-case (Galton, 1988; Galton and Knoll, 2006; Fig. 2A). The pattern of facets on the skull roof in the referred specimen ZIN PH 281/16 supports this reconstruction of the boundaries in the holotype. The parietal has two posterolateral processes that are sutured ventrolaterally to the dorsal surface of the paroccipital processes and medially to the supraoccipital (the latter contact is hard to trace; Fig. 1E). The posterolateral processes are an-teroposteriorly thin and oriented almost perpendicular to the sagittal plane of the skull. The posterior surface of the posterolateral processes is slightly posteroventrally-anterodorsally inclined. On the lateral aspect of ZIN PH 1/16, the skull roof appears to form an almost horizontal, slightly posteroventrally inclined contact with the braincase posterior to the capitate process of the lateros-phenoid and a gently anteroventrally inclined contact anteriorly (Fig. 2A). Posteriorly in lateral view, the parietal roofs a small vascular recess (nvr, Fig. 2A, B) and forms the dorsomedial wall of the adductor cavity. Here the skull roof reaches its greatest dorsoventral thickness of 21 millimeters.
oc
Fig. 1. ZIN PH 1/16, holotype of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. Photographs and corresponding CT-based models in dorsal (A, B), ventral (C, D), and occipital (E, F) views. Scale bars each equal 1 cm. Abbreviations: bo, basioccipital; bofe, basioccipital fenestra; bpt, basipterygoid process; bt, basal tuber; CN II — XII, cranial nerve foramina; fm, foramen magnum; fr, frontal; fvOC, foramen for orbitocerebral vein; fvSo, foramen for supraoccipital vein; MF, metotic foramen; nuca, nuchal caputegulum; nvf, neurovascular foramen; nvp, neurovascular pit; oc, occipital condyle; olff (CN I), olfactory fenestra; oto, otoccipital; p, parietal; paca, parietal caputegulum; pbsro-ios, fused parabasisphenoid rostrum and interorbital septum; pop, paroccipital process; proaf, proatlas facet; ptf, posttemporal fenestra; so, supraoccipital; tencr, tentorial crest.
Ventral surface of the basicranium. The basioccipi-tal and the parabasisphenoid meet at an angle of approximately 90o in ZIN PH 1/16; the suture between these bones is evident in lateral and ventral views (Figs. 1D, 2). Overall the basioccipital is massive and robust. The neck of the occipital condyle is barely defined. The ventral surface of the basioccipital is posteroventrally oriented, concave, and broad; it is slightly wider lateromedially than the corresponding surface of the parabasisphenoid. The basal tubera (= sphenoccipital tubera in Kurza-nov and Tumanova [1978] and Tumanova [1987]) are rounded, anteroposteriorly thin, and project laterally (bt, Fig. 1D, F). The basioccipital fenestra (bofe, Fig. 1D) is present as a distinct blind fissure on the ventral surface between the basal tubera. CT data show that two small, presumably vascular canals extend from it anteriorly and posteriorly inside the bone and gradually disappear in the trabeculae. The basioccipital fenestra is present in the same location ventral to the occipital condyle in present-day crocodylians; a small vein traverses this foramen (Owen, 1850).
The parabasisphenoid has a triangular, anteroven-trally oriented ventral surface (Fig. 1C, D). The surface between the basipterygoid processes is mediolaterally wider and gradually tapers anteriorly. The left basiptery-goid process is slightly incomplete (bpt, Fig. 1D). The basipterygoid process is a knob that projects ventro-laterally. It is oval in cross-section, with the longer axis directed anteriorly. Its anteroposterior length is nearly twice the mediolateral width at its base. The surface between the basipterygoid processes is relatively flat; there is a shallow depression on each side close to the base of the process. Only the base of the fused parabasisphenoid rostrum (= cultriform process) and the ossified/calcified interorbital septum is preserved. It is situated anterior to the basipterygoid processes (pbsro-ios, Fig. 1D). The base of the fused parabasisphenoid rostrum-interorbital septum extends obliquely anteriorly to the spheneth-moidal complex, where it merges with the septum that separated the olfactory bulbs (= mesethmoid in Miyas-hita et al. [2011]; Figs. 1D, 2E). Regarding the preserved part, the base of these elements is slightly transversally constricted at its mid-length and then expands anteriorly. On each side of the fused parabasisphenoid rostrum-interorbital septum are longitudinal depressions (possibly for the sphenopalatine artery; gaSP, Fig. 2D). A pronounced ridge ventral to the foramen for the optic cranial nerve (CN II) delimits the course of the longitudinal depression dorsally. No sutures in the region of the sphenethmoidal complex are discernable.
Occipital surface. The occipital surface is inclined at the angle of about 125o to the dorsal surface of the skull (Fig. 2A). When the specimen is held such that its skull roof surface is oriented horizontally, the occipital condyle is directed posteroventrally and barely projects
beyond the occipital plane. The articular surface of the condyle is crescent-shaped and transversely elongated (lateromedial length nearly 1.5 times larger its dorsoven-tral depth; Fig. 1E, F). The articular surface of the con-dyle is slightly eroded. The suture with the otoccipital is visible on the right lateral and posterior surfaces of the condyle (Figs. 1E; 2A); it indicates that the otoccipitals formed the dorsolateral corners of the occipital condyle. The posterior surface of the basioccipital ventral to the condyle is notably arched dorsally and overall faces pos-teroventrally (Fig. 2).
The foramen magnum is nearly circular. Its lateral and dorsal margins are formed by the otoccipitals; the supraoccipital appears to be excluded from the dorsal margin. Paired triangular surfaces (proatlas facets) project from dorsolateral corners of the foramen magnum (proaf, Fig. 1F). They merge medially and form a dorsal shelf over the foramen magnum. The proatlas facets overhang rounded notches that are sometimes interpreted as the path of the first spinal nerve (Kurzanov and Tumanova, 1978; Parish and Barrett, 2004). In addition, or as an alternative hypothesis, these sulci can correspond to the route of a venous vessel that branches off from the longitudinal venous sinus or its posterior expansion (occipital sinus) at the foramen magnum and courses ventrolaterally (Porter, 2015). Just dorsal to the proatlas facets, there are paired small foramina with associated grooves. These foramina pierce the occipital surface of the braincase directly to the endocranial cavity and likely transmitted small supraoccipital veins (fvSo, Fig. 1F). A venous foramen in a similar position above the foramen magnum was noted for "Amtosaurus magnus" (Kurzanov and Tumanova, 1978). Medial to these vascular foramina, on the assumed posterior surface of the supraoccipital, there is the base of the sagittal nuchal crest; dorsally, this surface is obscured by damage.
Paired rounded posttemporal fenestrae are present lateral to the sagittal nuchal crest (ptf, Fig. 1E, F). In general, the posttemporal fenestrae appear to lie near the contact of the parietal, supraoccipital, and otoccipital, but the precise sutural pattern is entirely obscured on the left side and is not clear on the right. The presumed parietal-otoccipital suture is situated at the ventrolateral margin of the posttemporal fenestra; thus, the ventral margin of the posttemporal fenestra is likely formed by the paroccipital process of the otoccipital, and its dorso-lateral margin by the parietal. It is likely that the supra-occipital contributed to the margin of the fenestra medially; alternatively, the medial margin of the fenestra may have been formed by the otoccipital and the parietal. The posttemporal fenestra pierces anteriorly into a small recess on either side of ZIN PH 1/16. This recess is evident in lateral view (nvr+g, Fig. 2A, B); it lies dorsal to the paroccipital process and medial to the adductor cavity. A notable groove is present at the anterior margin of the
Fig. 2. ZIN PH 1/16, holotype of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. Photographs and corresponding CT-based models in right lateral (A, B), left lateral (C, D), and oblique left lateral (E, F) views. Scale bars each equal 1 cm. Abbreviations: bo, basioccipital; bpt, basipterygoid process; bt, basal tuber; ca+vSO, canal for supraorbital artery and vein; ci, crista interfenestralis; CN II — XII, cranial nerve foramina; CN III / aOr, foramen for oculomotor nerve or orbital artery; CR+FO, columellar recess and fenestra ovalis; crp, crista prootica; fa+vSO, foramen for supraorbital artery and vein; faCC, foramen for cerebral carotid artery; faSP, foramen for sphenopalatine artery; fr, frontal; fvOC, foramen for orbitocerebral vein; gaSP, groove for sphenopalatine artery; ls, laterosphenoid; meth, mesethmoid; MF, metotic foramen; nvg, neurovascular groove; nvr+g, neurovascular recess and groove; olff (CN I), olfactory fenestra; ors, orbitosphenoid; oto, otoc-cipital; p, parietal; pbs, parabasisphenoid; pbsro-ios, fused parabasisphenoid rostrum and interorbital septum; pop, paroccipital process; pro, prootic; r, ridge; speth, sphenethmoid.
recess, suggesting the course of a blood vessel along the dorsomedial wall of the adductor cavity. Both the walls of the recess and the anterior groove are pierced by numerous small vascular foramina.
The preserved right paroccipital process extends laterally and slightly posteriorly and is incomplete distally (pop, Fig. 1). It is anteroposteriorly thin at its distal end and thick and robust at its base. The process is relatively narrow dorsoventrally; its depth equals the height of the foramen magnum. Two blunt ridges curve dorso-laterally and converge to form the ventral margin of the paroccipital process. The ventral margin of the paroc-cipital process is slightly arched dorsally and is nearly at the same level as the ventral border of the foramen magnum. Dorsally, the process is sutured to the skull roof. There is a small but pronounced depression at the posterior surface of the paroccipital process.
Lateral braincase wall. The elements forming the lateral wall of the braincase are fused (e.g., the sphen-ethmoidal complex, the orbitosphenoid, the lateros-phenoid, the parabasisphenoid, the prootic, and the otoccipital). No clear sutures can be observed, with the exception of the basioccipital-otoccipital suture on the condyle on the right side and the suture between the basioccipital and parabasisphenoid. All structures are paired, and the right and left sides of ZIN PH 1/16 have the same general structure and proportions. The lateral wall is penetrated by numerous neurovascular foramina (Fig. 2). These are clustered into two major groups and are relatively closely spaced within the cluster. The anterior group includes the foramina for CN II-VII and two primarily vascular foramina (for the cerebral carotid artery and the sphenopalatine artery and vein). The posterior group is situated ventral to the base of the par-occipital process and comprises the columellar recess/ fenestra ovalis, the metotic foramen, and the foramina for CN XII.
The two clusters of foramina are separated by a flattened strip of bone that extends ventrally between the basioccipital and the parabasisphenoid portion of the basal tuber. Dorsally, its posterior margin arches over the fenestra ovalis onto the ventral edge of the paroc-cipital process (crp, Fig. 2D). This structure corresponds to a poorly developed crista prootica (= otosphenoidal crest in Sampson and Witmer, 2007) that in diapsids separates the more anterior cranial nerve foramina from the posterior depression containing ear-related structures (fenestra ovalis plus metotic foramen). Generally in diapsids, the crista prootica arches posterodorsally from the parabasisphenoid lateral surface, just above the basipterygoid process. The crista prootica in Bissek-tipelta contacts ventrally the basal tubera instead of the basipterygoid proces. This is likely due to the highly divergent braincase structure of Bissektipelta (and other ankylosaurs) from the basic diapsid pattern, specifically
the posterior position of the basipterygoid processes close to the basal tubera.
The olfactory fenestrae are the only neurovascular foramina directed anteriorly instead of laterally (olff, Fig. 2D, F). They are paired and separated by a thick bony septum (= mesethmoid in Miyashita et al. [2011]). They are the largest neurovascular foramina and approach the foramen magnum in size. The olfactory fe-nestrae housed short paired olfactory bulbs and the ethmoid vessels, and they communicated directly with the olfactory region of the nasal cavity (Miyashita et al., 2011). The internal walls of the olfactory fenestrae are covered by numerous anteroposterior grooves, indicating that a large number of neurovascular bundles passed through them (nvg, Fig. 2E, F). The two separate cavities for the olfactory bulbs converge posteriorly into a single chamber that is separated from the rest of the endocra-nial cavity by a rounded constriction.
Only the base of the broken preorbital septum is preserved. The preorbital septum is a thin transversal bony lamina that separates the nasal and orbital cavities in ankylosaurs; it was first named by Maryanska (1977) (= ectethmoid in Miyashita et al. [2011]; see the description of ZIN PH 2329/16 below). Between the base of the preorbital septum and the anterior cluster of neurovas-cular foramina, the surface of the braincase wall bears no foramina and has dorsoventral striations. The largest foramen among the anterior cluster is that for CN V; the opening for CN II is the second largest. The foramina for the cerebral carotid artery and for the sphenopala-tine vessels are prominent and nearly equal in size (faCC and faSP, Fig. 2F). The large recess of the ganglion of CN V has a triangular projection from its dorsal margin that separates the anteriorly directed groove for CN V1 (ophthalmic branch of the trigeminal nerve) from postero-ventrally directed grooves for CN V2+3 (maxillary and mandibular branches of the trigeminal nerve; see Holli-day and Witmer [2007]). The small foramen for CN VII lies in the same large recess with that for CN V and is separated by a small ridge from the latter. The foramen for CN II is separated from the more posterior foramina by a thick vertical strut of bone. A small groove on the ventral margin of the foramen for CN II possibly indicates the course of a small vessel (Fig. 2D, E). There is a prominent depression on the lateral braincase wall dorsal to the foramen for CN IV and anterior to the adductor cavity (the postocular shelf is not preserved here in ZIN PH 1/16; see the description of ZIN PH 2329/16 below). The depression is pierced by two foramina for the orbitocerebral vein and a series of smaller vascular openings (fvOC, Fig. 2F).
The columellar recess/fenestra ovalis (CR/FO), the metotic foramen (MF), and three external foramina for CN XII are closely spaced and situated in a single depression ventral to the paroccipital process. This de-
pression is bordered by the crista prootica anteriorly, the basal tuber ventrally, and the prominent blunt ridge posteriorly (r, Fig. 2D). The latter connects with the ventral margin of the paroccipital process so that the foramina for CN XII are not evident in posterior view (Fig. 1E, F). The external openings of the CR/FO and MF are almost equal in size and large. The crista interfenestralis (= ventral ramus of opisthotic in more basal archosaurs; e.g., Gower, 2002; Sobral et al., 2016) separates FO and MF (ci, Fig. 2B). It is a slightly anteroventrally inclined lamina of bone that is barely visible in posterior view. The three foramina for CN XII are almost vertically arranged posterior to MF. The posteriormost foramen is the largest of the three. The anteriormost foramen for CN XII is the smallest and lies below MF.
Endocranial surface. The complex endocranial surface can be anteroposteriorly subdivided into four main concave regions (olfactory and cerebral fossae, and two fossae anterior and posterior to the otic capsule) separated by convex crests (Fig. 3). The anterior part of the endocranial cavity in ZIN PH 1/16 corresponds the pos-teriormost portion of the olfactory region of the nasal cavity (distinguished by rugose walls with numerous neurovascular grooves) and paired cavities of the olfactory bulbs that merge posteriorly into the cavity for the olfactory tract (olfbc and olftc, Fig. 3). The olfactory tract cavity is constricted laterally by paired blunt crests, which emphasize the division between the olfactory region anteriorly and the cerebral fossa posteriorly.
The cerebral cavity is circumscribed by the blunt olfactory crest anteriorly and by the tentorial crest (sensu Sedlmayr [2002]) posteriorly on each side (olfcr and tencr, Fig. 3A). Several neurovascular structures pierce the surface of the cerebral fossa, including the foramen for CN IV and two conspicuous foramina for the orbito-cerebral vein (Fig. 3A). The surface of the cerebral cavity has a gently corrugated texture but lacks prominent vascular valleculae, indicating that the brain was not in close relationship to the endocranial wall and loosely fitted the cerebral cavity (Evans, 2005). The large transverse groove for CN II is offset anteroventrally and opens posteriorly into the cerebral cavity (Fig. 3B). Its dorsal margin forms a blunt crest that arches posterodorsally onto the lateral endocranial surface on each side and merges with the tentorial crest. This oblique crest marks the subdivision of the cerebral fossa into two smaller fossae, roughly corresponding to the cerebral hemispheres anteriorly and the optic lobes posteriorly. The ventral margin of the CN II groove forms a sharp crest that denotes the anterior dorsal limit of the hypophyseal cavity.
The cerebral cavity merges into the hypophyseal cavity ventrally (hypc, Fig. 3). The hypophyseal cavity is comparatively shallow, being half the depth of the cerebral cavity. Foramina for the cerebral carotid and sphenopalatine arteries and for CN III pierce its surface
(Fig. 3). The internal foramen for CN III is unexpectedly situated ventrally, well in the limits of the hypophyseal cavity. A groove connects the internal openings of the sphenopalatine artery and CN III, raising the possibility that the latter may actually be a vascular foramen, perhaps for a branch of the cerebral carotid/sphenopalatine artery (e.g., the orbital artery of extant birds) or for the orbital/hypophyseal vein that drains into the cavernous sinus (Bruner, 1907; Porter and Witmer, 2015; Porter and Witmer, 2016a). In that case, the actual CN III would leave the braincase through the dorsally situated foramen for CN IV, as in Euoplocephalus (Miyashita et al., 2011). We tentatively follow the initial description by Averianov (2002) and maintain a conservative interpretation of the foramen in question as for CN III.
The dorsum sellae bulges over the hypophyseal cavity dorsally. It has a short anterior triangular projection surrounded by two grooves medially. This projection is also evident in the referred specimen of Bissektipelta ZIN PH 281/16, in "Amtosaurus magnus" (Kurzanov and Tumanova, 1978), and is possibly present, though less developed, in several other Mongolian taxa (Averianov, 2002; Parish and Barrett, 2004). We regard these grooves as vascular impressions that indicate the course of posterior venous branches of the cavernous sinus (caudo-ventral cerebral veins) or, as an alternative hypothesis, the course of the caudal encephalic arteries (Sedlmayr, 2002; Porter, 2015; Porter et al., 2016). Posteriorly to the dorsum sellae, the floor of the endocranial surface is essentially flat.
The tentorial crest is prominent; ventrally, it is confluent with the lateral aspect of the dorsum sellae, arches anterodorsally over the anterior margin of the foramen for CN V, and then curves posterodorsally and extends to the roof of the endocranial cavity (tencr, Fig. 3A). The internal opening for CN VI pierces the base of the tento-rial crest; the foramen for CN VII lies dorsolateral to it (Fig. 3B). The tentorial crest circumscribes a fossa dorsal to the foramen for CN V that was likely occupied by the cerebellum and a large venous vessel (middle cerebral vein; vMC, Fig. 3A). The latter opened externally through a series of foramina at the posterodorsal aspect of the fossa. The floccular (= auricular) fossa is very shallow.
The medial wall of the otic capsule is incompletely ossified (sc+ves in Fig. 3A). The amount of the exposure may have been exaggerated by postmortem fracture; however, both the referred specimens have largely medially open vestibules. The recesses for the vestibule, common crus, and lagena are medially open and confluent with the endocranial cavity. Paired unossified fossae with unfinished margins at the floor of the endocranial cavity mark the position of the lagenae (lagf, Fig. 3B). These fossae are comparatively large and probably contained additional structures such as supportive vascular plexus. A bifurcating groove extends posterodorsally
Fig. 3. ZIN PH 1/16, holotype of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. Parasagitally sectioned CT-based models showing left endocranial surface, in medial (A) and posteromedial (B) views. Scale bar equals 1 cm. Abbreviations: cerc, cerebral cavity; CN II — XII, cranial nerve foramina; CN III / aOr, foramen for oculomotor nerve or orbital artery; faCC, foramen for cerebral carotid artery; faSP, foramen for sphenopalatine artery; fvOC, foramen for orbitocerebral vein; hypc, hypophyseal cavity; lagf, lagenar fossa; MF, metotic foramen; olfbc, olfactory bulb cavity; olfcr, olfactory crest; olftc, olfactory tract cavity; sc+ves, cavities of semicircular canals and vestibule; tencr, tentorial crest; vMC, groove for middle cerebral vein.
from the internal foramen for CN VII and indicates the course of CN VIII (Fig. 3A); a similar reconstruction of this region was made for some ornithopods (Hopson, 1979; Sobral et al., 2012). The internal opening of the metotic foramen (MF) is just posterior to the otic capsule. The metotic foramen is undivided (see discussion in Rieppel [1985]; Gower and Weber, 1998; Sobral et al., 2012) and likely transmitted the perilymphatic sac, CN IX-XI, and the posterior cerebral vein (vagal vein in Sedlmayr [2002]). The wall between the vestibular recess and the MF is incised. This notch indicates the position of the incompletely ossified perilymphatic foramen that transmitted the perilymphatic sac from the otic capsule to the MF (Baird, 1960; Gower, 2002; Gower and Walker, 2002). Two larger internal foramina for CN XII pierce the endocranial wall posterior to the MF; a single small opening is just ventral to it. An extensive shallow depression with a deeper pit above these structures indicates the position of the occipital venous sinus (Fig. 3B; Sedlmayr, 2002; Witmer et al., 2008; Porter, 2015).
Endocast. The endocast of ZIN PH 1/16 generated from a CT scan data is complete, undistorted, and relatively detailed (Fig. 4; Table 2). It comprises casts of the endocranial cavity, cranial nerves, both endos-seous labyrinths, and vascular canals. The morphology of the inner ear and braincase vasculature of ZIN PH 1/16 are described in separate sections below. The brain of Bissektipelta loosely fitted the endocranial cavity as is common for many non-avian dinosaurs and for reptiles in general (Hopson, 1979; Witmer et al., 2008). Thus, the produced endocast is more a cast of the meninges (including endocranial venous sinuses) rather than the brain itself. Nevertheless, it appears to be a faithful inference of gross morphology of the brain as is suggested by recent research on extant archosaurs (Watanabe et al., 2019). Additionally, the endocranial vessels of various extant diapsids have a rather conservative pattern (Porter, 2015; see Vasculature and Fig. 9 below), and their disposition revealed on the endocast of Bissektipelta is a reliable proxy for recognition of major brain divisions.
olft ch
Fig. 4. ZIN PH 1/16, holotype of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. Cranial endocast with endosseous labyrinth of the inner ear in dorsal (A), ventral (B), and left lateral (C) views. Scale bar equals 1 cm. Abbreviations: aCC, cerebral carotid artery and vein; aSP, sphenopalatine artery and vein; cbl, cerebellum; ch, cerebral hemisphere; lab, endosseous labyrinth; MF, metotic foramen passage; olfb, olfactory bulb; olfc, olfactory cavity cast; olft, olfactory tract; optt, optic tectum of the midbrain; vn?, vomero-nasal bulbs?; II — XII, cranial nerves.
The endocranial cast is elongate and relatively straight, with low angles of cerebral and pontine flexures of about 30o; the cranial nerves and the fenestra ovalis of the labyrinth are correspondingly linearly arranged (Fig. 4C). The olfactory bulbs diverge anteriorly at an angle of approximately 80o from a short and broad olfactory tract (CN I; olfb and olft in Fig. 4A). The bulbs and the tract are nearly circular in cross-section. The olfactory ratio (the ratio of the greatest diameter of the olfactory bulb to the greatest diameter of the cerebral hemisphere regardless of their orientation; Zelenitsky et al., 2009) equals nearly 63-69 %. It is comparable to that of other ankylosaurs (see Discussion below) and suggests proportionally large olfactory bulbs in Bissektipelta and other ankylosaurs. Anteriorly, the endocast of each olfactory bulb terminates in a rounded expansion with wrinkled walls that corresponds to the olfactory region of the nasal cavity (olfc, Fig. 4A). The wrinkles most likely represent neurovascular bundles passing to and from the endocranial cavity, and part of them was visualized as the vascular olfactory plexus (olfP, Fig. 6; see "Vasculature" section below). Paired cavities within the ossified ethmoidal region, ventral to the olfactory tract, were segmented (vn?, Fig. 4A, B). A similar cavity was also found ventral to the broken mesethmoid in the referred specimen ZIN PH 281/16 (see Figs. 11-14). These struc-
tures lie below the olfactory complex of Bissektipelta and could correspond to the part of the vomeronasal bulb or vascular sinuses. These structures are reported for the first time in dinosaurs and require additional study to elucidate their nature.
The short olfactory tract posteriorly merges into a rounded cast of the cerebral cavity. The anterior part of the cavity's endocast reaches maximum dorsoventral depth and lateromedial breadth (Table 2) and corresponds to the cerebral hemispheres (ch, Fig. 4A). The cerebral hemispheres are relatively discrete on the endo-cast. The absence of a dorsal groove between the hemispheres indicates that the dorsal longitudinal venous sinus occupied the space above the latter.
The endocast of Bissektipelta lacks the dorsal dural peak present on endocasts of stegosaurs (Galton, 1988; Galton, 2001; Leahey et al., 2015: Fig. 10), most sauro-pods (Witmer et al., 2008), and some theropods (e.g., Sampson and Witmer, 2007). Dorsal expansions of dinosaur endocasts have been interpreted as an unossified gap plugged with cartilage in life (Hopson, 1979) or as corresponding to extensive dural sinuses that may have surrounded the pineal complex (Sampson and Witmer, 2007; Witmer et al., 2008). In Bissektipelta, the only structure of the endocast that may correspond to the pineal complex is a conspicuous median vessel (medVs,
Table 2. Endocast measurements of Bissektipelta archibaldi. All linear measurements in millimeters; volume in cm3
Parameter ZIN PH 1/16 ZIN PH 281/16
Whole endocast length 103 70
Endocast length without the cast of the olfactory region of the nasal cavity 85 -
Endocast volume (without vessels and nerves) 53
Endocast width across cerebral hemispheres 33 29
Olfactory bulb maximum cross-sectional diameter 15 -
Olfactory tract width 19.5 12
Pituiatary depth 19 18
Pituitary diameter 16.5 12.5
Fig. 7). Its canal pierces the skull roof all the way through to the endocast and connects to the anterior branching plexuses laterally (see Vasculature below). The median vessel emerges at the surface of the endocast just an-terodorsal to the inferred division between the cerebral hemispheres and the optic lobes (assessed by the position of the cerebrotectal venous sinus and the disposition of crests on the endocranial surface), at the level of the optic chiasm and the pituitary. This position broadly corresponds to that of the pineal complex in other di-apsids (e.g., Sphenodon; Dendy, 1911). Notably, the pineal complex, optic chiasm, and the neurohypophysis are all diencephalic derivates. However, the external pineal (parietal) foramen was lost early in archosauriform evolution (Hopson, 1979; character 63 in Nesbitt, 2011); extant birds have a pineal that lies internally within the braincase, adjacent to the skull roof (Ralph, 1970). Thus, we doubt that the median canal of Bissektipelta contained a pineal/parapineal organ that was exposed on the dorsal surface of the skull roof and consider the structure a vascular canal. However, noting its remarkable position, we hypothesize this canal enclosed vessels that may have been connected to pineal vasculature. The point of emergence of the median vessel from the endocast thus marks a possible position of the pineal complex in Bissektipelta.
The optic chiasm is located on the ventral surface of the endocast of Bissektipelta, below the cerebral hemispheres and just anterior to the hypophysis (Fig. 4). Each CN II leaves the braincase by a separate lateral foramen. The endocasts of the canals for CN II and of the optic chiasm form a single straight trunk that is oriented strictly perpendicular to the longitudinal axis of the braincase (Fig. 4B).
The complete cast of the hypophyseal (pituitary) fossa is present just posterior to the optic chiasm and ventral to the hemispheres (Fig. 4B, C). The pituitary projects vertically from the ventral surface of the en-docast. Overall, the pituitary cast is a tubular structure with an even diameter throughout; the stalk itself is not
expressed. It is relatively short dorsoventrally (its dorso-ventral depth equals nearly 19 mm and is half the depth of the cerebral cavity above it), broad, and nearly circular in cross-section (Table 2). The hypophyseal fossa apparently contained the infundibulum (hypophyseal stalk) and the hypophysis, which were likely surrounded by the cavernous venous sinus, as it in extant archosaurs (Neumeier and Lametschwandtner, 1994; Sedlmayr, 2002; Porter et al., 2016; Porter and Witmer, 2016a). The hypophyseal fossa of Bissektipelta was well vascularized. Large cerebral carotid arteries entered the hypophyseal cavity transversely as in most other ankylosaurs (e.g., Paulina-Carabajal et al., 2018); the sphenopalatine arteries branched off of them and left the hypophyseal cavity slightly anteriorly (a+vCC and a+vSP in Fig. 4B, C). The cerebral carotid and sphenopalatine veins that drained the cavernous sinus and the orbit/palate apparently shared canals with similarly named arteries. The stalk of CN III appears roughly at the mid-height of the endo-cast of the hypophyseal cavity, which occupies an unusually ventral position compared to those on most other dinosaur endocasts (Fig. 4C). A swelling on the lateral surface of the pituitary endocast connects the CN III trunk with the sphenopalatine artery endocast, which, combined with its low position, possibly indicates that the former represented a vessel (e.g., orbital artery and vein) rather than a nerve.
The optic lobes of the midbrain (optt, Fig. 4C) are not directly discernable on the endocast of ZIN PH 1/16 as they were likely overlain by sizable dural venous sinuses. The approximate position of the midbrain could be determined through the disposition of major encephalic vessels and general topographic cues of the diapsid brain (reviewed by Hopson [1979], Witmer et al. [2008], and others). The optic tectum of the midbrain in Bissektipelta apparently laid between the cerebral hemispheres anteriorly and the cerebellum posteriorly (Fig. 4C); thus, the brain had a linear arrangement that is similar to that in extant crocodiles, plesiomorphic for dinosaurs in general, and characteristic of many ornithischians in par-
ticular (Hopson, 1979; Balanoff and Bever, 2017). The endocranial cast of Bissektipelta is slightly constricted mediolaterally at the level of the optic tectum; its dorsal outline smoothly arches posteroventrally in lateral view. The short trunk of CN IV projects anterolaterally and slightly ventrally from the endocast above the pituitary (Fig. 4C). If we assume that the canal for CN III housed vascular structures rather than the actual cranial nerve, CN III must have left the braincase through the canal for CN IV.
The cerebellum is not distinctly expressed, and there is no prominent flocculus on the endocast of ZIN PH 1/16 (cbl, Fig. 4A, C). The region of the endocast that corresponds to the cerebellum is posterior to a groove reflecting the position of the tentorial crest. The cerebellum was circumscribed by extensive dural vessels, e.g., the middle cerebral vein anterodorsally and the longitudinal sinus (torcular Herophili part) dorsally (Figs. 6A, 7A, 9A).
Part of the endocast corresponding to the medulla oblongata is nearly as broad mediolaterally as it is anteroposterior^ long. The structure of the medulla ob-longata is obscured by extensive occipital venous sinus (sOc, Fig. 7A). The ventral surface of the brainstem is essentially flat and straight in lateral view; it is only slightly notched behind the endocast of the hypophyseal fossa anteriorly (Fig. 4B, C).
The single large trunk of CN V expands shortly after its emergence from the lateroventral surface of the endocast and superficially subdivides into three lobes (Fig. 4C). This expansion of CN V endocast likely corresponds to the Gasserian ganglion, and the three lobes reflect its main branches — the ophthalmic (CN VI), maxillary (CN VII), and mandibular (CN VIII) nerves (see Holliday and Witmer [2007] for a survey of the diapsid condition). The middle cerebral vein exited the brain-case together with CN V (vMC, Fig. 9). The endocast of CN VI extends anteroventrally and slightly laterally from the ventral surface of the brainstem. It comes off at the level of CN V, passes by the hypophyseal cavity, and exits the braincase through a separate foramen anterior to CN V (Figs. 2, 3, 4). The trunk of CN VII emerges between the endocasts of CN V and the inner ear and parallels the course of CN V. CN VIII was not digitally rendered; however, a groove at the endocranial surface of ZIN PH 1/16 that extends posterodorsally from the internal foramen of CN VII into the inner ear recess, just below the ampullary spaces, corresponds to the course of CN VIII (Fig. 3A). CN IX and CN X share the same exit via the metotic passage. The endocast of the MF is directly posterior to that of the inner ear and is relatively large (comparable to the endocast of CN II and slightly smaller than that of CN V). Three trunks of CN XII are evenly spaced posterior to the MF endocast; the anteri-ormost trunk is the smallest and lies adjacent to the MF.
Inner ear. The endosseous labyrinth of the inner ear was digitally reconstructed for both sides of ZIN PH 1/16 (Figs. 4, 5). The endosseous labyrinth is the endocast of inner skull cavities that carried the endolymphat-ic (otic or membranous) labyrinth surrounded by the perilymphatic (periotic) labyrinth (Baird, 1960; Witmer et al., 2008). Part of the perilymphatic labyrinth associated with semicircular canals is uniform among reptiles and closely matches the semicircular ducts of the endolymphatic labyrinth in shape. The lower part of the perilymphatic labyrinth that surrounds the saccule and the cochlear duct (lagena) of the endolymphatic system has a more complex structure that obscures the form of the endolymphatic labyrinth (Baird, 1960). The endosseous labyrinths of Bissektipelta, as well as those of other dinosaurs, reflect the structure of both the endolymphatic and perilymphatic systems as a whole. The perilymphatic labyrinth of Bissektipelta has an extracapsular portion (perilymphatic sac) that extends posteromedi-ally into the undivided metotic passage (MF) to participate in a compensatory secondary tympanic membrane (pls, Figs. 4B, 5B). This is a common condition for many diapsids including Sphenodon, basal archosaurs, and dinosaurs (Baird, 1960; Gower, 2002; Gower and Walker, 2002; Witmer et al., 2008). The position of extracapsular portion of the perilymphatic sac is marked by a notch between the vestibular recess of the inner ear cavity and the MF and was digitally visualized as part of the endosseous labyrinth (pls, Fig. 5B, C).
The endosseous labyrinths from both sides of ZIN PH 1/16 are undistorted and symmetrical. The medial aspects of both labyrinths are incomplete due to incomplete endocranial ossification of the otic capsules (Fig. 5C). The semicircular canals are robust. The am-pullar regions are not discrete and are present as expansions at proximal ends of the canals. Each semicircular canal lies in a single plane and does not curve beyond its limits. The anterior canal is the tallest and the largest of the three; it is roughly circular in shape (asc, Fig. 5B). The angle between the anterior and posterior canals equals approximately 90o. The posterior semicircular canal has a marked elliptical shape and is relatively low (psc, Fig. 5B); the anterior canal is one and a half times taller than the posterior one. The common crus is low (nearly equals the depth of the posterior canal) and broad (nearly twice the average canal breadth) (crc, Fig. 5). The lateral semicircular canal is ovoid in shape and appears to be equal to or only slightly smaller than the posterior canal (lsc, Fig. 5D). The utricular and sac-cular compartments of the endolymphatic labyrinth are not apparent as they were laterally covered by the peri-otic cistern of the perilymphatic labyrinth (as in other diapsids; Baird, 1960).
Below the level of semicircular canals, the endosse-ous labyrinth is markedly constricted anteroposteriorly.
Fig. 5. ZIN PH 1/16, holotype of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. Digital reconstruction of the endos-seous labyrinth of the left inner ear in anterior (A), lateral (B), medial (C), and oblique dorsolateral (D) views. Scale bar equals 1 cm. Abbreviations: asc, anterior semicircular canal; cochd, endosseous cochlear duct; CR, columellar recess; crc, crus communis; CR+FO, external openning of columellar recess leading to fenestra ovalis; lsc, lateral semicircular canal; pls, perilymphatic sac; psc, posterior semicircular canal.
This point is level with the columellar recess and the fenestra ovalis and marks the approximate line of division of the endosseous cochlear duct (lagena). The cochlear duct curves ventromedially below the vestibular portion of the labyrinth (cochd, Fig. 5A). Large unossified spaces below the endocranial cavity on both sides of ZIN PH 1/16 (lagf, Fig. 3B) were segmented as parts of the inner ear labyrinth. Although these structures are natural, symmetric, and observed in all studied specimens, we doubt that the actual cochlea was that elongate in Bissek-tipelta and curved below the endocast as in birds (e.g., Witmer et al., 2008). We suggest that these spaces could have contained either enlarged outgrowths of the peri-lymphatic sac and/or supportive neurovascular tissues of the inner ear. Overall, they just might have been plugged
with cartilage. Accounting for the uncertainty regarding the actual extent of the cochlear duct in ZIN PH 1/16, we perform two sets of measurements of its length (Table 3): a conservative assessment that does not include the medioventral portion of the cochlear endocast and an extended assessment that accounts for a complete model length. The presence of the extracapsular portion of the perilymphatic sac in Bissektipelta (which is the continuation of the scala tympani that encircles the la-gena medially in archosaurs; see Baird, 1960: Fig. 2) suggests that small portions of the perilymphatic labyrinth bulged out intracranially and, thus, were not visualized in the model. According to our approximate estimates, the length of the endosseous cochlear duct equals nearly 10-11 mm under a conservative measurement and 11-
Table 3. Endosseous labyrinth measurements and hearing properties of Bissektipelta archibaldi. The best frequency of hearing and the high-frequency hearing limit are calculated based on the equations from Gleich et al. (2005). We assume that the basilar papilla represents only two thirds of the cochlear duct length as in Gleich et al. (2005). For ZIN PH 1/16, two types of measurements were conducted — a more conservative approach (when a straighter line through the cochlea was measured) and an extended approach (when a strongly ventromedially curved line through the cochlea was measured). A single set of cochlear dimensions was taken from ZIN PH 281/16. All linear measurements in millimeters, hearing frequencies in hertz
Parameter ZIN PH 1/16 ZIN PH 281/16
Left labyrinth cochlear duct length, conservative 10.8
Left labyrinth cochlear duct length, extended 13.9 13.1
Right labyrinth cochlear duct length, conservative 10.1
Right labyrinth cochlear duct length, extended 11.6 14.4
Mean cochlear duct length for both labyrinths, conservative 10.45
Mean cochlear duct length for both labyrinths, extended 12.75 13.75
Best frequency of hearing, conservative 1002
Best frequency of hearing, extended 682 576
High-frequency hearing limit, conservative 2889
High-frequency hearing limit, extended 2299 2105
14 mm under an extended assessment, which amounts to 38-41 % and 41-53 % of the overall height of the en-dosseous labyrinth (the height of the vestibular part is around 15-16 mm). Thus, the lagena of Bissektipelta was moderately elongate.
A long canal extends laterally from the cochlear duct of ZIN PH 1/16 (CR, Fig. 5A). It has two parallel oblique sharp margins along its sides. We hypothesize that this structure represents the stapedial recess partly enclosed in bone due to extensive ossification of the lateral wall of the braincase in ZIN PH 1/16. In dorsal and anterior views, the distal part of the recess delimited by the aforementioned margins resembles the shape of the oblique stapedial footplate (Fig. 5A, D). Thus, the actual fenestra ovalis was likely displaced internally from the lateral surface of the braincase.
Vasculature. The CT data allowed digital reconstruction of a complex pattern of blood vessels in the ho-lotype of Bissektipelta archibaldi (Figs. 6-9). Endocranial vasculature and the system of vessels piercing the skull roof and lateral braincase wall has been reconstructed for Bissektipelta based on relative osteological correlates such as grooves and canals within bone (Witmer, 1995; Porter, 2015). Major vessels that are external to the braincase and did not leave direct bony features are only briefly mentioned here and are discussed later (see Discussion). As it is often hard to discriminate which component (arterial/venous) is prevalent in a given os-teological structure (save for well-known features such as the cerebral carotid canal predominated by the arterial component or grooves for the dural venous sinuses at the endocranial surface; see Porter [2015]), we have not distinguished between the types of blood vessels that pierced the skull roof and the braincase wall in our model. However, many of them are considered mainly or exclusively venous as encephalic arteries form a closed network around the brain under the dura matter and do not communicate with the orbital and temporal vessels (Sedlmayr, 2002; Almeida and Campos, 2010, 2011), with the exception of the ethmoid artery that communicates anteriorly with the supraorbital artery to form the nasal artery (Porter et al., 2016; Porter and Witmer, 2016a).
In extant diapsids, the main artery that supplies the braincase is the internal carotid artery and two of its branches — the cerebral carotid and the stapedial arteries (Porter and Witmer, 2015, 2016; Porter et al., 2016) (aIC, aCC, and aST in Fig. 8). In Bissektipelta, each cerebral carotid artery enters almost at the floor of the hypophyseal cavity (a+vCC, Fig. 6A). Small vessels branching off of the cerebral carotid curve anteroven-trally along the floor of the hypophyseal cavity (Fig. 4B). These small lobose vessels, though visualized as parts of the cerebral carotid artery endocast, possibly represent ventral parts of the cavernous sinus that drains into the
cerebral carotid vein (compare Fig. 4B with Neumeier and Lametschwandtner, 1994: Fig. 15). The latter vein shares the canal with the cerebral carotid artery; thus, both vessels are represented by a single trunk in the endocast of ZIN PH 1/16. The cerebral carotid arteries were likely connected medially because extant birds and crocodylians show anastomizing vessels/plexuses in the posteroventral region of the hypophyseal cavity (Sedlmayr, 2002; Porter et al., 2016; Porter and Witmer, 2016a). A horizontal swelling at the posterior surface of the pituitary endocast of ZIN PH 1/16, between cerebral carotid arteries, possibly corresponds to the intercarotid anastomosis (Fig. 4B). Just anterior to its entrance into the hypophyseal cavity, the cerebral carotid artery gives off the sphenopalatine artery (a+vSP, Fig. 6A). It is a distinct but smaller-caliber vessel compared to the cerebral carotid. A possible anterior course of the sphenopalatine artery is marked by a notch and depression on each side of the parabasisphenoid (gaSP in Fig. 2D, aSP in Fig. 8). The artery courses anterodorsally into the nasal region. A similar route of the sphenopalatine artery is present in extant birds (Porter and Witmer, 2016a). The dorsal courses of the common encephalic artery (= cerebral carotid after branching off sphenopalatine artery) and its branches that ramify around the brain inside the endocranial cavity are hard to trace. Possible osteologi-cal correlates are paired grooves on the dorsum sellae that could correspond to the caudal encephalic artery (Fig. 3A).
Another branch of the internal carotid is the sta-pedial artery, which continues anteriorly through the temporal region as the temporoorbital artery and then divides into three main orbital vessels (supra-, infraorbital, and ophthalmotemporal arteries; Sedlmayr, 2002; Porter, 2015) (aST, aTO, aSO, aOpt + aIO in Fig. 8). Anterior to the supratemporal fossa, a large curved canal within the lateral wall of the braincase, dorsal to the foramina for CN II-IV, is interpreted as the passage for the supraorbital artery and vein (preserved on the left side of ZIN PH 1/16 and opened by fracture on the right) (ca+v and fa+vSO in Fig. 2B, F; a+vSO in Fig. 6A; aSo in Fig. 8; vSO in Fig. 9). These vessels accompany each other through their course over the anterior surface of the laterosphenoid and ventral surface of the frontal in extant archosaurs (Porter et al., 2016; Porter and Witmer, 2016a). The supraorbital vessels course external to the bone surface in extant taxa; however, some dinosaurs with heavily ossified skulls (e.g., pachycephalo-saurids) show evidence for the bony enclosure of their branches into canals (Porter, 2015). The same is likely true for ZIN PH 1/16. The supraorbital artery/vein canal communicates via small-caliber vascular canals with the anterior branching plexus of the cranial roof dorsally and with endocranial vessels medially and posteromedi-ally (Figs. 6, 7, 9). The latter corresponds to numerous
Fig. 6. ZIN PH 1/16, holotype of Bissektipelta archibaldifrom the Bissekty Formation (Turonian), Uzbekistan. CT-based models showing braincase vasculature in left lateral view; endocast with surrounding vessels (A), semitransparent view of the braincase showing vessels within the skull roof and lateral braincase wall (B), solid braincase (C). Scale bar equals 1 cm. Abbreviations: a+vCC, cerebral carotid artery and vein; a+vSO, supraorbital artery and vein; a+vSP, sphenopalatine artery and vein; ABP, anterior branching plexus; olfABP, olfactory part of the anterior branching plexus; olfP, olfactory plexus; PBP, posterior branching plexus; PBP-vCD, anastomotic vessel between the posterior branching plexus and the dorsal head vein; PbsVs, parabasisphenoid vasculature; sOc, occipital venous sinus; sP, parietal venous sinus; vCD, dorsal head vein; vg, venous groove; vOC, orbitocerebral vein; vSo, supraoccipital vein; vTOc, transverso-occipital vein. Black arrow heads mark small anastomotic connections between main vascular elements.
Fig. 7. ZIN PH 1/16, holotype of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. CT-based models showing braincase vasculature in dorsal view; endocast with surrounding vessels (A), semitransparent view of the braincase showing vessels within the skull roof (B), solid braincase and skull roof (C). Scale bar equals 1 cm. Abbreviations: ABP, anterior branching plexus; anastABP, anastomotic connection between left and right anterior branching plexuses; medVs, medial vessel; olfABP, olfactory part of the anterior branching plexus; PBP, posterior branching plexus; sOc, occipital venous sinus; sP, parietal venous sinus; vCD, dorsal head vein; vOC, orbitocerebral vein; vSo, supraoccipital vein; vTOc, transverso-occipital vein. Black arrow heads mark small anastomotic connections between main vascular elements. White arrow heads mark connections between the dorsal head vein and the middle cerebral vein.
Fig. 8. ZIN PH 1/16, holotype of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. CT-based model of the braincase in right lateral view showing reconstructed pattern of arteries. Red vessels are reconstructed upon corresponding osteological correlates. Black vessels inferred from broader studies of diapsid vascular patterns. Not to scale. Abbreviations: aCC, cerebral carotid artery; aET, ethmoidal artery; aIC, internal carotid artery; aMan, mandibular artery; aNas, common nasal artery; aOc, occipital artery; aOpt+aIO, ophthalmotemporal and infraorbital arteries; aSO, supraorbital artery; aSP, sphenopalatine artery; aST, stapedial artery; aTO, temporoorbital artery; braSO, dorsal branches of supraorbital artery.
venous communications with endocranial dural veins (mainly, the dorsal longitudinal sinus with its tributaries and the cerebrotectal sinus, see Fig. 9). The supraorbital artery/vein canal also receives a number of small canals from the lateral surface of the braincase. The anterior course of the supraorbital vessels is marked by an antero-ventrally directed groove and the orbitonasal foramen in the preorbital septum (Fig. 8). The latter structure is broken in the holotype but preserved in the referred specimen ZIN PH 2329/16 (Fig. 15). The communication of the supraorbital and ethmoid vessels (situated dorsal to the olfactory tract and bulbs in extant taxa; Almeida and Campos, 2010, 2011; Porter et al., 2016; Porter and Wit-mer, 2016a) occurred through some of the small canals piercing the lateral wall of the braincase (Figs. 6A and 7A) and further anterior to the orbitonasal foramen (see Figs. 8-9). The latter communication of the supraorbital and ethmoid vessels gave rise to the nasal vessels that supplied and drained the nasal cavity.
The encephalic vessels seldom leave direct traces on the endocranial surface (with some notable exceptions; see Evans [2005]); however, their basic pattern appears to be rather conservative among known diapsids (Bruner, 1907; Dendy, 1909; Sedlmayr, 2002; Witmer et
al., 2008; Porter, 2015; Porter and Witmer, 2015, 2016; Porter et al., 2016). Major endocranial veins are recognized as swellings on the endocast surface that communicate with external vasculature via vascular or nervous canals (Hopson, 1979; Sampson and Witmer, 2007; Witmer et al., 2008; Porter, 2015). The latter are important landmarks that trace the course of the vessel. The digital endocast of the holotype of Bissektipelta allows recognition of several dural venous vessels/sinuses and their communications with external vasculature (Fig. 9).
The dorsal longitudinal sinus appears as a shallow but broad prominence on the top of the endocast that extends from the olfactory tract anteriorly to the level of the otic capsules posteriorly (Figs. 7, 9). Anterior to the olfactory tract, the dorsal longitudinal sinus apparently splits into a pair of vessels (olfactory veins in Dendy [1909]; ethmoid vein in Porter and Witmer [2016]; Porter et al., 2016) that overlaid the olfactory bulbs and continued forward to drain the olfactory cavity as nasal veins (vET + vNas in Fig. 9). A large number of neurovascular grooves are preserved around the olfactory bulbs/posterior portion of the olfactory region of the nasal cavity in ZIN PH 1/16 (Fig. 2F). These grooves indicate the presence of a vascular plexus around the ol-
olfP sLon
Fig. 9. ZIN PH 1/16, holotype of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. CT-based models of the cranial endocast and braincase in left lateral view showing reconstructed pattern of veins. A, encephalic veins that drain endocranial cavity, labeled on the endocast; B, semitransparent view of the braincase showing veins within the skull roof and lateral braincase wall (solid), encephalic veins (transparent), and their intercommunications; C, non-transparent view of the braincase showing external braincase veins (solid) and their parts within canals (transparent). Note controversial interpretation of veins at the supratemporal fossa; see Discussion for the preferred reconstruction. Blue vessels are reconstructed upon corresponding osteological correlates. Black vessels inferred from broader studies of diapsid vascular patterns. Not to scale. Abbreviations: ABP, anterior branching plexus; anastv, circum-occipital anastomotic loop between dorsal head vein, transverso-occipital vein, and parietal venous sinus; olfABP, olfactory part of the anterior branching plexus; olfP, olfactory plexus; PBP, posterior branching plexus; sCT, cerebrotectal sinus; sLon, dorsal longitudinal sinus; sOc, occipital venous sinus; sOc/vCP, occipital venous sinus/ posterior head vein; sP, parietal venous sinus; vCD, dorsal head vein; vET, ethmoidal vein; vMC/sT, middle cerebral vein and transverse venous sinus; vNas, common nasal vein; vOC, orbitocerebral vein; vOpt+vIO, ophthalmotemporal and infraorbital veins; vSC, superior cerebral veins; vSo, supraoccipital vein; vSO, supraorbital vein; vST+vTO, stapedial and temporoorbital veins; vTOc, transverso-occipital vein.
factory bulbs that drained into the ethmoid vein and the anterior branching plexus (olfP in Figs. 6-7, 9). In extant birds, the olfactory bulbs are surrounded by a dense venous plexus that eventually drains into the longitudinal sinus via the ethmoidal veins and/or the olfactory sinus and their anastomoses (Sedlmayr, 2002; Porter and Wit-mer, 2016a).
A conspicuous median vessel arises sagittally at the top of the endocast of Bissektipelta, dorsal to the cerebral hemispheres (medVs, Fig. 7). The vessel extends dor-sally within a canal inside the bone and opens through a foramen on the dorsal surface of the skull roof. Along the way, it issues left and right major branches as well as smaller branches in a slightly asymmetrical manner; each major branch communicates with a corresponding anterior branching plexus (Fig. 7A). The median vessel was apparently connected to the dorsal longitudinal sinus ventrally. Undoubtedly this structure was vascular (most likely, venous), as it commences from the endo-cast at the inferred position of the longitudinal sinus and connects to vascular branching plexuses. We are unaware of any similar dorsal extensions of the encephalic vessels in extant diapsids. As noted earlier in the description of the endocast, the position of the median vessel in Bissektipelta generally corresponds to the position of the pineal complex in extant lepidosaurs and birds. In extant diapsids, the pineal complex is well vascularized, supplied by branches of the posterior cerebral artery and drained by the dorsal longitudinal sinus (Dendy, 1909; Ralph, 1970). We doubt that the canal housed the pineal/ parapineal organ itself but hypothesize that the median vessel and its branches may represent dorsal continuations of the pineal complex vasculature.
The dorsal longitudinal sinus is most prominent posteriorly where it received the middle cerebral veins and appears as a broad triangle in the dorsal view (tor-cular Herophili; Dendy, 1909; Sedlmayr, 2002; sOc, Fig. 7A). Posterior to this, the dorsal longitudinal sinus likely bifurcated into two sinus-like posterior cerebral veins (vena cerebralis posterior, vena cephalica posterior, vagal vein, occipital venous sinus of different authors) (sOc, Fig. 7A; sOc/vCP, Fig. 9A). The posterior cerebral veins likely left the endocranial cavity through the foramen magnum, as in most extant diapsids (Bruner, 1907; Sedlmayr, 2002; Witmer et al., 2008; Porter and Witmer, 2015, 2016; Porter et al., 2016). However, Sphenodon shows an important variation of the course of the posterior cerebral vein, which leaves the endocra-nial cavity through the metotic foramen (Dendy, 1909). The same route for the posterior cerebral vein through the metotic foramen was reconstructed for basal croco-dylomorphs (Walker, 1990) and other pseudosuchians (Gower, 2002; Gower and Nesbitt, 2006; Sulej, 2010). Both pathways for the venous drainage (via the metotic foramen/foramen magnum) are likely traced in extant
crocodylians through their development (Dendy, 1909; Sedlmayr, 2002). For Bissektipelta, we imply that most, if not all, of the venous blood left the endocranial cavity posteriorly through the foramen magnum, with possible additional venous drainage through the metotic foramen via the posterior cerebral vein (sOc and sOc/vCP, Fig. 9A). Sobral et al. (2012) arrived at a similar conclusion regarding the pathway of the posterior cerebral vein in the ornithopod Dysalotosaurus.
A pair of small foramina (fvSo in Fig. 1) directly above the foramen magnum of Bissektipelta apparently transmitted small veins and accompanying arteries (su-praoccipital veins; vSo in Figs. 6A, 7A and 9A). In Sphen-odon, these vessels drain from the dura matter and the dorsal part of the occipital sinus extracranially through similarly distributed foramina (Dendy, 1909).
Along its course, the dorsal longitudinal sinus receives several transverse veins that drained lateral aspects of the endocranial cavity. We assume the presence of a number of superior cerebral veins that extended along the lateral aspects of the olfactory tract and the anterior cerebrum, dorsal to CN II, as was described for Sphenodon (venae cerebrales superiores; Dendy, 1909) (vSC, Fig. 9A). Additionally, the presence of corresponding but unidentified vessels dorsal to CN II was discussed for Caiman and reported for fossil endocasts (Hopson, 1979). The presence of superior cerebral veins in Bissektipelta is established by numerous small vascular canals that connected the anterior branching plexus and the canal for supraorbital vessels with endocranial dural veins (Fig. 7A). These veins joined the corresponding ethmoid vein/dorsal longitudinal sinus dorsally.
Posteriorly, at the level of CN IV, conspicuous swellings on the endocast and a pair of the orbitocerebral veins on each side indicate the course of a transverse venous sinus (sCT, Fig. 9A, B). The latter received confusing terminology in the literature: vena cerebri posterior in Hopson (1979); sphenotemporal sinus in Sedlmayr (2002); sphenoparietal sinus in Witmer et al. (2008); and cerebrotectal sinus in Porter et al. (2016) and Porter and Witmer (2016). In extant archosaurs, this venous sinus extends dorsally along the tentorial crest to join the dorsal longitudinal sinus and wedges in between the posterior region of the cerebrum and the optic tectum (Hopson, 1979; Sedlmayr, 2002). A series of veins on each side of the brain in the same region (venae begime-nales superiores) was described for Sphenodon (Dendy, 1909). We use the term "cerebrotectal sinus", as it clearly reflects the anatomical position of the vessel. In Bissek-tipelta, the cerebrotectal sinus extends transversally as a swelling on the posterior aspect of the cerebral endocast (compare Figs. 4C, 6A, and 9A). The orbitocerebral veins drain into the cerebrotectal sinus from the orbital cavity (vOC in Figs. 6A, 9B). The cerebrotectal sinus directly communicates via small vascular canals with the
middle cerebral vein and dorsal head vein/parietal sinus, the posterior branching plexus, and the canal for supraorbital vessels (Figs. 6, 9).
The succeeding large transverse tributary of the dorsal longitudinal sinus with complicated nomenclature is the middle cerebral vein (vena cerebralis media in Bruner [1907]; transverse sinus in Dendy [1909] and Porter and Witmer [2015]; recessus lateralis of longitudinal sinus in Hopson [1979]; rostral petrosal sinus in Sedlmayr [2002]; cerebellotectal sinus in Porter and Witmer [2016] and Porter et al. [2016]) (vMC, Fig. 9). In extant diapsids, it is a large vessel that extends between the optic tectum and cerebellum, just in front of the otic capsule. At the point of its divergence from the longitudinal sinus, the middle cerebral vein is sinus-like and broad, and thus its dorsal portion was designated the transverse, rostral petrosal, or cerebellotectal sinus (Dendy, 1909; Sedlmayr, 2002; Sampson and Witmer, 2007; Porter and Witmer, 2016a; Porter et al., 2016). Ventrally, the sinus drains into one or several smaller and more defined veins (= middle cerebral vein sensu stricto, e.g., Sampson and Witmer, 2007; trans-versotrigeminal vein of Porter and Witmer [2015]; rostral middle cerebral vein in Paulina-Carabajal et al. [2016]) that frequently pass through the trigeminal foramen ex-tracranially. We use the simpler term "middle cerebral vein" for both portions of the vessel (the transverse sinus and its continuations) in an effort to keep the terminology as concise as possible and to ensure compatibility with previous accounts on dinosaurian cranial vasculature (Sampson and Witmer, 2007; Witmer et al., 2008; Miyas-hita et al., 2011; Leahey et al., 2015; Paulina-Carabajal et al., 2016, and others).
In Bissektipelta, the middle cerebral vein/transverse sinus extends dorsally from the foramen for CN V as a bulge on the endocast surface, then arches posterodor-sally, parallel to the anterior semicircular canal, and finally joins the dorsal longitudinal sinus (Figs. 6A and 9). In Bissektipelta, there is no separate branch of the middle cerebral vein that passes extracranially in the lateral direction through its own canal (= rostral middle cerebral vein of some authors). Thus, the middle cerebral vein likely exited the braincase via the large foramen of CN V. Dorsally, a conspicuous posterodorsally curved swelling on the endocast marks the course of the middle cerebral vein. Here, the middle cerebral vein is laterally confluent with the dorsal longitudinal/occipital sinus (Fig. 9A). Three short vascular branches extend posterodorsally and laterally from the middle cerebral vein on each side of ZIN PH 1/16 (Fig. 7A). These vascular branches connect the middle cerebral vein with the external veins of the temporal and occipital regions of the skull (dorsal head vein, transversooccipital vein, parietal sinus; Figs. 6A, 7A, 9B). Additionally, a separate vessel extends from the anterior-most of the three described vascular branches and connects with the cerebrotectal sinus anteriorly (precisely,
with the dorsal orbitocerebral vein, Figs. 6A, 9B). As described, the course of the middle cerebral vein in Bissek-tipelta is consistent with observations on extant diapsids (Bruner, 1907; Dendy, 1909; Porter and Witmer, 2015; Porter and Witmer, 2016a; Porter et al., 2016) and various dinosaurs (see Discussion).
The dorsal head vein and the transverso-occipital vein (caudal middle cerebral vein of some authors) exit the braincase of Bissektipelta via separate foramina on the lateral (nvr+g, Fig. 2B; vCD, Fig. 6B, C) and occipital (ptf, Fig. 1; vTOc, Fig. 7B) surfaces of the skull, correspondingly. However, their endocast suggests that they either represent a single vessel or a continuous anasto-motic loop that extends from the temporal to the occipital region of the skull and maintains the connection with the middle cerebral vein/transverse sinus (Fig. 7A). In Bissektipelta, the groove passes anterior from the foramen of the dorsal head vein (nvr+g in Fig. 2B). This groove corresponds to the continuation of the dorsal head vein; we term this continuation as the parietal sinus (sP, Figs. 6B, 7A, 9B-C) following the terminology of extant squamates (Bruner, 1907; Porter and Witmer, 2015; see also Discussion).
Numerous small openings at the dorsal surface of the skull roof of Bissektipelta lead into the canals within bones that eventually converge ventrally (Fig. 7). This pattern of vascular canals is herein referred to as branching plexus. There are paired anterior and posterior branching plexuses that supplied and drained the skull roof and overlying dermis in Bissektipelta (ABP, Figs. 6-7 and 9). The anterior branching plexus can be subdivided into two parts: one part that lies above the olfactory bulb and the olfactory cavity and is connected ventrally to the ethmoid vessels (olfABP in Fig. 9B) and the other part that lies posteriorly and communicates with the supraorbital vessels ventrally and dural veins medially (ABP in Fig. 9B). Some parts of these canals likely transmitted small branches of the supraorbital artery that pierce the frontal and emerge onto the outer surface of the skull in extant birds (Porter and Witmer, 2016a) and some dinosaurs (Porter, 2015) in a similar way (braSO in Fig. 8). The posterior plexus is situated above the dorsal head vein (PBP in Figs. 6, 9); it is less distinct compared to the anterior plexus and was not visualized on the right side of ZIN PH 1/16. As previously described, small vascular canals integrate the anterior and posterior branching plexuses as well as various endo- and extracranial vessels into a single vascular network around the brain (see Discussion for physiological implications).
Description of ZIN PH 281/16 (Figs. 10-14)
General comments. ZIN PH 281/16 is exquisitely preserved, with fine features of the external and endocra-nial surfaces and clear sutures and facets. It appears to
be only slightly smaller compared to the holotype; most of its measurements are only 5-15 % smaller than those for ZIN PH 1/16 (Table 1). The braincase is externally and internally non-pneumatic, as is evident from the CT data.
Skull roof. ZIN PH 281/16 does not preserve bones of the skull roof and has slightly rugose fine facets on its dorsal surface (Figs. 10A, 11A). This indicates that the skull roof was not completely co-ossified with the braincase in this particular specimen. A lack of fusion between the skull roof and braincase is present in adults of Pinacosaurus (Maryanska, 1977; Tumanova, 1987) and Minotaurasaurus (Miles and Miles, 2009; Penkalski and Tumanova, 2017), and the two cranial components are strongly sutured in adult individuals of other anky-losaurs. The unfused skull roof in ZIN PH 281/16, along with open sutures between individual neurocranial elements, indicates that the specimen probably represents a somatically subadult individual.
Ventral surface of the basicranium. The ventral aspect of the specimen is formed by the basioccipital posteriorly and the parabasisphenoid anteriorly; the suture between these bones is clearly visible on both the lateral and ventral surfaces (Fig. 10C). Unlike in the holotype, the two bones join at an obtuse angle of approximately 120o. The ventral surface of the basioc-cipital is smoothly arched and bears a vascular foramen (basioccipital fenestra). The triangular ventral surface of the parabasisphenoid bears the base of the fused para-basisphenoid rostrum-interorbital septum anteriorly and small, bump-like basipterygoid processes (left one is broken off), which are offset posteriorly, close to the suture between the basioccipital and parabasisphenoid (Fig. 10D).
Occipital surface. The occipital surface of the specimen is formed by the supraoccipital, basioccipital, and paired otoccipitals; the sutures between these bones are easily recognized, unlike in the holotypic cranium and most other known ankylosaurs (Fig. 11C). The occipital surface forms the same angle of about 125o with the skull roof (inferred from the plane of corresponding facets) as in the holotype. The occipital condyle barely projects beyond the occipital plane. It has a more rounded shape compared to those of the holotype and ZIN PH 2329/16. The otoccipitals form the dorsolateral portions of the condyle. The otoccipital-basioccipital suture is evident on both sides of the specimen (Figs. 11C, 12). The suture extends onto the lateral and endocranial surfaces, where it gradually disappears toward the external and internal openings of the metotic foramen, respectively. The foramen magnum is bounded by the basioccipital ventrally and by the otoccipitals laterally and dorsally. The supraoccipital was probably excluded from the dorsal margin of the foramen magnum by a short dorsal contact between the otoccipitals; however, the latter is
not preserved. The supraoccipital-otoccipital suture is apparent dorsal to the foramen magnum and further anterolaterally where the supraoccipital reaches the prootic and probably the laterosphenoid on either side (Figs. 12A, 13A).
The supraoccipital bears a clear sagittal crest with two depressions on its sides (scr, Fig. 11D). These depressions could correspond to the ventral border of the posttemporal fenestra (ptf? in Fig. 11D) and the course of the transverso-occipital vein; they are, however, too close to the sagittal plane compared to the position of the posttemporal fenestra in the holotype (Fig. 1F). Paired canals within the paroccipital processes, just ventral to the contact with the parietal and lateral to the suture with the supraoccipital, almost certainly transmitted vascular elements. These canals could have transmitted some tributaries of the dorsal head/transverso-occipital veins or the occipital artery (vf in Figs. 11D, 12B, 13B). The latter canals are absent in the holotype ZIN PH 1/16; thus, the arrangement of vascular foramina on the occipital surface is variable among the specimens assigned to Bissektipelta. Small paired foramina for the supraoc-cipital vein are present at the suture between the supra-occipital and otoccipital (fvSo, Fig. 11D).
Lateral braincase wall. In general, the structure of the lateral wall of the braincase of ZIN PH 281/16, including the distribution of the neurovascular foramina, closely matches that of the holotype. The neurovascular foramina are grouped into anterior and posterior clusters divided by a flattened crista prootica.
The otoccipital forms most of the posterior aspect of the lateral wall of the braincase and encloses much of the posterior cluster of foramina: the undivided metotic foramen (MF), two or three external foramina of the CN XII (varying between the two sides of the specimen), and partly the fenestra ovalis (FO) and the columellar recess (Fig. 12). These openings are incised ventral to the broad base of the paroccipital process. A pair of grooves begins from the FO and MF and extends distally on the ventral surface of the paroccipital process. The openings for CN XII and the MF are completely enclosed by the otoccipital; the basioccipital is apparently excluded from the external ventral border of the MF (Fig. 12E). The MF is separated from the foramina for CN XII by a laminalike process of the otoccipital that descends anteroven-trally toward the basal tuber. The metotic foramen is separated from the anteriorly situated FO by an oblique crista interfenestralis (= ventral ramus of opisthotic in more basal archosauriforms). The crista interfenestralis is a minor process that is not visible in occipital view. The columellar recess and the FO are bounded by the prootic anteriorly and by the otoccipital posteriorly, which is a common condition for diapsids in general (e.g., Sobral and Müller, 2016). The columellar recess is more open laterally that in the holotype ZIN PH 1/16. It leads into
Fig. 10. ZIN PH 281/16, referred specimen of Bissektipelta archibaldifrom the Bissekty Formation (Turonian), Uzbekistan. Photographs and corresponding CT-based models in dorsal (A, B) and ventral (C, D) views. Scale bars each equal 1 cm. Sutures are represented by solid lines; possible sutures are represented by dashed lines. Abbreviations: bo, basioccipital; bpt, basipterygoid process; cap, capitate process; CN II — XII, cranial nerve foramina; fvOC, foramen for orbitocerebral vein; hypc, hypophyseal cavity; ls, laterosphenoid; MF, metotic foramen; ors, orbitosphenoid; oto, otoccipital; pbs, parabasisphenoid; pop, paroccipital process; pro, prootic; so, supraoccipital; speth, sphenethmoid; vf, vascular foramen.
the FO; the latter communicates medially with the vestibular recess of the inner ear via a foramen (Fig. 13C).
The prootic forms the posterior border of the foramen for CN V, encloses the foramen for CN VII, and partially bounds the columellar recesss/FO (Fig. 12A, C, E). It broadly adheres to the anterior surface of the par-occipital process, as in various archosauriforms except crocodylomorphs (character 105 in Nesbitt [2011]). The anterior contact of the prootic with the laterosphenoid is evident on both sides of the specimen. The prootic-supraoccipital contact is not clearly observable, and the ventral contacts of the prootic with the parabasisphenoid
are obliterated. The prootic forms a triangular projection that descends from the dorsal margin of the foramen for
CN V and partially subdivides it.
The suture between the laterosphenoid and prootic and a prominent capitate process (cap in Figs. 11B, 12B) mark the posterior extent of the laterosphenoid. The latter participates in the anterior margin of the foramen for CN V. It is likely that the laterosphenoid also encloses the orbitocerebral vein openings and the foramen for CN IV. A pair of foramina for the orbitocerebral veins is present on both laterosphenoids; additionally, a groove at the presumed laterosphenoid-frontal contact on both
Fig. 11. ZIN PH 281/16, referred specimen of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. Photographs and corresponding CT-based models in anterior (A, B) and posterior/occipital (C, D) views. Scale bars each equal 1 cm. Sutures are represented by solid lines; possible sutures are represented by dashed lines. Abbreviations: ameth, articular surface for mesethmoid; bo, basioccipital; bpt, basipterygoid process; bt, basal tuber; cap, capitate process; CN V, trigeminal cranial nerve foramen; oc, occipital condyle; cvn?, cavity for vomero-nasal bulb?; fCC, cerebral carotid artery and vein foramen; fm, foramen magnum; fvSo, supraoccipital vein foramen; ls, laterosphe-noid; oto, otoccipital; pbs, parabasisphenoid; pbsro-ios, fused parabasisphenoid rostrum and interorbital septum; pop, paroccipital process; proaf, proatlas facet; ptf?, posttemporal fenestra?; scr, sagittal crest; so, supraoccipital; speth, sphenethmoid; vf, vascular foramen.
sides of the specimen marks the course of a similar vascular element (fvOC in Fig. 12D). An internal vascular canal for the supraorbital vessels was reconstructed for the holotype at this region; the canal is absent in ZIN PH 281/16, indicating a lesser degree of ossification of the braincase wall. The capitate process is stout and bears a rounded head with an unfinished articular surface (Fig. 12A). The facet on its dorsal surface and round head indicate a synovial joint between the laterosphe-noid and the postorbital (Holliday and Witmer, 2008) that was nevertheless akinetic, as in extant crocodylians. The blunt crista antotica (Sampson and Witmer, 2007, and references therein; laterosphenoid buttress in Hol-liday and Witmer [2009]) descends from the capitate process and subdivides the orbital and adductor aspects of the external surface of the laterosphenoid (Fig. 12B,
D). On the left side of ZIN PH 281/16, a groove passes through the crista antotica. It likely indicates the course of the temporoorbital artery/vein (gTO in Fig. 12D).
Sutures cannot be distinguished between the preserved elements of the sphenethmoidal complex and between them and the parabasisphenoid. The medial septum that separated the olfactory bulbs is broken off in ZIN PH 281/16, but the preserved surface is symmetrical on both sides and most likely represents a facet (ameth, Fig. 11A). Thus, this medial septum was a separate element (mesethmoid in Miyashita et al. [2011]). It contacted the elements of the lateral wall of the braincase laterally, the parabasisphenoid ventrally, and the skull roof dorsally (based on the holotype that preserves both the medial septum and the skull roof) and was probably continuous anteriorly with the ossified nasal septum as
CR+FO CN VII bP*
bt
faCC
Fig. 12. ZIN PH 281/16, referred specimen of Bissektipelta archibaldifrom the Bissekty Formation (Turonian), Uzbekistan. Photographs and corresponding CT-based models with cranial endocast in right lateral (A, B) and oblique left lateral (C, D) views, with a close-up of the posterior cranial nerve foramina (E). Scale bars each equal 1 cm; E not to scale. Sutures are represented by solid lines; possible sutures are represented by dashed lines. Abbreviations: bo, basioccipital; bpt, basipterygoid process; bt, basal tuber; cap, capitate process; CN II — XII, cranial nerve foramina; cvn?, cavity for vomero-nasal bulb?; faCC, cerebral carotid artery and vein foramen; faSP?, sphenopalatine artery and vein foramen?; CR+FO, columellar recess and fenestra ovalis; fvOC, foramen for orbitocerebral vein; gTO, temporoorbital artery and vein groove; ls, laterosphenoid; MF, metotic foramen; ors, orbitosphenoid; oto, otoccipital; pbs, parabasisphenoid; pro, prootic; so, supraoccipital; speth, sphenethmoid; vf, vascular foramen.
Fig. 13. ZIN PH 281/16, referred specimen of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. Photograph and corresponding CT-based model in oblique anterodorsal view (A, B) and parasagitally sectioned CT-based model showing left endocranial surface in medial view (C). Scale bars each equal 1 cm. Sutures are represented by solid lines; possible sutures are represented by dashed lines. Abbreviations: bo, basioccipital; cap, capitate process; cerc, cerebral cavity; CN II — XII, cranial nerve foramina; cvn?, cavity for vomero-nasal bulb?; dsOc, occipital venous sinus depression; fCC, cerebral carotid artery and vein foramen; floc, floccular fossa; fvMC, middle cerebral vein foramen; fvOC, foramen for orbitocerebral vein; lagf, lagenar fossa; Is, laterosphenoid; olfc, olfactory cavity; oto, otoccipital; pbs, parabasi-sphenoid; pro, prootic; so, supraoccipital; speth, sphenethmoid; ves, vestibular cavity; vf, vascular foramen.
in other ankylosaurs (Miyashita et al., 2011). Features of the complex external surface of the sphenethmoidal complex include prominent vertical striations dorsal to the foramen for CN II (possible site of attachment of the preorbital septum; ectethmoid in Miyashita et al. [2011]) and multiple grooves, ridges, and bumps around the foramen for CN II (indicating courses of neurovas-cular elements and possible attachment sites of ocular musculature).
The parabasisphenoid constitutes most of the an-teroventral aspect of the lateral surface of the braincase (Fig. 12A, C). Paired grooves mark the course of CN VI from the dorsum sellae toward the external surface of the braincase, bypassing the hypophyseal cavity (Figs. 12D, 13). The participation of the prootic in the canal for CN VI is not clear. A large foramen for the cerebral carotid artery is present on either side of the specimen (faCC in Fig. 12B, D). The breakage of the specimen anterior to the cerebral carotid foramen on both sides makes the
interpretation of certain foramina challenging. On the left side, the anterior rounded margin of a foramen is preserved (faSP?, Fig. 12D). This opening is comparable in size to the foramen for the cerebral carotid artery. Dorsal to it, the bone is broken, and no additional foramina could be identified. On the right side, the margin of a small-sized foramen is preserved anterior to the foramen for CN VI (faSP?, Fig. 12B). It is half as large as the abovementioned foramen on the left side but is located at the same level as the latter. Considering the position of these foramina, both of them could equally likely represent foramina for the sphenopalatine artery or foramina for CN III. In the latter case, the sphenopal-atine artery would have branched off from the cerebral carotid artery before the latter entered the hypophyseal cavity in ZIN PH 281/16. However, as the nature of foramen for CN III is controversial in the holotype (see above), we assume that these foramina represent exits of the sphenopalatine artery (faSP?, Fig. 12B, D). Thus, the
Fig. 14. ZIN PH 281/16, referred specimen of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. Cranial endocast with endosseous labyrinth of the inner ear in right lateral (A), ventral (B), dorsal (C), and oblique ventrolateral (D) views. Scale bars each equal 1 cm. Abbreviations: aCC, cerebral carotid artery and vein; aCE/vCC?, caudal encephalic artery/caudoventral cerebral vein; aSP?, sphenopalatine artery and vein; cbl, cerebellum; ch, cerebral hemisphere; CN II — XII, cranial nerves; cochd, endosseous cochlear duct; hyp, hypophysis (pituitary); lab, endosseous labyrinth; MF, metotic passage; mo, medulla oblongata; olfb, olfactory bulb; olft, olfactory tract; pls, perilymphatic sac; vCD, dorsal heard vein; vn?, vomero-nasal bulbs?; vOC, orbitocerebral veins.
presence and position of separate foramina for CN III in ZIN PH 281/16 are uncertain.
Endocranial surface. The endocranial surface of ZIN PH 281/16 does not differ significantly from that of the holotype in the general division into regions or the distribution of neurovascular foramina (Fig. 13). Its surface is mostly smooth, indicating a loose infilling by the brain. Vertical striations at the surface of the cavity for the olfactory tract probably result from a closer contact between the brain and its dura with the walls of the brain-case (Fig. 13A). Most sutures on the endocranial surface are obliterated; however, the basioccipital-otoccipital, basioccipital-parabasisphenoid, and prootic-otoccipital sutures are visible (Fig. 13A). The straight basioccipital-otoccipital sutures extend ventral to the internal foramina for CN XII and disappear towards the MF and la-genar fossae. The suture between the basioccipital and parabasisphenoid extends transversally between large, unossified lagenar fossae. The basioccipital apparently forms most of the ventral endocranial surface. Possible
prootic-otoccipital sutures extend as ridges on the preserved surface of both otic capsules.
A single transverse canal for CN II opens laterally on either side in a separate foramen (Fig. 13C). The hypophyseal cavity is comparatively shallow, nearly half the dorsoventral depth of the cerebral cavity above it. The tentorial crest separating the cerebral and cerebellar cavities appears to be prominent and sharp at its dorsal part; it is broken off ventrally on both sides and the canal for CN VI is exposed (Fig. 13B). The dorsum sellae preserves the central triangular projection. Posterodor-sally in the prootic, a foramen leads into two canals, one anteroventral (for CN VII) and the other posterodorsal into the otic capsule (for CN VIII) (Fig. 13C). The medial walls of both otic capsules are largely unossified, and two huge lagenar fossae are present on the floor of the endo-cranium (ves+lagf in Fig. 13B). Anterodorsal to the otic capsule, a distinct fossa is present on either side, which is not particularly evident in the holotype (floc, Fig. 13C). These depressions correspond in their position to the
floccular (auricular) fossae in other archosaurs (Gower, 2002; Sampson and Witmer, 2007; Witmer et al., 2008; Sobral et al., 2016). Each fossa has a pair of sediment-filled openings at the bottom that apparently connect to the inner ear cavities within the bone (the CT data offer insufficient resolution to trace these structures with confidence). These foramina likely transmitted vessels to the inner ear labyrinth. Although the external foramina for CN XII vary in number (two or three) between both sides of the specimen, there are three internal openings on either side (Fig. 13C).
Endocast. The digital
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https://dinopedia.fandom.com/wiki/Zhejiangosaurus
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Zhejiangosaurus
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Zhejiangosaurus (meaning "Zhejiang lizard") is an extinct genus of nodosaurid dinosaur from the Upper Cretaceous (Cenomanian stage) of Zhejiang, eastern China. It was first named by a group of Chinese and Japanese authors Junchang Lü, Xingsheng Jin, Yiming Sheng and Yihong Li in 2007 and the...
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https://static.wikia.nocookie.net/dinosaurs/images/4/4a/Site-favicon.ico/revision/latest?cb=20240402071814
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Dinopedia
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https://dinopedia.fandom.com/wiki/Zhejiangosaurus
|
Zhejiangosaurus (meaning "Zhejiang lizard") is an extinct genus of nodosaurid dinosaur from the Upper Cretaceous (Cenomanian stage) of Zhejiang, eastern China. It was first named by a group of Chinese and Japanese authors Junchang Lü, Xingsheng Jin, Yiming Sheng and Yihong Li in 2007 and the type species is Zhejiangosaurus lishuiensis ("from Lishui", Chinese administrative unit on which the fossil was found).[1]
Description[]
Zhejiangosaurus could grow up to 4.5 m (17 ft) in length and was 1.4 metric tons in weigh.[3]
Material[]
Material for Zhejiangosaurus consists of the holotype, ZNHM M8718, a partial skeleton which has preserved a sacrum with eight vertebrae, a complete right ilium and partial left ilium, a complete right pubis, the proximal end of the right ischium, two complete hindlimbs, fourteen caudal vertebrae, and some unidentified bones. These remains come from Liancheng, in the Chinese administrative unit of Lishui on the province of Zhejiang and they were collected from the Cenomanian-age Chaochuan Formation.[1]
Systematics[]
On the species description, Lü et al. (2007) found Zhejiangosaurus to belong to the ankylosaurian family Nodosauridae.[1] It is the only known nodosaurid from As
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https://alchetron.com/Zhejiangosaurus
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Alchetron, The Free Social Encyclopedia
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2017-08-18T08:30:48+00:00
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Zhejiangosaurus (meaning Zhejiang lizard) is an extinct genus of nodosaurid dinosaur from the Upper Cretaceous (Cenomanian stage) of Zhejiang, eastern China. It was first named by a group of Chinese and Japanese authors Junchang L, Xingsheng Jin, Yiming Sheng and Yihong Li in 2007 and the type sp
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en
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/favicon.ico
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Alchetron.com
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https://alchetron.com/Zhejiangosaurus
|
Zhejiangosaurus (meaning "Zhejiang lizard") is an extinct genus of nodosaurid dinosaur from the Upper Cretaceous (Cenomanian stage) of Zhejiang, eastern China. It was first named by a group of Chinese and Japanese authors Junchang Lü, Xingsheng Jin, Yiming Sheng and Yihong Li in 2007 and the type species is Zhejiangosaurus lishuiensis ("from Lishui", Chinese administrative unit on which the fossil was found). It has no diagnostic features, and thus is a nomen dubium.
Contents
Material
Systematics
References
Material
Material for Zhejiangosaurus consists of the holotype, ZNHM M8718, a partial skeleton which has preserved a sacrum with eight vertebrae, a complete right ilium and partial left ilium, a complete right pubis, the proximal end of the right ischium, two complete hindlimbs, fourteen caudal vertebrae, and some unidentified bones. These remains come from Liancheng, in the Chinese administrative unit of Lishui on the province of Zhejiang and they were collected from the Cenomanian-age Chaochuan Formation.
Systematics
On the species description, Lü et al. (2007) found Zhejiangosaurus to belong to the ankylosaurian family Nodosauridae.
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https://www.wikiwand.com/en/Talarurus
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en
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Talarurus
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Talarurus is a genus of ankylosaurid dinosaur that lived in Asia during the Late Cretaceous period, about 96 million to 89 million years ago. The first remains of Talarurus were discovered in 1948 and later described by the Russian paleontologist Evgeny Maleev with the type species T. plicatospineus. It is known from multiple yet sparse specimens, making it one of the most well known ankylosaurines, along with Pinacosaurus. Elements from the specimens consists of various bones from the body; five skulls have been discovered and assigned to the genus, although the first two were very fragmented.
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en
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Wikiwand
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https://www.wikiwand.com/en/Talarurus
|
Talarurus ( TAL-ə-ROOR-əs; meaning "basket tail" or "wicker tail") is a genus of ankylosaurid dinosaur that lived in Asia during the Late Cretaceous period, about 96 million to 89 million years ago. The first remains of Talarurus were discovered in 1948 and later described by the Russian paleontologist Evgeny Maleev with the type species T. plicatospineus. It is known from multiple yet sparse specimens, making it one of the most well known ankylosaurines, along with Pinacosaurus. Elements from the specimens consists of various bones from the body; five skulls have been discovered and assigned to the genus, although the first two were very fragmented.
It was a medium-sized, heavily built, ground-dwelling, quadrupedal herbivore, that could grow up to 5–6 m (16–20 ft) long and weighed about 454 to 907 kg (1,001 to 2,000 lb), nearly a ton. Like other ankylosaurs it had heavy armour and a club on its tail, limiting its speed. Talarurus is classified as a member of the Ankylosauria, in the Ankylosaurinae, a group of derived ankylosaurs. Talarurus is known from the Bayan Shireh Formation, being likely niche partitioned with Tsagantegia, as indicated by its muzzle, which has a rectangular shape specialized for grazing. These represent the oldest known ankylosaurines from Asia, although they are not very closely related to each other. It appears that the closest relative of Talarurus was Nodocephalosaurus, an ankylosaurin with similar facial osteoderms.
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https://copylists.com/animals/list-of-dinosaurs/
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PDF Excel - CopyLists.com
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2021-12-19T05:22:49+00:00
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Copy or download a list of dinosaurs in popular formats.Aardonyx, Abelisaurus, Abrictosaurus, Abrosaurus, Abydosaurus, Acanthopholis, Achelousaurus
|
en
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CopyLists.com
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https://copylists.com/animals/list-of-dinosaurs/
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Copy or download this list of Dinosaurs A-Z in many popular formats. Dinosaurs are one of the most popular topics in the world. They have been a fascination for people for centuries and continue to be so today. There are many different types of dinosaurs, but they all share some common traits. Dinosaurs were reptiles that lived on Earth from about 230 million years ago to 65 million years ago. They were the dominant species on Earth for over 160 million years and then went extinct.
Paleontology is the study of the age of dinosaurs and other prehistoric life. It is a field of science that deals with fossils and all other evidence of the earth’s natural history. The word paleontology comes from the Greek words “palaio” which means old, and “onitis” which means study.
People are fascinated by dinosaurs because they offer a window into the past. They help us understand how life came to be and how it evolved. Also, they are great giant real monsters and just really cool!
17
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This is the list read by a female voice in mp3 format. Click the 3 dots on the left of the player to download the mp3 file or change the speed of the playback.
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dbpedia
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https://www.rareresource.com/Zhejiangosaurus-dinosaur.html
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Zhejiangosaurus Dinosaur, facts
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Zhejiangosaurus Dinosaur
Zhejiangosaurus (phonetically spelled as Zay-jee-ang-o-sore-us) is a Lishuiensis type species named by Junchang Lu, Xingsheng Jin, Yiming Sheng & Yihong Li in 2007. The classification of the species is as follows:
Chordata â Reptilia â Dinosauria â Ornithischia â Thyreophora â Ankylosauria â Nodosauridae
The size of the Zhejiangosaurus is unknown and they followed an herbivorous diet. They existed during the Cenomanian period of the Cretaceous era. The fossil representation includes a partial skeleton including vertebrae, right ilium, partial left ilium, right pubis, end of the ischium and both hind legs.
Zhejongosaurus was first named by a group of Chinese and Japanese authors Junchang Lü, Xingsheng Jin, Yiming Sheng and Yihong Li in 2007. The type species is Zhejiangosaurus lishuiensis named from Lishui, a Chinese administrative unit on which the fossil was found. Zhejiangosaurus is an extinct genus of nodosaurid dinosaur from the Cenomanian stage of the Upper Cretaceous of Zhejiang, eastern China.
The remains of the holotype of Zhejiangosaurus come from Liancheng, in the Chinese administrative unit of Lishui on the province of Zhejiang and they were collected from the Cenomanian-age Chaochuan Formation. The holotype of this species is ZNHM M8718 which consists of a partial skeleton which has preserved a sacrum with eight vertebrae, a complete right ilium and partial left ilium, a complete right pubis, the proximal end of the right ischium, two complete hindlimbs, fourteen caudal vertebrae, and some unidentified bones.
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dbpedia
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https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8667728/
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en
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The phylogenetic nomenclature of ornithischian dinosaurs
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[
"Daniel Madzia",
"Victoria M. Arbour",
"Clint A. Boyd",
"Andrew A. Farke",
"Penélope Cruzado-Caballero",
"David C. Evans"
] |
2021-08-25T00:00:00
|
Ornithischians form a large clade of globally distributed Mesozoic dinosaurs, and represent one of their three major radiations. Throughout their evolutionary history, exceeding 134 million years, ornithischians evolved considerable morphological disparity, ...
|
en
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https://www.ncbi.nlm.nih.gov/coreutils/nwds/img/favicons/favicon.ico
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PubMed Central (PMC)
|
https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8667728/
|
Phylogenetic Nomenclature of Ornithischian Clades
For the sake of clarity, all clade names are provided in alphabetical order. The definitions are summarized in . The extent of all clade names is further depicted on that shows the relationships of taxa included in the present study as specifiers (both, internal as well as external) and additionally on – that represent selected ornithischian-wide phylogenies published within recent years: Madzia, Boyd & Mazuch (2018: Fig. 4B), Dieudonné et al. (2020: Figs. 1 and 2), and Yang et al. (2020: Fig. 12).
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About: Ankylosaurus
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Ankylosaurus is a genus of armored dinosaur. Its fossils have been found in geological formations dating to the very end of the Cretaceous Period, about 68–66 million years ago, in western North America, making it among the last of the non-avian dinosaurs. It was named by Barnum Brown in 1908; it is monotypic, containing only A. magniventris. The generic name means "fused lizard", and the specific name means "great belly". A handful of specimens have been excavated to date, but a complete skeleton has not been discovered. Though other members of Ankylosauria are represented by more extensive fossil material, Ankylosaurus is often considered the archetypal member of its group, despite having some unusual features.
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http://dbpedia.org/resource/Ankylosaurus
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dbo:abstract
الأنكيلوصور أو العظاءة الملتحمة أو العظاءة المنصهرة (بالإنجليزية: Ankylosaurus) هي جنس منقرض من العظاءات حاملات الدروع. ويحتوي على نوع واحد، هو (A. magniventris). تم العثور على أحافير الأنكيلوصور على هيئة ترجِع إلى أواخر نهايةالعصر الطباشيري فترة (علم الجيولوجيا) (منذ حوالي من 65.5 إلى 66.5 مليون سنة) بِغرب أمريكا الشمالية. وعلى الرغم من عدم اكتشاف هيكل عظمي كامل للأنكيلوصور مع تميز ديناصورات عديدة أخرى بمواد أحفورية شاملة، إلا أنه يُعتَبَر الديناصور دِرع (علم الحيوان) . وتشترك الديناصورات الأخرى التي تنتمي إلى فصيلة (Ankylosauridae) في ملامحها المعروفة وهي—الجسد الذي يحتوي على درع ثقيل والذيل العظمي الضخم—إلا أن الأنكيلوصور كان أكبر الأعضاء المعروفة في العائلة. (ar)
Ankylosaurus („Neohebný/srostlý ještěr“) byl rod ptakopánvého dinosaura, jež zahrnuje druh (A. magniventris). Fosilie tohoto mohutného obrněného býložravce jsou známy z nejmladších vrstev křídového útvaru (věk maastricht, asi před 68 až 66 miliony let). Ankylosaurus byl popsán v roce 1908 paleontologem Barnumem Brownem. Fosilie tohoto rodu byly dosud objeveny v souvrství Hell Creek, souvrství Lance, souvrství Scollard, souvrství Frenchman a v souvrství Ferris. (cs)
L'anquilosaure (Ankylosaurus magniventris) és una espècie de dinosaure ornitisqui de la família dels anquilosàurids, l'única del gènere Ankylosaurus. Se n'han trobat restes fòssils en formacions geològiques de l'oest de Nord-amèrica que daten del límit superior del Cretaci, fa 66-68 milions d'anys. Fou, doncs, un dels últims dinosaures no aviaris. Cap dels espècimens que se n'han descobert és un esquelet complet. Sovint és considerat l'arquetip de l'infraordre Ankylosauria malgrat que n'hi ha altres espècies que estan representades per una major quantitat de fòssils i que l'anquilosaure presenta diversos caràcters inusuals en aquest grup. Altres anquilosaures comparteixen els seus ben coneguts trets de cos fortament protegit amb llarga cua òssia amb forma de garrot, però l'anquilosaure és el membre més conegut de la família. (ca)
Οι αγκυλόσαυροι (Ankylosaurus) είναι γένος δεινοσαύρων, το οποίο έζησε στο ύστερο Κρητιδικό, πριν 66,5 με 65,5 εκατομμύρια χρόνια στην Βόρεια Αμερική. Ήταν μεγάλοι φυτοφάγοι θωρακισμένοι δεινόσαυροι. Αν και δεν έχει βρεθεί κάποιος ολοκληρωμένος σκελετός του, ο αγκυλόσαυρος θεωρείται ο αρχέτυπος θωρακισμένος δεινόσαυρος. Σύμφωνα με εκτιμήσεις, ο αγκυλόσαυρος έφτανε σε μήκος τα 8 με 9 μέτρα και βάρος 4-7 τόνους. Μια εκ νέου περιγραφή με βάση το μεγαλύτερο κρανίο που έχει βρεθεί τοποθετεί το μέγιστο μήκος του σε 6,25 μέτρα. Το πιο εμφανές χαρακτηριστικό του αγκυλόσαυρου είναι η πανοπλία του, η οποία κάλυπτε το σώμα του δεινόσαυρου και αποτελείτο από ακίδες και οστέινες πλάκες. Ακόμη και τα βλέφαρά του είχαν οστέινες προστατευτικές πλάκες. Επίσης είχαν και ένα οστέινο ρόπαλο στην άκρη της ουράς τους για περισσότερη προστασία που ζύγιζε 30 κ. Εξαιτίας του μεγέθους και της πανοπλίας του ο αγκυλόσαυρος ήταν μια δύσκολη λεία. (el)
Ankylosaurus is a genus of armored dinosaur. Its fossils have been found in geological formations dating to the very end of the Cretaceous Period, about 68–66 million years ago, in western North America, making it among the last of the non-avian dinosaurs. It was named by Barnum Brown in 1908; it is monotypic, containing only A. magniventris. The generic name means "fused lizard", and the specific name means "great belly". A handful of specimens have been excavated to date, but a complete skeleton has not been discovered. Though other members of Ankylosauria are represented by more extensive fossil material, Ankylosaurus is often considered the archetypal member of its group, despite having some unusual features. Possibly the largest-known ankylosaurid, Ankylosaurus is estimated to have been between 6 and 8 meters (20 and 26 ft) long and to have weighed between 4.8 and 8 metric tons (5.3 and 8.8 short tons). It was quadrupedal, with a broad, robust body. It had a wide, low skull, with two horns pointing backward from the back of the head, and two horns below these that pointed backward and down. Unlike other ankylosaurs, its nostrils faced sideways rather than towards the front. The front part of the jaws was covered in a beak, with rows of small, leaf-shaped teeth farther behind it. It was covered in armor plates, or osteoderms, with bony half-rings covering the neck, and had a large club on the end of its tail. Bones in the skull and other parts of the body were fused, increasing their strength, and this feature is the source of the genus name. Ankylosaurus is a member of the family Ankylosauridae, and its closest relatives appear to be Anodontosaurus and Euoplocephalus. Ankylosaurus is thought to have been a slow-moving animal, able to make quick movements when necessary. Its broad muzzle indicates it was a non-selective browser. Sinuses and nasal chambers in the snout may have been for heat and water balance or may have played a role in vocalization. The tail club is thought to have been used in defense against predators or in intraspecific combat. Specimens of Ankylosaurus have been found in the Hell Creek, Lance, Scollard, Frenchman, and Ferris formations, but appears to have been rare in its environment. Although it lived alongside a nodosaurid ankylosaur, their ranges and ecological niches do not appear to have overlapped, and Ankylosaurus may have inhabited upland areas. Ankylosaurus also lived alongside dinosaurs such as Tyrannosaurus, Triceratops, and Edmontosaurus. (en)
Ankylosaurus ist eine Gattung aus der Gruppe der Vogelbeckensaurier (Ornithischia) aus der Oberkreide Nordamerikas. Er ist Namensgeber und gleichzeitig einer der größten und jüngsten Vertreter der Ankylosauria. (de)
Ankylosaurus (antzinako grezieraz "narrasti fusionatua" edo "narrasti zurruna") dinosauro genero bat da. Kretazeo Berantiarrean bizi izan zen, Maastrichtiar estaian, duela 66-68 milioi urte artean. Ipar Amerikaren mendebaldean aurkitu dira fosilak. Generoko espezie bakarra Ankylosaurus magniventris da, 1908an deskribatu zuena. Ordutik ale gutxi batzuk bildu dira baina ez da hezurdura osorik aurkitu oraindik. Gorputz sendoa zuen armadura sendo batez babestua. Osteodermo izeneko plakek osatzen zuten armadura hori. Zortzi eta hamar metro arteko luzera zuen eta lau hankatan ibiltzen zen. Isatsaren muturrean mailu handi bat zeukan. Buruko hezurrak fusionaturik zituen, bai eta gorputzeko hainbat ataletako hezurrak ere, eta horrek erresistentzia handitzen zion ankylosauroari. Ezaugarri honetatik datorkio izena. (eu)
Ankylosaurus magniventris (en gr. «lagarto acorazado de vientre grande») es la única especie conocida del género fósil Ankylosaurus de dinosaurios tireofóros anquilosáuridos, que vivió a finales del período Cretácico, hace aproximadamente 68 a 66 millones de años, durante el Maastrichtiense, en lo que hoy es Norteamérica.Al igual que otros anquilosáuridos, Ankylosaurus se distinguía por su pesada y un gran mazo caudal, siendo probablemente el más grande de su grupo. Aunque hace falta descubrir esqueletos completos y algunos de sus parientes poseen muchos más fósiles recobrados, Ankylosaurus es considerado el dinosaurio acorazado más destacado y representativo. Siendo el anquilosáurido conocido más grande, Ankylosaurus medía hasta 6,25 m de largo, 1,7 de altura, y pesaba 6 toneladas. Fue un animal cuadrúpedo, con un cuerpo amplio y robusto. Tenía un cráneo grande y ancho, con dos cuernos apuntando hacia atrás desde la parte posterior de la cabeza, y dos cuernos por debajo de estos que apuntaban hacia atrás y hacia abajo. La parte delantera del rostro estaba cubierto de un pico, con hileras de dientes pequeños, en forma de hoja más detrás de él. Estaba cubierto de placas de armadura u osteodermos, con medios anillos óseos que cubren el cuello, y tenía un gran mazo en el extremo de su cola. Huesos en el cráneo y otras partes del cuerpo se fusionaron, lo que aumentaba su resistencia, y esta característica es la fuente del nombre del género. Ankylosaurus es un miembro de la familia Ankylosauridae y sus parientes más cercanos parece ser Anodontosaurus y Euoplocephalus. Ankylosaurus se cree que ha sido un animal lento, capaz de hacer movimientos rápidos cuando sea necesario. Su hocico ancho indica que realizaba un pastoreo no selectivo. Los senos paranasales y fosas nasales en el hocico pueden haber sido para el intercambio de calor y agua o haya desempeñado un papel en la vocalización. La porra de la cola se cree que fue utilizada para la defensa contra los depredadores o en el combate intraespecífico. Ankylosaurus se ha encontrado en las formaciones Hell Creek, lance y , pero parecen haber sido poco frecuentes en su entorno. A pesar de que vivía junto a un nodosáurido, sus rangos y nichos ecológicos no parecen haber solapado y Ankylosaurus pueden haber vivido en zonas de montaña. Ankylosaurus vivió junto con dinosaurios como el Tyrannosaurus, Triceratops y Edmontosaurus. (es)
Dineasár beag ceathairchosach, armúrtha le plátaí dronuilleogacha cnámhacha ar feadh na colainne is an eireabaill. An ceann beag, na fiacla laghdaithe nó as láthair. D'itheadh sé fásra. Faightear iarsmaí de ón tréimhse Chailceach i Meiriceá Thuaidh. (ga)
Ankylosaurus magniventris • Ankylosaures Ankylosaurus Reconstitution d'un Ankylosaurus. Genre † AnkylosaurusBrown, 1908 Espèce † Ankylosaurus magniventrisBrown, 1908 Ankylosaurus, les ankylosaures en français, littéralement « lézard rigide », est un genre éteint de dinosaures ornithischiens herbivores de l'infra-ordre des Ankylosauria et de la famille des Ankylosauridae. Des fossiles d'Ankylosaurus ont été découverts en Amérique du Nord dans les sédiments de la fin du Crétacé supérieur (Maastrichtien), soit il y a environ entre 68 et 66 millions d'années. On a aussi trouvé un grand nombre de fossiles à l'est de la Hongrie, des traces de pas en Bolivie. C'est un des dinosaures qui a disparu lors de la grande extinction de la fin du Crétacé, survenue il y a 66 millions d'années. Une seule espèce est rattachée au genre : Ankylosaurus magniventris, décrite par Barnum Brown en 1908. Bien qu'aucun squelette complet n'ait été trouvé et que plusieurs autres espèces disposent de plus de fossiles, Ankylosaurus est souvent considéré comme l'archétype du dinosaure à armure (les thyréophores). Il a donné son nom à la famille et à l'infra-ordre auxquels il appartient, respectivement les ankylosauridés et les ankylosauriens. D'autres ankylosauridés partageaient ses caractéristiques, dont une queue en forme de massue, mais Ankylosaurus était le plus grand membre de sa famille. (fr)
Ankylosaurus adalah salah satu jenis dinosaurus yang hidup pada periode kapur akhir sekitar 68 juta hingga 65 juta tahun yang lalu di Amerika Utara. Memiliki tubuh sepanjang 8 meter ( 26 kaki), tingginya sekitar 2 meter ( 6.6 kaki), dan beratnya mencapai setidaknya 4 ton. Ankylosaurus memiliki tubuh yang dilindungi oleh semacam cangkang keras yang membuat tubuhnya tidak bisa diserang dengan mudah, bahkan oleh Tyrannosaurus rex. Perisai tulang setebal 5 cm melindungi tubuhnya yang pendek, yang disangga oleh empat kakinya yang kuat, pendek dan gemuk. Di ekornya terdapat bola dari batu seberat 5 kg. Jika Ankylosaurus diadang oleh lawannya, ia akan menyerang lawan tersebut dengan ekor kerasnya dan dalam sekejap lawannya akan terjatuh. Para ilmuwan dan ahli palaeontologi biasanya menyebut Ankylosaurus dengan sebutan 'Anky kecil'. (in)
Ankylosaurus (Brown, 1908 - il cui nome dal greco significa letteralmente "lucertola fusa") è un genere estinto di dinosauro ornitischio tireoforo vissuto nel Cretaceo superiore, circa 70-65 milioni di anni fa (Maastrichtiano), in quella che oggi è il Nord America, il che lo rende uno degli ultimi dinosauri a comparire prima della grande estinzione dei dinosauri. Ankylosaurus fu descritto per la prima volta nel 1908, da parte del paleontologo Barnum Brown, e la specie tipo, A. magniventris, è l'unica specie ascritta al genere Ankylosaurus. Dalla sua scoperta, sono stati scoperti solo una manciata di esemplari frammentari, ma non è mai stato ritrovato uno scheletro completo. Anche se la maggior parte dei membri di Ankylosauria sono rappresentati da materiali fossili ben più completi, Ankylosaurus è spesso considerato come l'archetipo del suo gruppo. Con una lunghezza massima di 6,25 m (20,5 piedi), un'altezza di 1,7 m (5,6 piedi) e un peso di 6 tonnellate (13.000 lb), nonostante lo scarso materiale fossile, Ankylosaurus è ancora oggi considerato il genere più grande degli Ankylosauria. Il suo corpo ampio e robusto era sorretto da quattro corte ma robuste zampe, ed era ricoperto da una spessa corazza di osteodermi, disposti ad anelli che circondavano anche la regione del collo. La sua lunga coda corazzata finiva di una grossa mazza caudale fatta d'ossa fuse. La sua testa larga e bassa, disponeva di due piccole corna che diramavano all'indietro e un altro paio della stessa forma ma che invece puntavano verso il basso. La parte anteriore delle mascelle erano coperte in un becco corneo, dotato di una fila di piccoli denti a forma di foglia. Le ossa del cranio erano fuse, il che dava maggiore resistenza alle ossa, tale caratteristica dà anche il nome al genere. Ankylosaurus è un membro della famiglia degli Ankylosauridae, e i suoi parenti più stretti sembrano essere Anodontosaurus ed Euoplocephalus. A causa della sua imponente corazza, l'Ankylosaurus non doveva essere un animale molto veloce, quindi si pensa che fosse un animale lento, in grado di fare movimenti rapidi e scattanti solo in caso di necessità. Il suo muso largo e appiattito indica che era un erbivoro, poco selettivo in grado di masticare e strappare qualunque tipo di vegetale. I seni e le cavità nasali erano eccezionalmente sviluppati il che indica uno sviluppatissimo senso dell'olfatto, oltre ad aver fatto supporre ai paleontologi che l'animale potesse emettere vari tipi di vocalizzazione. La grande mazza caudale posta alla fine della coda di Ankylosarus era probabilmente usata dall'animale in vita come arma di difesa e offesa contro i predatori o nei combattimenti intraspecifici. I fossili di questo animale sono stati ritrovati nelle formazioni Hell Creek, Lance e Scollard, ma in tutte le formazione sembra essere stato un animale piuttosto raro nel suo habitat. Anche se ha vissuto accanto ad un altro anchilosauride Edmontonia, pare che i due animali non siano mai entrati in competizione in quanto occupavano nicchie ecologiche differenti, infatti, l'anchilosauro sembra abbia abitato le zone più montane. Questo animale visse negli stessi luoghi e nello stesso periodo in cui vissero altri giganti come Tyrannosaurus, Triceratops e Edmontosaurus. (it)
アンキロサウルス (Ankylosaurus) は、中生代白亜紀後期（約6,800万年前 - 6,600万年前）の現北アメリカ大陸に生息した植物食恐竜の属の一つ。鳥盤目 - 曲竜下目 -アンキロサウルス科に属する。属名は「連結したトカゲ」の意。 (ja)
안킬로사우루스(Ankylosaurus)는 백악기 후기(6800만년 전~6550만년 전) 에 살았던 공룡이다. 미국, 캐나다, 호주 등지에서 화석이 발견되고 있다. 갑옷과 같은 등껍질과 꼬리에 있는 곤봉으로 육식 공룡의 공격으로부터 자신을 보호하였다. 안킬로사우루스의 몸길이는 최대 7m에 몸무게 6톤에 달했다. 안킬로사우루스는 최후의 곡룡류이며 ‘ 도마뱀’ 또는 또는 이라는 뜻이다. 무엇보다 딱딱한 융합된 골편이 몸으로 뒤덮었고 가장 강력한 방어 무기인 꼬리 곤봉도 달려있다. 안킬로사우루스는 배에 골편이 없었기 때문에 육식공룡들은 배를 주로 공격했다. (ko)
Ankylosaurus is een geslacht van uitgestorven plantenetende ornithischische dinosauriërs, behorend tot de Thyreophora, dat tijdens het Laat-Krijt (Maastrichtien) leefde in het gebied van het huidige Noord-Amerika. De typesoort van het geslacht, tevens de enige soort, Ankylosaurus magniventris, werd in 1906 ontdekt en kreeg in 1908 zijn naam die 'gebogen sauriër met de grote buik' betekent. Daarna is er nog een klein aantal fossielen van de soort gevonden. Ze zijn allemaal onvolledig zodat maar een gedeelte van het skelet bekend is. Door te kijken naar andere Ankylosauridae, de groep waartoe Ankylosaurus behoort, kunnen we ons een beeld vormen van hoe het dier eruitzag. Ankylosaurus was ruim zes meter lang, misschien zo lang als acht meter. Oudere boeken geven een lengte van wel elf meter maar die schattingen bleken achteraf veel te hoog te zijn. Hij had een erg platte en ronde romp met zeer brede heupen en korte poten. De nek was kort en droeg een korte brede kop voorzien van opvallend gekromde kaken waarin kleine tandjes stonden en die eindigden in een grote kromme snavel. Met die snavel en tanden rukte Ankylosaurus stukken plant af die hij bijna heel doorslikte en verder in zijn grote buik verteerde. De kop, romp en staart van Ankylosaurus waren bovenop bedekt met kleinere en grotere platte beenplaten. Die beschermden hem tegen de beten van het belangrijkste roofdier in zijn omgeving, de reusachtige Tyrannosaurus rex. Ankylosaurus kon zich daartegen ook goed verdedigen met een zware beenknots aan het uiteinde van zijn staart waarmee hij dodelijke klappen kon uitdelen. (nl)
Ankylozaur (Ankylosaurus) – rodzaj dinozaura pancernego. Obejmuje jeden gatunek – A. magniventris. Skamieniałe szczątki odnajdywano w formacjach geologicznych datowanych na koniec okresu kredowego, położonych na zachodzie Ameryki Północnej. Mimo iż nigdy nie odkryto kompletnego szkieletu ankylozaura, a kilka innych rodzajów reprezentuje o wiele szerszy materiał, ankylozaura często uznaje się za archetyp dinozaura pancernego. U innych przedstawicieli jego rodziny również występowały cechy tak charakterystyczne, jak ciężko uzbrojone ciało o grubym pancerzu i masywnej wieńczącej ogon, jednakże żaden z nich nie dorównywał mu wielkością. (pl)
Ankylosaurus ou anquilossauro é um gênero de dinossauro encouraçado herbívoro que viveu durante o final do período Cretáceo, no território que corresponde hoje à América do Norte. Trata-se de um gênero monotípico descoberto em 1908 por Barnum Brown, cuja a única espécie é denominada Ankylosaurus magniventris. O nome do gênero significa "lagarto fundido", e o nome específico significa "grande barriga". Um punhado de espécimes foi escavado até agora, mas um esqueleto completo não foi descoberto. Embora outros membros do clado Ankylosauria sejam representados por um material fóssil mais extenso, o Ankylosaurus é frequentemente considerado o membro arquetípico de seu grupo, apesar de ter algumas características incomuns. Estes animais pesavam entre 7 a 9 toneladas e media cerca de 9 metros de comprimento e 2 metros de altura. Possivelmente o maior anquilossaurídeo conhecido, estima-se que o Anquilossauro tenha entre 6 e 8 metros de comprimento e tenha pesado entre 4,8 e 8 toneladas. Era quadrúpede, com um corpo largo e robusto. Tinha um crânio largo e baixo, com dois chifres apontando para trás na parte de trás da cabeça, e dois chifres abaixo destes que apontavam para trás e para baixo. Ao contrário de outros anquilossauros, suas narinas estavam voltadas para os lados e não para a frente. A parte da frente das mandíbulas estava coberta por um bico, com fileiras de pequenos dentes em forma de folha mais atrás. Ele estava coberto de placas de armadura, ou osteodermas, com meio-anéis ósseos cobrindo o pescoço, e tinha um grande porrete na ponta de sua cauda. Ossos do crânio e outras partes do corpo foram fundidos, aumentando sua força, e essa característica é a fonte do nome do gênero. O Ankylosaurus é um membro da família Ankylosauridae, e seus parentes mais próximos parecem ser o e o Euoplocephalus. Acredita-se que o anquilossauro seja um animal lento, capaz de fazer movimentos rápidos quando necessário. Considera-se que a clava em sua cauda tenha sido usado na defesa contra predadores ou em competição intraespecífica. O anquilossauro foi encontrado nas formações Hell Creek, Lance, , e , mas parece ter sido raro em seu ambiente. Embora tenha vivido ao lado de outro anquilossauro nodosaurídeo, seus intervalos e nichos ecológicos não parecem ter se sobreposto, e o anquilossauro pode ter habitado áreas de planalto. Também viveu ao lado de dinossauros como Tyrannosaurus, Triceratops e Edmontosaurus. (pt)
Анкилоза́вр (лат. Ankylosaurus, от др.-греч. ἀγκύλος σαῦρος «согнутый ящер») — род вымерших растительноядных рептилий из надотряда динозавров, семейства анкилозаврид. Род является монотипическим, так как включает только один известный науке вид — Ankylosaurus magniventris. Ископаемые остатки анкилозавра были найдены в геологических формациях, датируемых наиболее поздней эпохой мелового периода (66,5—66 миллионов лет назад), на западе североамериканского материка. Несмотря на то, что до сих пор не был найден целый скелет анкилозавра, и на то, что родственные динозавры изучены лучше, именно анкилозавра принято считать (архетипом) представителем панцирных динозавров. Более того, это один из самых известных динозавров, несмотря на столь скудный окаменелый материал. Другие анкилозавриды обладали сходными чертами: телом, покрытым крепкими, тяжёлыми костными пластинами, сильным окостенением поверхности черепа и массивным утолщением, своеобразной «палицей», на конце хвоста. Анкилозавр является одним из самых больших известных анкилозаврид. (ru)
Ankylosaurus magniventris ("stel ödla med stor mage") är en art av dinosaurier som tillhör släktet Ankylosaurus, en ankylosaurid som var en fyrbent växtätare som lätt känns igen på sin kraftigt bepansrade kropp och den tydligt klubbformade svansen. Fossil efter Ankylosaurus har hittats i formationer som dateras tillbaka till slutet av kritaperioden i västra Nordamerika. Även om ett fullständigt skelett aldrig har hittats och flera andra släkten representeras av mer omfattande fossila material, anses Ankylosaurus ofta vara arketypen av bepansrade dinosaurier. Andra ankylosaurider delar dess välkända drag, så som den tungt bepansrade kroppen och dess massiva benklubba i änden av svansen, men Ankylosaurus var den största medlemmen i dess familj. (sv)
Анкілозавр (Ankylosaurus) — рід динозаврів. Довжина тіла анкілозавра досягала 9 метрів. Висота ж до 3 м. Важив він до 6 тонн. Існували анкілозаври в кінці крейдяного періоду близько 68 — 66 мільйонів років тому. Були поширені в західній частині Північної Америки. (uk)
甲龍屬（屬名：Ankylosaurus，意為「僵硬的蜥蜴」）是甲龍科恐龍的一個屬，當中只有一種，為大面甲龍（A. magniventris）。甲龍最大的特點是擁有全骨板覆蓋的身驅及巨型的尾部棒槌，也是甲龍科內最大型的物種，不過牠們的體重只有2噸到4噸，非常輕量級，身高也比同體長的恐龍低很多，方便中心下移、用甲殼保護自己柔軟的腹部。甲龍化石是在北美洲西部的地獄溪組被大量發現，屬於約6800萬到6550萬年前白堊紀末期的整個美國和墨西哥地區，和三角龍、暴龍、愛德蒙托龍和腫頭龍生活在同一個地質年代。 (zh)
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https://dinotoyblog.com/wuerhosaurus-haolonggood/
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Wuerhosaurus (Haolonggood) – Dinosaur Toy Blog
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2023-11-22T22:59:36+00:00
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https://dinotoyblog.com/wuerhosaurus-haolonggood/
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Wuerhosaurus is a genus of stegosaurid that lived during the early Cretaceous in China. Being from the early Cretaceous makes it notable as it means it’s one of the last living stegosaurid genera. While stegosaurids as a group flourished during the late Jurassic, they went completely extinct by the end of the early Cretaceous. A few figures of Wuerhosaurus exist including an old 2009 one by CollectA, a mini by PNSO, and one by Vitae. Today we’re looking at Haolonggood’s Wuerhosaurus, released in 2023.
Remains for Wuerhosaurus are scant and include a mostly complete pelvis and sacrum, some vertebrae, bits of the shoulder and forelimbs, and a couple plates. Wuerhosaurus is typically reconstructed with shallow plates like what’s preserved, but this is probably incorrect since the preserved plates are broken. Indeed, the most recent study on Wuerhosaurus phylogeny places it close to Stegosaurus and it’s likely that it had larger plates than what we typically see.
The Haolonggood Wuerhosaurus has the shallow plates that you would expect it to have. Chances are good that this is inaccurate, but we would need more material to say for sure. The shallow plates are somewhat of a trope for the genus and without them the figure wouldn’t really be recognizable as a Wuerhosaurus, so I’m cool with it for now.
Any reconstruction of Wuerhosaurus is going to be largely speculative anyway but Haolonggood’s figure does have the standard stegosaurid body plan that’s generally accurate for the group. It is presented low to the ground which is correct for the genus. It appears that five digits are sculpted on each forelimb, and it appears that digits 4 and 5 are clawless. I say “it appears” since the digits are so small it’s hard to study them closely. The hindlimbs have three digits. This is all correct as well. You’ll notice that this figure lacks shoulder spikes while Vitae’s has them but there is no evidence for them in Wuerhosaurus and their presence is unlikely.
The Haolonggood Wuerhosaurus measures 7” (17.78 cm) in length and stands 2.5” (6.35 cm) tall at the hips. It is estimated that Wuerhosaurus measured 23’ (7 meters) long which puts the figure at 1/39 in scale. The figure is presented striding forward with the neck bending slightly leftward and the tail with some subtle curves.
This figure comes with two different paintjobs, green and blue. I got the blue variant but it’s really more of a purple. I chose this one because purple dinosaurs are a rarity while green dinosaurs are not and are especially common in stegosaur figures. Patterning is the same on both figures. The main body is purple with a peach-colored underside that radiates in stripes across the body. Darker purple squiggles and spots add further complexity to the figure. The plates are edged in purple with an orange center, looking somewhat like a little sunset on each plate. The smallest plates on the neck are painted the same color as the body. The tail is tipped in vibrant orange in typical Haolonggood fashion. Claws are painted grey, and the eyes are simple shiny black specks.
Overall, I find this paintjob unique, but I don’t care for the darker squiggles. Also, the entire figure has a flat, hazy look to it, like it was dusted with flour. This makes the finer details harder to see and appreciate and it also makes it difficult to accurately represent the figure’s appearance in photographs. I’ve seen other collectors add a dark wash to their copies with great effect.
Although they’re hard to see the fine details are indeed there. The entire figure is covered in fine pebbly scales and saggy looking creases and skin folds. Larger feature scales run in rows along the flanks and clusters of larger scales are sculpted at the bases of the plates and tail spikes. The plates themselves have unique jagged edges and grooves that I enjoy. A cloaca is sculpted on the underside along with a round bulge on either side of it. These remind me of the hemipenal bulges seen in male lizards so I guess it’s a boy!
Even though the figure is largely speculative, and the shallow plates are likely inaccurate, I’m happy to finally have a Wuerhosaurus in my collection to boost my stegosaur diversity. The figure itself is masterfully sculpted and lifelike and I find nothing to take issue with aside from the hazy appearance that can be easily remedied if one felt so inclined. The Hoalonggood Wuerhosaurus is currently available and retails at about $17-20, an excellent price point for a figure of this quality.
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https://en.wikipedia.org/wiki/Cedarpelta
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Cedarpelta
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2006-03-07T22:34:48+00:00
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/static/apple-touch/wikipedia.png
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https://en.wikipedia.org/wiki/Cedarpelta
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Skull of Cedarpelta bilbeyhallorum, on display at the USU Eastern Prehistoric Museum, Price, Utah. Scientific classification Domain: Eukaryota Kingdom: Animalia Phylum: Chordata Clade: Dinosauria Clade: †Ornithischia Clade: †Thyreophora Clade: †Ankylosauria Family: †Ankylosauridae Genus: †Cedarpelta
Carpenter et al., 2001 Species:
†C. bilbeyhallorum
Binomial name †Cedarpelta bilbeyhallorum
Carpenter et al., 2001
Cedarpelta is an extinct genus of basal ankylosaurid dinosaur from Utah that lived during the Late Cretaceous period (Cenomanian to lower Turonian stage, 98.2 to 93 Ma) in what is now the Mussentuchit Member of the Cedar Mountain Formation. The type and only species, Cedarpelta bilbeyhallorum, is known from multiple specimens including partial skulls and postcranial material. It was named in 2001 by Kenneth Carpenter, James Kirkland, Don Burge, and John Bird. Cedarpelta has an estimated length of 7 metres (23 feet) and weight of 5 tonnes (11,023 lbs). The skull of Cedarpelta lacks extensive cranial ornamentation and is one of the only known ankylosaurs with individual skull bones that are not completely fused together.
Discovery and naming
[edit]
The partial remains of an ankylosaur were discovered by Evan Hall and Sue Ann Bilbey at the CEM site near the Price River in Carbon County, Utah while they were visiting an excavation in the surrounding area.[1] The site was originally interpreted as being from the top of the Ruby Ranch Member of the Cedar Mountain Formation,[1] but was later interpreted as being from the bottom of the Mussentuchit Member.[2] The age of the layer was originally thought to have been 104.46 ± 0.95 Ma,[3] but more recent estimates date it to 98.2 ± 0.6 to 93 Ma.[4] In 1998, the discovery was reported by Kenneth Carpenter and James Kirkland.[5] In 2001, it was subsequently described, along with other material, by Kenneth Carpenter, James Kirkland, Don Burge, and John Bird. The holotype specimen, CEUM 12360, consists of a partial skull that is missing the snout and lower jaws. Numerous osteoderms, postcranial material and a disarticulated skull were designated as paratype specimens. Both holotype and paratype specimens represent at least three individuals and are currently housed at the College of Eastern Utah, Prehistoric Museum, Utah.[5][1]
The generic name, Cedarpelta, is derived from the Cedar Mountain Formation and the Greek word "pelte" (small shield). The specific name, bilbeyhallorum, honours Sue Ann Bilbey and Evan Hall, who discovered the remains of Cedarpelta.[1]
In 2008, additional specimens were referred to Cedarpelta from the Price River II Quarry, which is about 24.5 km southeast of Price River, Utah and at the base of the Mussentuchit Member. The quarry also produced specimens pertaining to four individuals of a brachiosaurid, an iguanodontian, a turtle, a pterosaur, and specimens of the nodosaurid Peloroplites. The referred material includes: CEUM 10396, a cervical vertebra; CEUM 10412, CEUM 10404, caudal vertebrae; CEUM 10371, a coracoid; CEUM 10256, CEUM 11629, humeri; CEUM 10266, an ischium; CEUM 11334, a femur; and CEUM 11640, a tibia.[2]
Description
[edit]
Carepnter et al. (2001) originally gave Cedarpelta an estimated length of 7.5-8.5 metres (24.6-27.9 feet). However, Gregory S. Paul gave a lower estimate of 7 metres (23 feet) and a weight of 5 tonnes (11,023 lbs), while Thomas Holtz gave a higher estimation at 9 meters suggesting that it was rivalling Ankylosaurus.[6][7][8]
Carpenter et al. (2001) established several distinguishing traits of Cedarpelta. The body of the praemaxilla, the front snout bone, is short in front of its nasal branch. The outer sides of the two praemaxillae run more parallel compared to the snouts of later forms which are strongly diverging to behind. The cutting edge of the bone core of the upper beak is limited to the front of the praemaxilla. Each praemaxilla has six (conical) teeth. The quadrate, and with it the entire back of the skull, is inclined to the front. The head of the quadrate is not fused with the paroccipital process, contrary to the situation in Shamosaurus. The neck of the occipital condyle is long and sticking out to behind, like with nodosaurids, not obliquely to below as in typical ankylosaurids. The tubera basilaria, appending processes of the rear lower braincase, form a large wedge directed to below. The pterygoid is elongated from the front to the rear and has a saddle-shaped process on its outer edge oriented to behind and sideways. The coronoid process of the rear lower jaw has an oval process at the inside. The straight ischium has a knob-shaped boss at the inside near the pubic pedicle.[1]
Cedarpelta shows a mix of basal and derived traits. The presence of premaxillary teeth is a plesiomorphic character because it is inherited from earlier Ornithischia. In contrast, closure of the opening on the side of the skull behind the orbit, the lateral temporal fenestra, is an advanced, derived (apomorphic) character only known in ankylosaurid ankylosaurians.[1]
Two skulls are known, and the skull length for Cedarpelta is estimated to have been roughly 60 centimetres (24 in). One of the Cedarpelta skulls was found disarticulated, a first for an ankylosaur skull, allowing paleontologists a unique opportunity to examine the individual bones instead of being limited to an ossified unit. The skull is relatively elongated and does not show a strongly appending beak. Of the conical premaxillary teeth, the first is the largest. The maxilla bears eighteen teeth. The eye socket is surrounded by the lacrimal, a single supraorbital and a large postorbital, excluding the prefrontal and the jugal from the orbital rim. The postcranial skeleton was in 2001 not described in any detail.[1]
The skulls, though of large and thus not juvenile individuals, do not show a distinctive pattern of fused caputegulae, head tiles. This inspired Carpenter to propose an alternative hypothesis of ankylosaur skull osteoderm formation. Formerly, it had been assumed that such armour plates were either formed by direct skin ossification into distinct scutes which later fused to the skull (the more popular theory), or by a reaction of the skull bones to the pattern of overlying scales. The lack of a clear pattern in Cedarpelta suggested to Carpenter that the ossification took place in an intermediate layer between the scales and the skull roof itself, which he surmised to have been the periosteum.[1]
Classification
[edit]
Carpenter (2001) placed Cedarpelta within the family Ankylosauridae and offered two interpretations of its position. The first was that it could be the basalmost known ankylosaurid, i.e. the first discovered branch to split off from the ankylosaurid stem line. This would be in line with its plesiomorphic traits and the fact that the in 2001 supposed Barremian age made it one of the oldest known ankylosaurids. The second was that it formed an early ankylosaurid branch, or clade, Shamosaurinae together with Gobisaurus of north-central China and the eponymous Shamosaurus of Mongolia.[9] Thompson et al. (2012),[10] Chen et al. (2013),[11] Yang et al. (2013),[12] Han et al. (2014),[13] Arbour & Currie (2015),[14] Arbour et al. (2016),[15] Arbour & Evans (2017),[16] Yang et al. (2017),[17] Zheng et al. (2018),[18] Rivera-Sylva et al. (2018),[19] Park et al. (2019)[20] and Frauenfelder et al. (2022)[21] have all found Cedarpelta to be within Ankylosauridae, as either within a polytomy with Liaoningosaurus, Aletopelta, Chuanqilong, Gobisaurus and Shamosaurus or as sister taxon to Chuanqilong. The results of Arbour & Currie (2015) are reproduced below.
Vickaryous et al. (2004) interpreted Cedarpelta as the basalmost member of the family Nodosauridae, positioned even below the nodosaurids Pawpawsaurus, Silvisaurus, and Sauropelta.[22] Wiersma & Irmis (2018) also interpreted Cedarpelta as a nodosaurid.[23] The results of Vickaryous et al. (2004) are reproduced below.
See also
[edit]
Dinosaurs portal
Timeline of ankylosaur research
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https://en.wikipedia.org/wiki/Zhejiangosaurus
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Zhejiangosaurus lishuiensis on display at the Zhejiang Museum of Natural History Scientific classification Domain: Eukaryota Kingdom: Animalia Phylum: Chordata Clade: Dinosauria Clade: †Ornithischia Clade: †Thyreophora Clade: †Ankylosauria Clade: †Euankylosauria Genus: †Zhejiangosaurus
Lü et al., 2007 Species:
†Z. lishuiensis
Binomial name †Zhejiangosaurus lishuiensis
Lü et al., 2007
Zhejiangosaurus (meaning "Zhejiang lizard") is an extinct genus of ankylosaurian dinosaur from the Upper Cretaceous (Cenomanian stage) of Zhejiang, eastern China. It was first named by a group of Chinese authors Lü Junchang, Jin Xingsheng, Sheng Yiming and Li Yihong in 2007 and the type species is Zhejiangosaurus lishuiensis ("from Lishui", where the fossil was found).[1] It has no diagnostic features, and thus is a nomen dubium.[2]
Description
[edit]
Zhejiangosaurus could grow up to 4.5 m (17 ft) in length and was 1.4 metric tons in weigh.[3]
Material
[edit]
Material for Zhejiangosaurus consists of the holotype, ZNHM M8718, a partial skeleton which has preserved a sacrum with eight vertebrae, a complete right ilium and partial left ilium, a complete right pubis, the proximal end of the right ischium, two complete hindlimbs, fourteen caudal vertebrae, and some unidentified bones. These remains come from Liancheng, in the Chinese administrative unit of Lishui on the province of Zhejiang and they were collected from the Cenomanian-age Chaochuan Formation.[1]
Systematics
[edit]
On the species description, Lü et al. (2007) found Zhejiangosaurus to belong to the ankylosaurian family Nodosauridae.[1][4]
Zhejiangosaurus in a cladogram after Pond et al. (2023):[5]
See also
[edit]
Dinosaurs portal
China portal
Timeline of ankylosaur research
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https://fossil.fandom.com/wiki/Edmontonia
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Edmontonia
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Edmontonia (meaning "From the Edmonton Formation") is an extinct armoured dinosaur, a part of the nodosaur family from the Late Cretaceous period. It is named after the Edmonton Formation (now the Horseshoe Canyon Formation), the unit of rock it was found in. Edmontonia was bulky and tank-like...
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Edmontonia (meaning "From the Edmonton Formation") is an extinct armoured dinosaur, a part of the nodosaur family from the Late Cretaceous period. It is named after the Edmonton Formation (now the Horseshoe Canyon Formation), the unit of rock it was found in.
Description[]
Size and general build[]
Edmontonia was bulky and tank-like at roughly 6.6 meters (22 feet) long and 2 meters (6.5 feet) high.[citation needed] It had small, ridged bony plates on its back and head and many sharp spikes along its back and tail. The four largest spikes jutted out from the shoulders on each side, two of which were split into subspines in some specimens.[1] Its skull had a pear-like shape when viewed from above. (Compare 26 feet long and 8 feet high for the M1 Abrams army tank.)
Distinguishing traits[]
In 1990, Kenneth Carpenter established some diagnostic traits for the genus as whole, mainly comparing it with its close relative Panoplosaurus. In top view, the snout has more parallel sides. The skull armour has a smooth surface. In the palate, the vomer is keeled. The neural arches and neural spines are shorter than those of Panoplosaurus. The sacrum proper consists of three sacral vertebrae. In the shoulder girdle, the scapula and coracoid are not fused.
Carpenter also indicated in which way the main species differed from each other. The type species, Edmontonia longiceps, is distinguished from E. rugosidens in lacking sideways projecting osteoderms behind the eye sockets; having tooth rows that are less divergent; possessing a more narrow palate; having a sacrum that is wider than long and more robust; and in having shorter spikes at the sides. Also, an ossified cheek plate, known from E. rugosidens specimens, has not been found with Edmontonia longiceps.
Skeleton[]
The skull of Edmontonia, up to half a metre long, is somewhat elongated with a protruding truncated snout. The snout carried a horny upper beak and the front snout bones, the premaxillae, were toothless. The cutting edge of the upper beak continued into the maxillary tooth rows, each containing fourteen to seventeen small teeth. In each dentary of the lower jaws, eighteen to twenty-one teeth were present. In the sides of the snout large depressions were present, "nasal vestibules", that each possessed two smaller openings. The top of these was a horizontal oval and represented the bony external nostril, the entrance to the nasal cavity, the normal air passage. The more rounded second opening below and obliquely in front, was the entrance to a "paranasal" tract, running along the outer side of the nasal cavity, in a somewhat lower position. A study by Matthew Vickaryous in 2006 proved for the first time the presence of multiple openings in a nodosaurid; such structures had already been well established in ankylosaurids. The air tracts are however, much simpler than in the typical ankylosaurid condition, and are not convoluted while lacking bony turbinate bones. The nasal cavity is separated into two halves along the midline by a bone wall. This septum is continued to below by the vomers, which are keeled, the keel featuring a pendulum-shaped appendage. Another similarity with Ankylosauridae is the presence of a secondary bone palate, a possible case of parallel evolution. This has been shown too for Panoplosaurus.
The head armour tiles, or caputegulae, are smooth. Details differ between the various specimens but all share a large central nasal tile on the snout, bend large "loreal" tiles at the rear snout edges and a large central caputegula on the skull roof. The tiles behind the upper eye socket rim in Edmontonia longiceps do not stick out as much as in E. rugosidens, combined with a more narrow, pointed snout in the former. Some E. rugosidens specimens are known that possess a "cheek plate" above the lower jaw. Contrary to that discovered with Panoplosaurus, it is "free-floating", not fused with the lower jaw bone. E. schlessmani was described as having a wide rear skull.
The vertebral column contains about eight neck vertebrae, about twelve "free" back vertebrae, a "sacral rod" of four fused rear dorsal vertebrae, three sacral vertebrae, two caudosacrals and at least twenty, but probably about forty, tail vertebrae. In the neck the first two vertebrae, the atlas and axis, are fused. In the shoulder girdle, the coracoid has a rectangular profile, in contrast to the more rounded shape with Panoplosaurus. Two sternal plates are present, connected to sternal ribs. The forelimb is robust but relatively long. In Edmontonia longiceps and E. rugosidens the deltopectoral crest of the humerus is gradually rounded. The metacarpus is robust compared to that of Panoplosaurus. The hand very likely was tetradactyl, having four fingers.[4] The exact number of phalanges is unknown but the formula was by W.P. Coombs suggested to be 2-3-3-4-?.
Osteoderms[]
Apart from the head armour, the body was covered with osteoderms, skin ossifications. The configuration of the armour of Edmontonia is relatively well known, much of it having been discovered in articulation. The neck and shoulder region was protected by three cervical halfrings, each consisting of fused rounded rectangular, asymmetrically keeled, bone plates. These halfrings did not have a continuous underlying bone band. The first and second halfrings each had three pairs of segments. Below each lower end of the second halfring a side spike was present, a separate triangular osteoderm pointing obliquely forward. In the third halfring over the shoulders, the two pairs of central segments are bordered on each side by a very large forward-pointing spike that is bifurcated, featuring a secondary point above the main one. A third large spike behind it points more sideways; a smaller fourth one, often connected to the third at the base, is directed obliquely to behind. The row of side spikes is continued to the rear but there the osteoderms are much lower, curving strongly to behind, with the point overhanging the rear edge. Gilmore had trouble believing that the shoulder spikes really pointed to the front as this would have greatly hampered the animal while moving through vegetation. He suggested that the points had shifted during the burial of the carcass. However, Carpenter and G.S. Paul, trying to reposition the spikes, found that it was impossible to rotate them without losing conformity with the remainder of the armour. The side spikes have solid, not hollow, bases. The spikes differ in size between E. rugosidens individuals; those of the E. longiceps holotype are relatively small.
Behind the third halfring the back and hip are covered by numerous transverse rows of much smaller oval keeled osteoderms. These are not ordered in longitudinal rows. The front rows have plates oriented along the length of the body, but to the rear the long axis of these osteoderms gradually rotates sideways, their keels ultimately running transversely. Rosettes are lacking. The configuration of the tail armour is unknown. The larger plates of all body parts were connected by small ossicles. Such small round scutes also covered the throat.
Discovery and species[]
In 1915, the American Museum of Natural History obtained the nearly complete, articulated front half of an armoured dinosaur. In 1922, William Diller Matthew referred this specimen to Palaeoscincus. In 1940, Lori Russell referred it to Edmontonia rugosidens.
The type species of Edmontonia, E. longiceps was discovered in 1924 by George Paterson. It wasn't named until 1928 by C. M. Sternberg. E. rugosidens, formally named by Gilmore in 1930, is reported from the Aguja formation in Texas. Edmontonia species include:
E. longiceps, the type, is known from the middle Horseshoe Canyon Formation (Unit 2) dated to 71.5-71 million years ago.
E. rugosidens, is sometimes given its own genus, Chassternbergia, first coined as a subgenus by Dr. Robert T. Bakker in 1988 (Edmontonia (Chassternbergia) rugosidens) and based on differences in skull proportion from E. longiceps and its earlier time period. This subgenus or genus is not generally accepted; It is found in the lower Dinosaur Park Formation, dating about 76.5-75 million years ago.
And E. australis, which is known from cervical scutes only, and is considered to be a dubious name or a synonym of Glyptodontopelta mimus.
Usually included in this genus is Denversaurus schlessmani ("Schlessman's Denver lizard"). This taxon was erected by Bakker in 1988 for a skull from the Late Maastrichtian Upper Cretaceous Lance Formation of South Dakota, but considered by later workers to belong to Edmontonia rugosidens. The type specimen of Denversaurus is in the collections of the Denver Museum of Natural History (now the Denver Museum of Nature and Science), Denver, Colorado (for which the genus was named).
Phylogeny[]
C.M. Sternberg originally did not provide a classification of Edmontonia. In 1930, L.S. Russell placed the genus in the Nodosauridae, which has been confirmed by subsequent analyses. Edmontonia was generally shown to be a derived nodosaurid, closely related to Panoplosaurus. Russell in 1940 named a separate Edmontoniinae. In 1988 Bakker proposed that the Edmontoniinae with the Panoplosaurinae should be joined into Edmontoniidae, the presumed sister group of the Nodosauridae within Nodosauroidea which he assumed not be ankylosaurians but the last surviving stegosaurians. Exact cladistic analysis has not confirmed these hypotheses however, and the concepts of Edmontoniinae and Edmontoniidae are not in modern use.
The following cladogram shows the position of Edmontonia in the nodosaurid evolutionary tree according to an analysis in 2011 by paleontologists Richard Stephen Thompson, Jolyon C. Parish, Susannah C. R. Maidment and Paul M. Barrett.
Nodosauridae
Antarctopelta
Mymoorapelta
Hylaeosaurus
Anoplosaurus
Tatankacephalus
Horshamosaurus
Polacanthinae
Gargoyleosaurus
Hoplitosaurus
Gastonia
Peloroplites
Polacanthus
Struthiosaurus
Zhejiangosaurus
Hungarosaurus
Animantarx
Niobrarasaurus
Nodosaurus
Pawpawsaurus
Sauropelta
Silvisaurus
Stegopelta
Texasetes
Edmontonia
Panoplosaurus
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Plans to display Dinosaur Trackways in Washington D.C.
When asked to comment on dinosaur discoveries in the United States most experts may cite discoveries in the Badlands of Montana or the Cleveland-Lloyd Dinosaur quarry in Utah. Certainly, it is true to say that they are many fantastic Mesozoic fossil sites in the west of the USA but the eastern part of the United States, although perhaps a little under-represented in terms of fossil evidence, can still spring a few surprises.
Dinosaur Tracks
Now a new study of fossil trackways in Maryland, north-eastern USA has provided a glimpse into a thriving dinosaur based eco-system. Many of the trackways, have been found just a few miles drive out of Washington D.C. Trackways and footprints are called trace fossils. Trace fossils preserve evidence of the activity of animals such as their trackways, borings or burrows. The problem with most sets of footprints, even the very best preserved ones, is that, unless the animal is found fossilised at the end of the trackway, scientists can never be 100% certain as to the species or genus that actually left the prints.
Trace fossils such as footprints do have a significant advantage over other types of fossil such as fossil bones, most are direct in situ evidence of the environment at the time and place the organism was living.
Studying Trace Fossils
A total of over 900 fossilised footprints from a variety of dinosaurs all dated from the Cretaceous have been identified from the area. Theropods, ankylosaurs (Nodosauridae), sauropods and ornithopods are represented by the prints. Palaeontologists have estimated that the trackways were made between 121 and 98 million years ago.
Trace fossils of other animals have also been preserved in the this part of the USA, one trackway has been identified as a flying reptile, perhaps a pterosaur flew down to get a drink and its trail was preserved in the soft sediment. Mammal tracks have also been found, indeed one trackway indicates that some mammals were quite large, tracks of a quadrupedal mammal about the size of a large dog have been recorded.
Visit the Everything Dinosaur website: Everything Dinosaur.
Two Dozen Species of Dinosaur
“Based on the trace fossils, over two dozen species of dinosaurs were living in Maryland at that time,” co-author of the study, Ray Stanford commented. Ray specialises in studying fossil trackways, he began to discover tracks in the area whilst out looking for native Indian artifacts, in the stream-beds that criss cross the area.
He explained that as water and human development erode such beds, “floats” can result. These are pieces of track-bearing substrate that hydrodynamically dislodge from their natural stratigraphic context during stream bank flooding.
“This is one instance where building booms and storms can benefit science,” he said.
All of the discoveries were made either in Prince George’s county, near the capital, Washington D.C. or at the White Marsh Run area of Baltimore county.
An Illustration of the Track Made by an Ornithopod Dinosaur
Picture credit: Everything Dinosaur
Photographs show a number of footprint specimens, the peculiar, almost flower-shaped five-toed print in the foreground was most probably made by a nodosaur. Nodosaurs are members of the Ankylosauria, heavily built, slow-moving, plant-eaters with body armour and horns.
To read an early article about dinosaur tracks discovered on the North Yorkshire coast: Dinosaur Tracks Found by Young Boy.
Ray Stanford in conjunction with a Johns Hopkins University palaeontologist called Davide Weishampel hope to publish a journal paper on this new genus of nodosaur. The nodosaur print in the foreground is much smaller than the cast print in the very centre of the image (the print which the model nodosaur is facing), this indicates that some of the trackways may have been made by young, immature animals. This area may have provided a Cretaceous nursery for many species, a popular nesting and breeding ground for a variety of dinosaurs.
Providing an Insight into Dinosaur Behaviour
The scientists state that they may even have uncovered trackway evidence showing youngsters following adults, a possible insight into animal’s behavioural and social relationships.
So far, Stanford has described and published Maryland’s first dinosaur track species (called an ichnospecies which translates to ‘trace species’). It consists of both front and back footprints of a hypsilophodontid dinosaur. He named the new dinosaur footprint type or species Hypsiloichnus marylandicus, meaning “trace of a hypsilophodontid dinosaur from Maryland.”
An overview of these, and other, finds was recently published in the journal Ichnos.
Analysis of the region’s geology indicates that during that dinosaur era, fresh water sources and plant life would have been plentiful. Stanford has excavated fossilised pollen for ancient plants, along with fossilised wood for a large, now-extinct fern tree similar to today’s cycads.
The Smithsonian Museum of Natural History in Washington D.C. is investigating the possibility of putting some of the tracks on display in a special exhibition. There are certain obstacles to overcome, such as how best to present the casts so that their fine detail can be seen, but such an exhibit be popular with museum visitors. After all, it would give the residents of Washington D.C. an opportunity to learn more about some of the previous residents in the neighbourhood.
Everything Dinosaur stocks a wide selection of dinosaur models including replicas of ornithopods and nodosaurids: Everything Dinosaur Models and Replicas.
Earthquake shakes the Country – Epicentre 4km north of Market Rasen, Lincolnshire
At shortly before 1am this morning (GMT) an earthquake with a magnitude of 5.2 struck the United Kingdom. The epicentre (the point on the Earth’s surface directly above the centre of the earthquake), was 4 kilometres north of the town of Market Rasen, Lincolnshire. Reports have been received of a Market Rasen earthquake!
There is one report of an injury, the British Geological Society (BGS) had by 7am received over 1,400 reports from members of the public, the media and the emergency services. Some structural damage has been caused, chimneys falling off, walls collapsing close to the epicentral area, but this tremor was felt across a large part of the UK. Many residents in English and Wales towns were awoken by the shaking, the quake has been felt as far away as southern Scotland.
Market Rasen Earthquake
In this country we are not immune from earthquakes, each year the BGS records around 200, but only about 10% are big enough to be felt by local residents. Fortunately, most of the quakes have epicentres which are offshore. The largest earthquake recorded in the British Isles took place in 1931. This quake had a local magnitude of 6.1, but fortunately it was centred on the Dogger Bank area of the North Sea. Even so, the quake and the aftershocks were powerful enough to cause structural damage to many buildings on the east coast of England.
Finding the Epicentre
The precise epicentre of the Market Rasen quake has been calculated to be latitude 53.419 degrees north and longitude 0.354 degrees west. It is understood to have taken place approximately 5,000 metres underground.
Earthquakes are monitored by the British Geological Survey, part of the Natural Environment Research Council (NERC). There is a network of 146 seismometer stations across the UK sending data to the head office based in Edinburgh four times per day. However, during times of earthquake activity data can be sent on demand and staff at the BGS can access data and analyse results from home. They are on call 24-hours a day, as scientists don’t know when a quake will strike.
Earthquakes of this magnitude occur approximately ever 30 years or so, in the world there are about 1,300 quakes of this magnitude or bigger each year. This latest quake is the biggest since 1984, when on the 19th July North Wales was struck by an earthquake that had a magnitude of 5.4. It too caused structural damage to many buildings with cities such as Liverpool 120 kilometres from the epicentre being affected.
106 Tremors
None of the team members at Everything Dinosaur felt the quake (all sound asleep in our beds). However, one member of staff recalled the Manchester earthquakes that struck in the Autumn of 2002. A series of tremors were recorded with an epicentre in and around Manchester over a period of five weeks. The magnitude ranged from 1.1 to 3.9 ML (local magnitude). In total 106 tremors were recorded, the biggest of which (3.9 ML) hit on October 21st. Our colleague remembers particular incident very well, as he was travelling in a lift in an office block in the centre of Manchester at the time – very scary.
To read more about the work of the BGS and the latest on this mornings quake you can visit the BGS website.
Visit Everything Dinosaur’s award-winning website: Everything Dinosaur.
Rises in Oxygen Levels may Explain “Cambrian Explosion”
A new study from a multi-national team of scientists provides evidence of the link between the explosion of early life forms and the oxidation of the deep oceans. The rise of oxygen levels within the ocean between 635 and 551 million years ago may have helped trigger the increase and rapid diversification of early lifeforms, leading ultimately to the “Cambrian Explosion”.
The “Cambrian Explosion” is a term used by scientists to describe the huge increase in life that occurred around 545 million years ago, at this stage of the history of life on Earth, all life was associated with marine environments. It was during the Cambrian that most of the major groups of animals that exist today evolved.
Speedy Evolution
Soft bodied animals and the stromatolites (colonies of bacteria) were partly replaced and superseded by the evolution of organisms with hard parts such as exoskeletons and shells. The first forms of life that could be biomineralised evolved, this meant that the hard parts of their bodies could be preserved as fossils and thus this period of ancient history not only marks the increasing abundance and diversity of organisms but also marks the start of an enriched fossil record, providing palaeonotologists with more evolutionary evidence.
Complex organisms had been in existence prior to the beginning of the Palaeozoic, but the fossil record is extremely poor. Multi-cellular life forms have been recorded in rocks of approximately 600 million years of age, but these creatures seemed to have lacked any hard parts and as soft-bodied creatures, palaeontologists have only a few tantalising fossils to work with.
A Rise in Oxygen Levels
The rise in oxygen levels and the oxidation of deep oceans in the late Precambrian has been accepted for a number of years. However, it had been thought that the increase in photosynthetic bacteria such as cyanobacteria (formerly known as blue-green algae), assisted by other non-biological means such as the breakdown of water into hydrogen and oxygen by ultraviolet rays penetrating to the surface of the Earth through the ozone devoid atmosphere had led to the increase.
Now, new research from scientists studying the geochemical structure of the Duoshantuo Formation in southern China reveals that life on Earth may have been influenced by two distinct pulses of oxygen. The first increase in oxygen predates the “Cambrian Explosion” by a significant amount of time but may have led to an increase in microscopic life forms. The second burst of oxygen aerating the oceans seemed to have occurred around 550 million years ago and in geological terms immediately pre-dates the increase in life during the Early Cambrian.
Trilobites Thrived During the Cambrian
Picture credit: Everything Dinosaur
An international team of scientists from Virginia Tech, the University of Maryland, University of Nevada (Las Vegas) and the Chinese Academy of Sciences set out to test the relationship between the evolution of more complex and diverse life forms and environmental change. To do this the team needed to find sedimentary strata that pre-dates the Cambrian and a sequence of strata (stratigraphic column) that would show deposition and formation as a timeline, one that had not been altered or changed by other chemical or geological processes.
Finding pristine Precambrian strata is a challenge in itself but such locations are known, one being the Doushantuo Formation in the Yangtze Gorges area, Guizhou Province, southern China. The strata consists of phosphate and dolomite sequences, laid down at the bottom of a sea. China at this time was made up of two separate and submerged continental sheets, that lay in shallow, warm tropical waters off the coast of the super-continent Gondwana. The first part of what was to become China, closest to Gondwana, straddled the Equator, the second part lay across the Tropic of Cancer.
Mapping Oxygen Levels
By mapping the levels of oxygen at various levels in the stratigraphic column, the team could measure the amount of oxygen in the marine environment and then associate this with the biostratigraphic column (fossils used to date and correlate strata), this would provide evidence to support the increase in oxygen leading to a diversity and increase in lifeforms.
To calculate when there was enough oxygen to support animal life in the ocean, the researchers asked, “What kind of geochemical evidence would there be in the rock record?” said Shuhai Xiao, associate professor of geosciences at Virginia Tech.
Scientists hypothesized that there was a lot of dissolved organic carbon in the ocean when oxygen levels were low. If oxygen levels rose, some of this organic carbon would be oxidized into inorganic forms, some of which can be preserved as calcium carbonate (CaCO3 ) in the rock record. “We measured the carbon isotope signatures of organic and inorganic carbon in the ancient rocks to infer oxidation events,” said co-author Ganqing Jiang, assistant professor of geology at the University of Nevada at Las Vegas.
The stratigraphic column exposed during the construction of the dams in the Yangtze Gorges area represents a large slice of ancient geological history. The researchers carefully took samples from each strata of rock, the deeper the strata, then, unless the strata has been overturned, as can sometimes happen during mountain building processes for example, the older the rocks will be. This is an important geological principle it is called the “Law of Superposition”. Many hundreds of different samples were taken, representing marine deposits laid down during the Precambrian and Early Cambrian.
The researchers cleaned and crushed the small samples to powder, which they reacted with acid to release carbon dioxide from carbonate minerals, and then burned the residue to get carbon dioxide from organic matter. “The carbon dioxide that is released was measured with mass spectrometers to gives us the isotopic signature of the carbonate and organic carbon that was present in the rock,” a researcher commented.
“The relative abundances of the carbon-12 and carbon-13 isotopes, which are stable and do not decay with time, provide a snapshot of the environmental processes taking place in the ocean at the different times recorded in the layers of rock”.
The stratigraphic pattern of carbon isotope abundances suggested to these researchers that the ocean, which largely lacked oxygen before animals arrived on the scene, was aerated by two discrete pulses of oxygen.
The first pulse that occured in the Precambrian seemed to have little impact on a large organic carbon reservoir in the deep ocean, but did spark changes in microscopic life. The second event, which occurred around 550 million years ago, immediately prior to the palaeontological event known as the “Cambrian Explosion”, resulted in the reduction of the organic carbon reservoir. This indicates that the ocean became fully oxidized just before the evolution and diversification of many of Earth’s earliest animals. Perhaps this dramatic increase in the level of available oxygen provided the fuel for the rapid burst of evolution.
Certainly, scientists have speculated why all of our sudden around 545 million years ago evolution seems to have pressed the accelerator when for much of the Precambrian (Cryptozoic), evolution seemed to be progressing at a very slow pace. You could say that evolution, prior to the second pulse of oxygen had progressed at a snail’s pace but to be fair to the Gastropods (snails) these animals did not really get going until the Early Cambrian.
Photographs show a field of view 0.15 millimetres in diameter of a beautifully preserved eukaryotes fossil from the Doushantuo formation (635-550 million years old). Eukaryotes are cells with their genetic material enclosed in a cell nucleus. Eukaryotes are believed to have first appeared in the fossil from strata dated to 2,100 million years ago, but evidence from molecular biology indicates that they may have been present earlier than this but left little or no fossil evidence.
“The Doushantuo Formation has a wonderful fossil record. It allows us to look at major fossil groups, when they appear and when they disappear, and to see a relationship between oxidation events and biological groups”, a researcher commented.
“This study supports the growing view that life and environment co-evolved through this tumultuous period of Earth history,” said geochemist Alan J. Kaufman, a co-author of the study from the University of Maryland.
The triggers for the oxidation events remain elusive, scientists are still not sure what set off these oxidizing events. Members of the research team have suggested that these two events recorded in marine sediments were probably related to oxygen in the atmosphere reacting with sediments on land as rocks are eroded away. The lack of biological activity on the land would have resulted in weathered rocks and soils on the continents releasing certain dissolved ions, such as sulphate, into rivers. These would then be transported to the sea where they might be used by bacteria to oxidize the organic carbon pool in the deep oceans.
This article has been adapted from materials published by Virginia Tech, USA. The full article entitled “Pulsed oxidation and biological evolution in the Ediacaran Doushantuo Formation,” was written by Kathleen A. McFadden; Jing Huang and Xuelei Chu of the Institute of Geology and Geophysics, Chinese Academy of Sciences; Ganqing Jiang; Alan J. Kaufman; Chuanming Zhou and Xunlai Yuan of the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences; and Shuhai Xiao.
It is due to be published in March.
CollectA have recently introduced a range of models of invertebrates reflecting iconic animals from the fossil record including trilobites and members of the Mollusca: Replicas of Iconic Fossil Animals, Models, Toys and Games.
Yorkshire Lad goes “Walking with Dinosaurs” (Dinosaur Footprints Discovered)
For many palaeontologists discovering a perfectly preserved set of dinosaur footprints may mark a high point in their careers but for young Rhys Nichols of Scarborough finding dinosaur trackways is as easy as taking a walk along the beach.
Whilst walking with his father, on the beach at Burniston Rocks, north of the seaside resort of Scarborough on the North Yorkshire coast, Rhys noticed that part of cliff face had fallen away and it was here that he found the footprints. Rhys’ very proud father, Richard stated that “Rhys loves dinosaurs so for him to find something like that was wonderful. He was over the moon – I couldn’t get him away from it!
Dinosaur Footprints
“We are always coming down here beach-combing and hunting for fossils.”
Experts have agreed that these fossil prints are a “great find”. The footprints are what is known as a trace fossil. Trace fossils preserve evidence of the activity of animals, such as their tracks; unlike many dinosaur fossil bones that may have been transported after death a long way from where the dinosaur originally lived, most trace fossils such as these prints are direct “in situ” evidence of the environment at the time and the place where the dinosaurs roamed.
Picture credit: Everything Dinosaur
Pictures show two beautifully preserved fossil footprints with the three toes of dinosaur seen clearly. The raised appearance of the fossil is typical of this sort of trace fossil. Footprint fossils can either be a depression-type fossil made by the weight of the animal or a cast of the “hole” made by the animal’s foot as it walked along. Sediments can fill the print up and it is these that are fossilised and the cast preserved giving the raised appearance. Mr Nichols measured the footprints estimating that they were around 21 cm in diameter, other footprints have been found but they had been heavily eroded.
Fossilised Trackways
A number of fossilised dinosaur trackways have been found in the North Yorkshire area , much of the coast of the North East from Scarborough to near Redcar is comprised of exposed areas of delta mudstone and sandstone and thin coals that were laid down in the Middle Jurassic approximately 160 million years ago. The rock strata where the prints have been found is well known for producing dinosaur trackways and isolated footprints, in fact geologists term this strata as the Burniston Footprint Bed. As blocks of silty sandstone fall onto the beach, split apart from the cliff face by erosion, these blocks frequently come to rest at the base of the cliff upside down revealing the finely detailed tracks of dinosaurs from the Jurassic period.
It is not known what actual species of dinosaur made the prints, as with most fossil trackways, unless the culprit is found fossilised at the end of the trackway, Ichnologists (scientists who specialise in studying tracks), can only speculate what sort of creature it was. Local museum staff have stated that this dinosaur may have been an Iguanodon. The three-toed prints are indicative of an Iguanodontidae, however, the mid Jurassic date is very early for such an animal, more normally associated with the Early Cretaceous. Perhaps it could be a trackway made by a dryosaur, these animals grew to lengths in excess of 3 metres, were relatively light and had a bi-pedal stance. Dryosaurs seem to have been relatively ubiquitous, with fossil being found in both the Northern and Southern Hemispheres. Like many trackways the scientists may just have to resort to referring to these wonderfully, well preserved prints as belonging to an “in-determinant ornithopod”.
Three Toes Clearly Observed
Photographs show a close up of the footprint shown in the foreground in the picture with Rhys. The animal would have been walking from right to left as the page is viewed. The three-toes can clearly be seen, but there is little evidence of a claw mark, adding weight to the thought that this is the footprint of an ornithopod. Based on comparisons with the fossils of ornithopods such as the large amount of Iguanodontidae material available, it has been estimated that the dinosaur walking across the delta 160 million years ago would have been roughly the same size as young Rhys. It is not known whether the animal was a fully grown adult or juvenile. The pictures of the footprints indicate a bipedal stance, but as to what actual animal made these tracks, this will probably remain a mystery, unless of course Rhys happens to find another set of prints whilst beach-combing but this time with the fossils of the dinosaur which made them at the end of the track.
Here’s hoping… in the meantime well done to Rhys, palaeontology remains the only science where by going for a walk you can change the way the world views itself.
Everything Dinosaur stocks a wide range of ornithopod models and figures: Everything Dinosaur – Dinosaur Models and Figures.
Giant Frog challenges Scientists over movement of Continents
Frogs are the most common type of amphibian alive today, with an estimated 5,500 separate species, making them the most diverse and successful clade of the Lissamphibians. They are known from all the continents except Antarctica but their fossil record is quite poor. Although very much extant, scientists still debate how many actual families make up the order containing frogs and toads – Anura. With discussion ongoing as to how to classify frogs and toads around today, it is no wonder that difficulties arise when trying to piece together the development and relationships between elements of Anura when you consider how sparse the fossil evidence is.
Now the discovery of a giant, Late Cretaceous frog from Madagascar that may be related to the horned frogs of South America, has opened up the debate once again over frog family ancestry and the break up of the super-continent Gondwanaland.
Frog and Toad Evolution
Frogs and toads are very specialised Lissamphibians with a body shape (morphology) unlike their living relatives and their ancient amphibian ancestors from the Palaeozoic. In comparison with other amphibian groups, they have dramatically reduced skeletons, lacking ribs, a tail, with a simple pelvic girdle and relatively few vertebrae. One of the earliest known frogs was also found in Madagascar, called Triadobatrachus; this animal dates back to the Triassic.
Frogs and toads were probably relatively abundant during the Mesozoic but the lack of fossil evidence inhibits palaeontologists when it comes to working out Anura evolution. Fossil bones have been recorded from a number of Mesozoic sites but they are usually isolated fragments, ilia, humeri (limb bones) and the more robust skull elements such as the frontoparietals and squamosals – elements from the top and towards the rear of the skull respectively. Some upper jaws bones (maxillae) have also been located and it is the jaws and the partial skull elements that provide the greatest assistance to palaeontologists when they attempt to work out the relationships between extinct genera and species.
Researchers Study Fossil Bones
Researchers from New York’s Stony Brook University aided by a team from University College, London headed up by vertebrate morphologist and palaeontologist Susan Evans; have published their findings on this new species of Madagascan giant frog in the journal Proceedings of the National Academy of Sciences.
This discovery, led by David Krause of Stony Brook University may undermine current scientific thinking over the isolation of Madagascar that was believed to have taken place in the Cretaceous. Conventional theory states that by approximately 95 million years ago, the land mass that was to eventually form India and Madagascar had split away from Africa, part of the break up of Gondwanaland (Australia, New Zealand, Africa, Antarctica and South America). Over the next 30 million years or so a rising plume of hot magma forced its way through a fault rifting apart India and Madagascar. Madagascar was left an isolated island with its own distinct indigenous fauna and flora and India went northwards to collide with Asia.
Beelzebufo ampinga
The fossilised bones of Beelzebufo ampinga, a frog the size of a partially deflated beach ball and tipping the scales at around 4 kilogrammes, making it the largest frog found to date, resemble the bones of the extant frog group – the Ceratophyrinae. The Ceratophyrinae, termed the “horned frogs” as many members of this family have soft extensions of skin growing out from the upper eyelid, which resemble little horns; are associated with South America.
Study of the fossils have indicated that this ancient animal is not related to any of the frog species living on Madagascar today. If Madagascar was very much an isolated island when Beelzebufo was hopping around, then how do scientists explain a member of the Ceratophyrinae group on an island thousands of miles away from their ancestral home of the South Americas.
Beelzebufo – the Frog from Hell
Picture credit: Associated Press
The diagram above shows an artist’s impression of Beelzebufo, with a modern frog and a pencil for scale. Although only partial elements of the skeleton have been recovered Krause and his team estimate that this animal was 40 cm long and would have weighed as much as a large domestic cat.
The most characteristic feature of the Ceratophyrinae is not their horned eyelids (some members of the group do not possess this feature), but their large heads, huge mouths and blunt snouts. They are voracious and unfussy hunters, lying half submerged in mud waiting for any unsuspecting small animal to wander by. Basically, anything that can fit into their mouths is on the menu, mice, frogs, snakes, fish and such like.
Wide Mouth and Powerful Jaws
Beelzebufo had a very wide mouth and powerful jaws, plus teeth. The skull material recovered has ridges and groves on it; perhaps indicating that this animal had bony armour or a protective head shield.
David Krause commented: “This frog, if it has the same habits as its living relatives in South America, was quite voracious. It’s even conceivable that it could have taken down some hatchling dinosaurs.”
The name Beelzebufo is a derivative of the Greek word for Devil and bufo is the Latin for toad. The “Devil Toad” would be an apt title for a frog capable of swallowing whole baby dinosaurs.
Krause and his team began finding fragments of abnormally large frog bones whilst studying the late Cretaceous sediments of the Mahajganga basin in north-western Madagascar in 1993. Amongst the various dinosaur and crocodilian fossils a total of 60 fossil frog bone fragments were located during a number of expeditions to the area by the New York team.
A Relative of Extant Horned Frogs
The unusually large frog bones were sent to the University College, London for specialist Susan Evans to examine. The London researchers were not able to piece together a complete skeleton but they had enough of the skull elements to make a diagnosis and interpret Beelzebufo as a relative of the horned frogs group.
The giveaway clinching evidence was the skull material indicating a “short, fat skull with a huge mouth”, says Evans.
Scale Drawing of Beelzebufo Skeleton compared to Living Frogs
Picture credit: Journal of Proceedings of the National Academy of Sciences
The drawing depicts the skeleton of Beelzebufo ampinga (A) compared to the largest extant member of the South American Ceratophyrs (B) and the largest frog species found on Madagascar today (C). The skeletal material in white represents bones found, those parts of Beelzebufo skeleton in grey are a scientific impression as to what the remainder of the skeleton would have looked like.
A Palaeontology Puzzle
The link to South America raises a palaeontology puzzle. Standard theory for how the continents drifted apart show what is now Madagascar would have been long separated by ocean from the Americas during Beelzebufo’s time. Frogs with their soft permeable skins cannot survive long in salt water, so reaching Madagascar by swimming can be ruled out.
Krause contends that the giant frog provides evidence for competing theories that some bridge still connected the land masses that late in time, perhaps via Antarctica that was much warmer than today. Perhaps Gondwanaland stayed together for longer than scientists currently think, could India/Madagascar have been linked to South America by an Antarctica land bridge as recently as the late Cretaceous.
Evans says that when she first began to suspect the Madagascar fragments came from a frog related to South American Ceratophryinae, she was very cautious about the claim. “We knew it would be controversial,” she says. “There are people who believe everything on Madagascar today must have been there when it broke with Gondwanaland 160 million years ago.”
Blair Hedges, a biologist at Pennsylvania State University in University Park, agrees that Beelzebufo is an important find. “The new fossil frog, besides being large and odd-shaped, is quite unexpected because of its apparent relationship with South American species,” he says.
But he says he isn’t yet convinced that the new find is related to the South American frogs. Molecular clock data suggests that these frogs split from a common ancestor more recently than 66 million years ago, he says. “Based on molecular evidence of frog relationships, the specific resemblance to some living wide-mouthed frogs is more likely from [evolutionary] convergence than actual relationship.” Convergent evolution, where unrelated species occupying similar niches tend to look the same, is common in frogs, he says.
Even if they are related, he adds, this doesn’t mean that the frogs necessarily had to walk on land from one location to another before Gondwana split. “Any organism, including a frog, can raft on dead vegetation,” he says.
Flood events and tropical storms can wash relatively large pieces of vegetation out to sea, some of these “rafts” get washed up on foreign shores. A number of animals migrate between islands today under these circumstances.
Susan Evans and her team remain convinced that Beelzebufo is a relative of the horned frogs of the Americas and refutes the convergence evolution theory. “It is the same family. I have no doubt of that,” she says.
It seems that this large, voracious frog – a trouble maker back in the Mesozoic, eating anything that could fit in its mouth, is going to be causing just as much trouble in scientific circles here in the Holocene.
Visti Everything Dinosaur’s website: Everything Dinosaur.
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New Ankylosaurian Cranial Remains From the Lower Cretaceous (Upper Albian) Toolebuc Formation of Queensland, Australia
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Australian dinosaur research has undergone a renaissance in the last 10 years, with growing knowledge of mid-Cretaceous assemblages revealing an endemic high...
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Frontiers
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https://www.frontiersin.org/journals/earth-science/articles/10.3389/feart.2022.803505/full
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Introduction
Ankylosaurians are a rare component of Gondwanan Cretaceous terrestrial ecosystems with five described species: Antarctopelta oliveroi, Minmi paravertebra, Kunbarrasaurus ieversi, Spicomellus afer, and Stegouros elengassen (Molnar, 1980; Salgado and Gasparini, 2006; Leahey et al., 2015; Maidment et al., 2021; Soto-Acuña et al., 2021). A. oliveroi and M. paravertebra have been considered nomen dubia due to a perceived lack of autapomorphies (Leahey et al., 2015; Arbour and Currie, 2016). However, comparisons between M. paravertebra and K. ieversi support their differentiation and the specific validity (Leahey et al., 2015). Despite the currently limited taxonomic diversity of Gondwanan Cretaceous ankylosaurs, skeletal fragments ascribed to the clade are found throughout Argentina (Coria and Salgado, 2001; Murray et al., 2019), New Zealand (Molnar and Wiffen, 1994), and Madagascar (Russel et al., 1976), with further ichnological evidence from Bolivia (Apesteguía and Gallina, 2011; Riguetti et al., 2021), and Brazil (Francischini et al., 2018). These widespread occurrences demonstrate that ankylosaurs were a rare yet pervasive component of Gondwanan dinosaur faunas. In Australia, ankylosaurian occurrences include two named taxa, M. paravertebra (Molnar, 1980) and K. ieversi (Molnar, 1996; Leahey et al., 2015), along with footprints (Salisbury et al., 2016) and several isolated skeletal elements that span most Australian dinosaur-bearing formations (Molnar, 1996; Molnar, 2001; Barrett et al., 2010; Leahey and Salisbury, 2013; Leahey et al., 2015; Bell et al., 2018a).
Despite brief mentions in the literature for almost 40 years, ankylosaurs from the middle–upper Albian marine Toolebuc Formation of central Queensland have never previously been studied in detail (Molnar, 1996; Leahey and Salisbury, 2013; Leahey et al., 2015). Three specimens were previously reported from two localities, Julia Creek (QM F33286) and Hughenden (AM F35259, AM F119849), and all were tentatively referred to the genus Minmi (Figures 1A,B; Molnar, 1996). QM F33286 is a partially disarticulated ankylosaur comprising “thoracic and pelvic” elements associated with ventral or lateroventral dermal ossifications (Molnar, 2001; Leahey and Salisbury, 2013). A brief report mentioned that the dermal ossicles were tightly arranged as square tiles that presumably covered most of the neck and trunk (Molnar, 2001). AM F35259 includes only a series of incomplete ribs with ossicles. AM F119849 likewise comprises a single block with vertebrae, ribs, and dermal ossifications (Leahey and Salisbury, 2013).
FIGURE 1
Here we provide the first comprehensive description of ankylosaur cranial remains recovered from the Toolebuc Formation based on a previously undocumented specimen, SAMA P40536, collected by BPK in 2005 from Warra Station near Boulia in western Queensland. SAMA P40536 is preserved within a series of yellow limestone concretions typical of the Toolebuc Formation (Kellner et al., 2010) but had eroded out and were dispersed throughout the blacksoil weathering residuum. The concretions contain remnants of the skull, pelvis, and limbs associated with isolated vertebrae, ribs, and dermal armor fragments. Our initial assessment of the remains focuses on the cranium and dentition, which are important because they represent only the second example of an ankylosaur skull recovered from Australia to date.
Institutional Abbreviations
QM, Queensland Museum, Brisbane, Queensland, Australia; SAMA, South Australian Museum, Adelaide, South Australia, Australia; AM, Australian Museum, Sydney, New South Wales, Australia.
Methods for Synchrotron Scanning and 3D Modeling
Microtomographic measurements of SAMA P40536 were performed using the Imaging and Medical Beamline (IMBL) at the Australian Nuclear Science and Technology Organisation’s (ANSTO) Australian Synchrotron, Melbourne, Victoria, Australia. For this investigation, acquisition parameters included a 40.29 × 40.29 μm pixel size, monochromatic beam energy of 70 keV, a sample–detector distance of 500 mm, and the “Ruby” detector. The latter consists of a PCO. edge sCMOS camera (16-bit, 2,560 × 2,160 pixels) and a Nikon Makro Planar 100 mm lens coupled with a 20 µm thick Gadox/CsI(Tl)/CdWO4 scintillator screen. As the height and width of the specimen exceeded the detector field-of-view, the specimen was aligned axially relative to the beam, its center of rotation shifted toward one edge of the detector. Twelve successive scans were required to cover the full specimen volume, each consisting of 1,500 equally spaced angle shadow radiographs with an exposure length of 0.35 s, obtained every 0.12° as the sample was continuously rotated 180° about its vertical axis. Horizontal translation of the specimen between tomographic scans was 80 mm, and 25 mm between vertical scans. One-hundred dark (closed shutter) and beam profile (open shutter) images were obtained for calibration before and after shadow-radiograph acquisition. The total time for the scan was 140 min.
The raw 16-bit radiographic series were normalized relative to the beam calibration files and stitched with the in-house software “IMBL-Stitch” to yield a 32-bit series with a 178 × 160 mm field-of-view. Reconstruction of the 3D dataset was achieved by the filtered-back projection method using the CSIRO’s X-TRACT (Gureyev et al., 2011). The image stack (Available on MorphoSource; ark:/87602/m4/392687) was segmented, and the volume data was rendered using Mimics Innovation Suite (v. 21.4; Materialise HQ, Leuven, Belgium). As various sections are preserved as bone impressions, with the original bone lost, thin surface models (∼0.8 mm thick) were added during segmentation to illustrate the relative placement of these bones. Finally, to create positive molds of the tooth row and properly study the dental anatomy, the negative left dentary tooth row impression was molded in Pinkysil® (fast set silicone) and cast in Easycast® (Rigid Polyurethane System). Manual preparation techniques using an air scribe (Paleotools® Micro Jack 3) were used to extract a single preserved tooth crown, subsequently scanned using the General Electric (GE) Phoenix V|tome|xs microCT scanner at the University of New England, Armidale. This tooth did not show in the original synchrotron scan.
Systematic Palaeontology
Dinosauria Owen, 1842
Ornithischia Seeley, 1888
Thyreophora Nopcsa, 1915
Ankylosauria Osborn, 1923
Kunbarrasaurus Leahey et al., 2015cf. Kunbarrasaurus sp.
Referred Material—SAMA P40536, a partial skull incorporating impressions of the left and right maxillae, the vomer, both palatines, the right (and part impression of the left) ectopterygoid, a possible right mandibular ramus, impressions of both maxillary tooth rows, and eight isolated teeth (Supplementary Materials S1, S2 available at figshare; https://figshare.com/articles/figure/3D_PDF_of_SAMA_P40536-_Frauenfelder_et_al_/16892908). Associated postcranial elements include parts of the pelvis, limb elements, vertebrae, ribs, scutes, and numerous dermal ossicles. However, much of the postcranial skeleton remains encased in limestone matrix and thus requires preparation before describing it.
Horizon and locality—The limestone concretions containing SAMA P40536 had weathered out of organic-rich marine shales that are lithostratigraphically representative of the Toolebuc Formation. This unit crops out over a vast area west of the township of Boulia in southwestern Queensland (Figure 1). The Toolebuc Formation strata around Boulia have long been recognized as abundant sources of vertebrate fossils with documented finds including chimaeroids, lamniform sharks, a diverse range of actinopterygians, ceratodont dipnoans (e.g., Lees, 1986; Rozefelds, 1992; Kemp, 1993; Bartholomai, 2004; Kear, 2007; Wilson et al., 2011; Bartholomai, 2012; Bartholomai, 2015a; Bartholomai, 2015b), marine reptiles incorporating elasmosaurid, polycotylid, and pliosaurid plesiosaurs, ophthalmosaurid ichthyosaurs and protostegid turtles (e.g., Kear, 2003; Kear, 2006; Kear and Lee, 2006; Zammit et al., 2010; Kear and Hamilton-Bruce, 2011; Kear, 2016; Kear et al., 2018), ornithocheiroid pterosaurs (Molnar and Thulborn, 1980; Molnar, 1987; Fletcher and Salisbury, 2010; Kellner et al., 2010, 2011; Pentland and Poropat, 2019), and enantiornithine birds (Molnar, 1986; Chiappe, 1996; Kurochkin and Molnar, 1997; Kear et al., 2003). These assemblages are associated with abundant pelagic cephalopod (ammonites, belemnites, and coleoids) and benthic invertebrate remains (as summarized in Henderson et al., 2000) that indicate a shallow off-shore marine setting subject to poorly oxygenated bottom water conditions (Henderson, 2004; Jiang et al., 2018). Chronostratigraphically, the Toolebuc Formation is correlated with the latest middle–upper Albian Coptospora paradoxa spore/pollen and Pseudoceratium ludbrookiae dinoflagellate zones (Figure 1C; McMinn and Burger, 1986; Moore et al., 1986).
Description
The skull of SAMA P40536 is incomplete and preserves a portion of the palatal region (Figure 2). The choanal width is 82 mm, similar to proposed values of 72 mm for K. ieversi (measured from Leahey et al., 2015 p. 22), suggesting a potential skull length of ∼270 mm and skull width of ∼280 mm. The holotype of K. ieversi was assumed to be near-mature to mature at the time of death based on its small size and lack of cranial fusion (Molnar, 1996; Leahey et al., 2015). Given the similarity in size, SAMA P40536 may have been of equivalent age.
FIGURE 2
Although not described herein, a large bony fragment is preserved on the ventral and right lateral surfaces, with the lateral extent unknown due to erosion. The bone is mediolaterally compressed anteriorly and forms a ventrally projecting hook-like process. We tentatively identify this fragment as belonging to the dentary due to its position, ventral to the maxillary tooth row. The paired choanae of SAMA P40536 are separated medially by the posterior portion of the vomer (Figure 2). They are crescent-shaped, curving away from the midline posterolaterally, and are slightly wider than they are long. Anteriorly, the vomer, maxillary tooth roots, and maxillary bone impressions form the boundary of both choanae. Typically, the secondary palate separates the choana from the maxillary tooth row (Bourke et al., 2018); however, here, these elements form the choanal boundary due to mediolateral compression of the maxillary tooth row. The palatines form the posterior boundary of the choanae, medially, along with the ectopterygoid, laterally (Figure 2). The contribution of ectopterygoid to the choanal boundary suggests that a “posteroventral” secondary palate, which generally braces the palatine against the medial surface of the maxilla, may not have been present (Vickaryous et al., 2004; but see Bourke et al., 2018 for a new interpretation of this palatal structure). The choanae are posteriorly positioned, with their preserved anterior extent roughly in line with the middle of the maxillary tooth row. This condition is similar to that seen in K. ieversi but differs from that of ankylosaurids and nodosaurids (Lee, 1996; Godefroit et al., 1999; Carpenter, 2004; Kilbourne and Carpenter, 2005; Leahey et al., 2015; Kinneer et al., 2016).
Facial Bones
Maxilla—The right and left ventromedial surfaces of the maxilla are preserved as impressions on the block; a small bony fragment of the left maxilla is also present along the anterior choanal margin (Figures 2B,C). The maxillary impressions form an arcuate contact with the ectopterygoid posteriorly and medially (Figure 3D), together forming the lateral and posterolaterally margins of the choanae, respectively. Anteriorly, the impressions enclose the posterior portion of the exposed maxillary tooth roots. Right, and left maxillary tooth rows are partially preserved as impressions of their medial surfaces (Figure 3C). The left maxillary tooth row is 82 mm in length, whilst the right is 90 mm. The tooth rows are straight for approximately two-thirds of their length, diverging posteriorly. The final third of the tooth row curves laterally, producing an overall curved tooth row in palatal view. An impression of the alveolar margin occurs on the left side of the block; however, it is not preserved on the right. Approximately 20 tooth impressions are preserved on the right tooth row, with replacement teeth visible dorsal to the impressions of the erupted teeth (Figure 3C). There are at least 15 preserved alveolar positions on the left tooth row. As both tooth rows are preserved as incomplete impressions, the total number of teeth within the maxillae is a minimum count.
FIGURE 3
Palatal Bones
Vomer—The vomer is an anteroposteriorly elongate and mediolaterally compressed bone; becoming dorsoventrally tall, posteriorly (Figure 4). Several portions of the nasal septum are missing, but this is likely due to poor mineralization (Witmer and Ridgley, 2008; Leahey et al., 2015). As a result, the dorsal and ventral extents of the vomer are unclear. The ventral nasal keel is partially preserved (Figure 4C). As observed in the CT scans, the anterior half of the vomer forms an elongate triangular process that tapers to a point anteriorly (Figure 4A). This process is dorsoventrally compressed (Figure 4C) with an inverted triangular cross-section and, unlike the posterior half, does not divide the nasal passage. At roughly its mid-point and the mediolateral choanal corner, the vomer mediolaterally expands and is bulbous in dorsal view (Figure 2C). A posterodorsally oriented groove is present on the left lateral surface of the expanded region. This groove is not present on the right lateral surface and is a notable asymmetrical feature (Figures 4C,D). Posterior to the posterodorsal groove, the vomer expands laterally, resulting in an hourglass shape in dorsal view (Figures 2B, 4A).
FIGURE 4
Palatines—The palatines form the posteromedial margins of the choanae and are preserved mostly as impressions, with the right preserving a fragment of bone (Figures 2C, 3B). The palatine is a thin, anteroventrally/posterodorsally angled element, similar to other ankylosaurs, such as Ankylosaurus magniventris and Gargoyleosaurus parkpinorum (Carpenter, 2004; Kilbourne and Carpenter, 2005). Together, the palatines form a U-shaped wall in posterior view that would have separated the palatal and orbital regions, similar to Pawpawsaurus campbelli (Figures 2C, 3B; Lee, 1996). Medially, the palatines contact the vomer. Examining the CT data and physical specimen, we cannot confirm if the palatines and vomer are fused (Figures 2B,C). The palatines extend posterior to the vomer, suggesting that they contacted each other for part of their length; however, the medial edges of the palatines are missing. Along the posterior margin of the choanae, the palatines contact the ectopterygoid along a straight suture that is angled ventrolaterally in posterior view, similar to Ankylosaurus (Figure 3B; Carpenter, 2004).
Ectopterygoid—A partial ectopterygoid is preserved as both bone and impression on the right lateral surface of the skull block, and the left ectopterygoid is partially preserved as an impression (Figures 2C, 3B). The ectopterygoid is sub-triangular in mediolateral view (Figures 5A,B), curved medially, with a concave dorsal margin forming the choanal posterolateral edge (Figures 2B, 5E). The ectopterygoid contacts the maxilla anteriorly, but the contact would have continued laterally to the ectopterygoid along a scarf joint (Figure 6). The ectopterygoid contacts the palatine medially, along the choanal posterior margin, and forms a straight joint angled ventrolaterally (Figure 6). The posteroventral corner is drawn out into a wedged-shaped process (as preserved) buttressed medially along its length (Figure 5).
FIGURE 5
FIGURE 6
Dentition
Eight isolated teeth are identified from the CT scans: one in situ left maxillary tooth crown and seven isolated teeth with roots of unknown position found “floating” within the matrix (Table 1; Supplementary Materials S1, S2). Of these, the in situ crown provides the best resolution and the basis for the following description (Figures 7H–J). The crown is asymmetrical in labial view and is labiolingually compressed (Figures 7I,J). The apex corresponds to the primary ridge/denticle (sensu Bell et al., 2018b) and is distally offset. The primary denticle is marginally larger than the remaining denticles and bears a shallow, mesiodistally oriented, and grooved wear facet on the apex. Two denticles are present distal to the primary denticle and three mesially. Denticles are apically pointed and curved away from the central tooth axis, giving the crown a fan-shaped appearance. Apicobasal grooves (or sulci) between denticles extend basally but do not appear to have reached the cingulum (Figure 7; the cingulum itself is only preserved on the more complete isolated teeth; see above). Individual denticles are difficult to identify on the isolated teeth due to the low resolution of the CT scans; however, small bumps on the crowns suggest five to seven denticles (including the primary denticle) were present per tooth (Figures 7A,E).
TABLE 1
FIGURE 7
The low denticle count is similar to K. ieversi and other nodosaurids but differs from the approximately 8–12 denticles per tooth seen in the upper Strzelecki group teeth (Molnar, 1996; Barrett et al., 2010; Leahey et al., 2015). The in situ crown is broken basally, and the presence of cingula are unknown; however, cingula are present on five isolated teeth. The lingual cingulum is prominent, forming a bulbous shelf, whereas the labial cingulum is less prominent and positioned apically (Figures 7A–F). Five teeth preserve large roots, two to three times taller than the crowns (Table 1), separated by a constriction. Roots are straight, bullet-shaped, and circular in cross-section, similar to other Australian ankylosaurian teeth (Figure 7G; Molnar, 1996; Barrett et al., 2010; Leahey and Salisbury, 2013). Two isolated teeth have partially resorbed roots, as the labial surface is either partially or completely missing (Figure 7B), suggesting root resorption proceeded in a labial–lingual direction. All tooth impressions from the right maxilla have prominent lingual cingula. Consistent with the in situ maxillary crown, tooth impressions have anywhere from five to seven denticles. Crowns of the 16 mesial-most teeth are fan-shaped, whereas the four distal-most teeth are more pointed (Figure 3C). The preserved crown impressions range from 4 to 6 mm wide and 5–8 mm long.
Phylogenetic Analysis
Methods
SAMA P40536 was scored into two phylogenetic datasets (Supplementary Material S3): Arbour and Currie (2016) (Modified from Thompson et al., 2012; Arbour and Currie, 2013; Arbour et al., 2014a; Arbour et al., 2014b) and Soto-Acuña et al. (2021) (Modified from Arbour and Currie, 2016; Arbour et al., 2016). The Arbour and Currie (2016) matrix consists of 178 characters and 45 ingroup taxa, including SAMA P40536 (scores provided in Table 2) and the non-ankylosaurian outgroup taxa Lesothosaurus diagnosticus, Sceliosaurus harrisonii, and Huayangosaurus taibaii. In this matrix, we updated the terminal taxon designation from Minmi sp. to Kunbarrasaurus ieversi, and “Zhejiangosaurus lishuiensis” replaced the incorrectly identified “Zhejiangosaurus luoyangensis” (Lü et al., 2007; Leahey et al., 2015). The Soto-Acuña et al. (2021) matrix consists of 190 characters and 66 ingroup taxa, including SAMA P40536 (scores provided in Table 2) and the non-ankylosaurian outgroup taxa Lesothosaurus diagnosticus, Scelidosaurus harrisonii, and members of Stegosauria (Huayangosaurus taibaii, Paranthodon africanus, and Stegosaurus stenops). Here, we updated the terminal taxon designation from Sauropelta edwardsi to Sauropelta edwardsorum, and “Argentinian ankylosaur” replaced the “Salitral Moreno ankylosaur” for consistency between matrices.
TABLE 2
To incorporate additional palatal variations noted in SAMA P40536, K. ieversi, and other ankylosaurians, we added a new character (178 in Arbour and Currie, 2016; and 190 in Soto-Acuña et al., 2021) that describes the position of the choanae within the palate relative to the maxillary tooth row: choanae with their anterior margins inline or within the anterior third of the maxillary tooth row (178/190:0); choanae posteriorly situated with their anterior margins at least mid-way along the tooth row (178/190:1). Ankylosaurians were scored from the literature (Eaton Jr, 1960; Sereno and Zhimin, 1992; Lee, 1996; Godefroit et al., 1999; Carpenter et al., 2001a, 2008, 2011; Vickaryous et al., 2001; Hill et al., 2003; Carpenter, 2004; Kilbourne and Carpenter, 2005; Parsons and Parsons, 2009; Arbour and Currie, 2013; Arbour et al., 2014a; Leahey et al., 2015; Kinneer et al., 2016; Arbour and Evans, 2017; Yang et al., 2017; Bourke et al., 2018; Paulina-Carabajal et al., 2018; Wiersma and Irmis, 2018; Norman, 2020; Park et al., 2020); however, we could not adequately code the outgroup taxon Huayangosaurus taibaii because the condition of its choanae is unknown. Character polarity was therefore determined from Hesperosaurus mjosi (Maidment et al., 2018) and a 3D cranial model of Stegosaurus armatus (specimen number; UMNH VPC 44, sketchfab. com/ivlpaleontology), which preserve the choanae approximately in line with the anterior-most maxillary tooth (Sereno and Zhimin, 1992; Carpenter et al., 2001b; Chengkai et al., 2007; Mateus et al., 2009). The anterior extent of the choanae in Lesothosaurus diagnosticus is likewise not directly observable. Nonetheless, the maxillae are mediolaterally compressed and unlikely to have formed a bony palate as in many ankylosaurs (Porro et al., 2015). Consequently, we interpret the anterior margins of the choana as probably being anteriorly placed. Finally, the choanae of Scelidosaurus harrisonii end anteriorly relative to the anterior-most maxillary tooth (Norman, 2020). Character 31 was recoded as “?” for K. ieversi in both matrices because the placement of the ectopterygoid cannot be adequately interpreted from the physical specimen or 3D tomographic renderings (Leahey et al., 2015). Moreover, given the absence of a “caudoventral” secondary palate in SAMA P40536, it is doubtful that such as structure was present in K. ieversi.
The updated character-taxon matrices were compiled using Mesquite version 3.61 (Maddison and Maddison, 2019) and analyzed in TNT version 1.5 (Goloboff and Catalano, 2016). All characters were assumed unordered and of equal weight, with Lesothosaurus diagnosticus designated the most distant outgroup taxon in both matrices. Both matrices were subjected to a phylogenetic analysis involving 1,000 replicates of a “traditional” search using random addition sequence starting trees and the Tree Bisection Reconnection (TBR) branch swapping algorithm. Following the initial search, we performed another round of branch swapping on the set of most parsimonious trees (MPTs) using TBR to more fully explore the tree space. We used iterative PCR to identify wildcard taxa that could be pruned to improve resolution in the strict consensus of the final set of MPTs (Pol and Escapa, 2009), while retaining SAMA P40536. Nodal supports for the resulting reduced strict consensus tree from each matrix were calculated from 1,000 bootstrap resampling replicates using the same initial “traditional” tree search strategy. Clade frequencies were summarised using the Groups present/Contradicted (GC) metric.
Results
The phylogenetic analysis of Arbour and Currie (2016) returned 1,630 MPTs of 423 steps [Consistency Index (CI): 0.536, Retention Index (RI): 0.704, Rescaled Consistency Index (RC): 0.377] from the initial tree search, and over 10,000 additional trees of equal length after another round of branch swapping. Sixteen wildcard taxa were identified for removal by iterative PCR: Aletopelta coombsi, Antarctopelta oliveroi, Bissektipelta archibaldi, Dongyangopelta yangyanensis, Glyptodontopelta mimus, Gobisaurus domoculus, Liaoningosaurus paradoxus, Minmi paravertebra, Pawpawsaurus campbelli, Sauroplites scutiger, Scolosaurus cutleri, Shamosaurus scutatus, Stegopelta landerensis, Taohelong jinchengensis, Zhejiangosaurus lishuiensis, and Zipaleta sanjuanensis. The reduced strict consensus tree is almost completely resolved (Figure 8A); Ankylosauridae is fully resolved, whilst Nodosauridae contains one polytomy (Sauropelta edwardsorum, Tianchisaurus nedegoapeferima, and the Argentinian ankylosaur). SAMA P40536 was recovered as the sister of K. ieversi, which together form the sister clade to Mymoorapelta maysi + Ankylosauridae + Nodosauridae (Figure 8A). Kunbarrasaurus ieversi and SAMA P40536 share two autapomorphies: the maxillary tooth row is medially convex (28:1), and the choanae are posteriorly placed approximately mid-way along the maxillary tooth row (178:1). As M. paravertebra was removed by IterPCR, the relationship with SAMA P40536 is currently unknown; however, when the analysis was run with M. paravertebra included, the tree collapsed into a large polytomy (results not included).
FIGURE 8
The initial search of Soto-Acuña et al. (2021) returned 20 MPTs of 693 steps (CI: 0.359, RI: 0.653, RC: 0.234). More than 10,000 trees of equal length were found after another round of branch swapping. Seven wildcard taxa were identified for removal by iterative PCR: Ahshislepelta minor, Denversaurus schlessmani, Donyangopelta yangyanensis, Hylaeosaurus armatus, Sauroplites scutiger, Taohelong jinchengensis, and Zhejiangosaurus lishuiensis. Both Ankylosauridae and Nodosauridae are well resolved in the resulting reduced strict consensus tree (Figure 8B), each containing one polytomy (Ziapelta sanjuanensis and Anodontosaurus lambei, and all three Struthiosaurus, respectively). Parankylosauria is monophyletic but unresolved, recovered as the sister-taxon to Ankylosauridae, Nodosauridae, and a clade formed by Cedarpelta bilbeyhallorum, Chuanqilong chaoyangensis, and Liaoningosaurus paradoxus. Parankylosauria contains Stegouros elengassen, K. ieversi, and A. oliveroi, as in Soto-Acuña et al. (2021), along with SAMA P40536.
Discussion
Comparisons with Kunbarrasaurus ieversi
SAMA P40536 shares five features with Kunbarrasaurus ieversi: a sinuous maxillary tooth row, posteriorly placed choanae, asymmetric tooth crowns, and tooth crown striations that are both confluent with the denticles and extend to the cingulum (Table 2 for clarification on matrix assignment). In addition, SAMA P40536 and K. ieversi share teeth with low denticle counts, cylindrical tooth roots, and both lingual and labial cingula (Molnar, 1996). Unfortunately, none of the K. ieversi autapomorphies identified on the holotype (QM F18101; Leahey et al., 2015) are preserved in SAMA P40536, precluding unambiguous referral. Nevertheless, their dental and choanal similarities are sufficient to recover both specimens as sister-taxa (Figure 8A) and within Parankylosauria with the added tooth characters (Figure 8B). Combined with their spatiotemporal proximity (Figure 9), we provisionally refer SAMA P40536 to cf. Kunbarrasaurus sp. pending further preparation and description of the postcranial skeletons.
FIGURE 9
Although our taxonomic assignment is tentative, the palatal osteology of SAMA P40536 fills anatomical gaps not observable in QM F18101 (Leahey et al., 2015). For example, the palatines of SAMA P40536 are vertically positioned and separate the palatal and orbital regions. Although the posterior extent of the palatines is not preserved, vertical sutures along the posterior margins of the choanae represent the contacts between the palatines and the ectopterygoids (Figures 3A,B). This arrangement indicates that the ectopterygoid forms the posterolateral margin of the choana, as in Edmontonia longiceps (Vickaryous et al., 2004). Furthermore, the palatines did not contact the maxillae along the choanal margin as reconstructed by (Leahey et al., 2015). The ectopterygoid contacts are unknown in K. ieversi due to poor preservation and encasing matrix (Leahey et al., 2015) but differ from the interpretation of (Leahey et al., 2015, p. 22, Figure 6), who placed the ectopterygoid more posteriorly between the maxilla and pterygoids.
Implications of Choanal Variation in Ankylosaurs
The palatal choanae form part of the nasal passages and are the boundary between the internal cranial nasal passages and the buccal region (Bourke et al., 2018). Therefore, it may be expected that choanal variations reflect the complexities of ankylosaurian nasal passages, such as mineralised soft tissue creating convoluted nasal passages and the paranasal sinus system formed by an extensive set of air sacs surrounding the nasal airways. (Brown and Kaisen, 1908; Vickaryous et al., 2004; Vickaryous, 2006; Witmer and Ridgley, 2008; Leahey et al., 2015; Paulina-Carabajal et al., 2016; Bourke et al., 2018). Surprisingly, the palate remains an anatomically under-sampled component of ankylosaur phylogenies. Indeed, only five of the 178 and 190 characters employed in both matrices capture palatal morphologies. We found notable variation in the relative placement of the choana within the palate. The majority of ankylosaurians display choanae that span most of the palatal region, and the anterior choanal margins are either in line with the anterior-most maxillary tooth (e.g., Pawpawsaurus campbelli; Paulina-Carabajal et al., 2016 p. 5) or within the anterior third of the tooth row (e.g., Ankylosaurus magniventris; Carpenter, 2004). As we used the literature to code most ankylosaurian taxa, we simplified the previous conditions into a single character relative to the outgroup (Sereno and Zhimin, 1992; Porro et al., 2015; Maidment et al., 2018; Norman, 2020), indicating that anteriorly placed choanae are primitive for ankylosaurs. By contrast, SAMA P40536 and K. ieversi exhibit a derived condition whereby the choanae are relatively posterior within the palate. The other parankylosaurian Stegouros ellengassen may also exhibit this derived condition, as the secondary palate that marks the anterior extent of the choanae extends posteriorly to approximately the mid-point of the preserved tooth row (Soto-Acuña et al., 2021). Unfortunately, the maxillary tooth row is incomplete posteriorly and so we cannot confirm its condition at this time, and it was coded as “?”. The derived condition stabilizes SAMA P40536 within both trees but otherwise does not dramatically affect the phylogenetic placements of other ankylosaurians. Without this character, the trees collapse into a large polytomy with virtually no basal resolution (results not provided). It is worth noting that if future postcranial observations of SAMA P40536 support an assignment to K. ieversi, this character may represent a new autapomorphy for K. ieversi.
Presently, only the ankylosaurids Cedarpelta bilbeyhallorum, and Gobisaurus domoculus (Vickaryous et al., 2001; Carpenter et al., 2008) exhibit posteriorly positioned choanae comparable to K. ieversi (Leahey et al., 2015) and SAMA P40536. The ankylosaurid Akainacephalus johnsoni and the nodosaurid Panoplosaurus mirus show the most extreme condition, whereby the anterior choanal margins are approximately in line with the posterior-most maxillary tooth (Bourke et al., 2018; Wiersma and Irmis, 2018). These occurrences suggest that posteriorly located choanae evolved independently at least four times: once before the Nodosauridae + Ankylosauridae split, once in nodosaurids, and at least twice in ankylosaurids. Note that there is currently no resolution on the relationships of C. bilbeyhallorum and G. domoculus in Arbour and Currie (2016) and that our safe taxonomic reduction analysis of their matrix excluded G. domoculus. However, our analysis of the Soto-Acuña et al. (2021) matrix places C. bilbeyhallorum outside of Ankylosauridae + Nodosauridae (Figure 8B), which would incur a more complex evolutionary path for the character. However, we refrain from over interpreting this tree given its labile nature (See next Section).
The phylogenetic placement of SAMA P40536 as an early-branching ankylosaurian has implications for interpreting choanal and palatal variations in ankylosaurs. The bones forming the choanal margins are often difficult to discern due to a tightly sutured or fused nature. The preserved palatal bones of SAMA P40536 are not fused, and contacts are visible. The choanal margins are formed by four bones, the maxilla, vomer, ectopterygoid, and palatines (Figure 2). Given that the ectopterygoid contributes to the choanal margin, the palatines were excluded from contacting the maxilla, suggesting that SAMA P40536 did not have a “posteroventral” secondary palate (Vickaryous et al., 2004) or lamina transversa (Bourke et al., 2018); these contacts are unknown in K. ieversi (Leahey et al., 2015). Therefore, our results indicate that the ankylosaurian lamina transversa evolved later in ankylosaur evolution, perhaps due to modifications to the nasal passages (Bourke et al., 2018). Functionally, the lamina transversa separates the olfactory and nasal regions and is associated with a heightened sense of smell (Bourke et al., 2018). Therefore, its absence in SAMA P40536 suggests that a more basic sense of smell was primitive for ankylosaurians.
Implications for Australian Ankylosaur Diversity
Australia has the largest abundance of Gondwanan ankylosaurs and species-level diversity documented from the mid-Cretaceous (Figure 9; Molnar, 2001; Barrett et al., 2010; Leahey and Salisbury, 2013; Leahey et al., 2015; Salisbury et al., 2016; Bell et al., 2018a). Occurrences include ankylosaur footprints from the Valanginian–Barremian Broome Sandstone of Western Australia (Salisbury et al., 2016), isolated skeletal elements from the uppermost Barremian–lower Aptian “Wonthaggi formation” strata of the upper Strzelecki Group in Victoria, Minmi paravertebra from the lower Aptian Bungil Formation of Queensland, Kunbarrasaurus ieversi from the upper Albian Allaru Mudstone of Queensland, and other indeterminate bones and teeth from the uppermost Albian–lower Cenomanian Mackunda and Griman Creek formations of Queensland and New South Wales, respectively (Molnar, 2001; Barrett et al., 2010; Leahey and Salisbury, 2013; Leahey et al., 2015; Bell et al., 2018a). Historically, all Australian ankylosaur fossils were referred to the genus Minmi (Molnar, 1980, 1996); however, the re-classification of K. ieversi (previously Minmi sp.; Leahey et al., 2015) as a separate genus and species implicates greater intra-clade diversity. Our attribution of SAMA P40536 to cf. Kunbarrasaurus sp. further extends the stratigraphical range of this taxon into the lower–upper Albian and evinces a novel sampling occurrence some ∼550 km to the SW of other earlier discoveries.
Previous phylogenies treated K. ieversi (QM F18101) and M. paravertebra (QM F10329) as a generic hypodigm, recovering it as either a sister to all other ankylosaurians (Kirkland, 1998; Carpenter, 2001) or ankylosaurids (Sereno, 1999; Hill et al., 2003; Vickaryous et al., 2004; Ősi, 2005; Burns et al., 2011; Thompson et al., 2012). However, K. ieversi is now considered an early-branching ankylosaurian outside of both Ankylosauridae and Nodosauridae (Arbour and Currie, 2016). Recently, the discovery of Stegouros ellengassen introduced a new phylogenetic hypothesis, whereby K. ieversi, S. ellengassen, and Antarctopelta oliveroi form a Gondwanan ankylosaur clade, Parankylosauria, which would have diverged before the Ankylosauridae + Nodosauridae split (Soto-Acuña et al., 2021). In testing the phylogenetic position of SAMA P40536, we were unable to reproduce the same tree from Soto-Acuña et al. (2021), even when the aforementioned specimen was removed. Our replication attempt produced 20 MPTs and contained several differences in the strict consensus, such as a polytomy outside of Ankylosauridae + Nodosauridae containing Chuanqilong chaoyangensis, Lianingosaurus paradoxus and Cedarpelta bilbeyhallorum, and slightly better resolution within Ankylosauridae and Nodosauridae (Supplementary Material S3; Supplementary Figure S3.2). Furthermore, an additional round of branch swapping on the initial 20 MPTs produced over 10,000 MPTs with considerably reduced resolution in the strict consensus. For instance, there was no clear split between Ankylosauridae and Nodosauridae and most ingroup taxa were reduced to a polytomy (Supplementary Material S3; Supplementary Figure S3.3). The reason for the difference in phylogenetic hypothesis observed in our replication of Soto-Acuña et al. (2021) is unclear; however, it likely stems from the overall poorly supported nature of ankylosaur phylogenetics. Fortunately, these broader differences had no effect on the phylogenetic placement of SAMA P40536 and Parankylosauria was nonetheless recovered in all analyses, supporting the existence of an early-branching, exclusively Gondwanan ankylosaurian clade. Given the placement of SAMA P40536 as the sister taxon of K. ieversi in the Arbour and Currie (2016) phylogeny, it is not surprising that it is also occurs within Parankylosauria. All four terminal units share four dental features: asymmetric tooth crowns, striations confluent with the denticles and extend to the cingula, and cingulum present on either maxillary or dentary teeth. Of note, the first three features are also present in the Argentinian ankylosaur (Coria and Salgado, 2001), although current hypotheses place it deep within Nodosauridae (Figure 8).
Previous studies hypothesized that Australian ankylosaurians evolved independently from the Late Cretaceous ankylosaurs found elsewhere in Gondwana (i.e., A. oliveroi and the Argentinian ankylosaur), which were previously considered to be more closely related to Laurasian nodosaurids (Figure 8A; Coria and Salgado, 2001; Salgado and Gasparini, 2006; Arbour and Currie, 2016; Arbour et al., 2016). However, the proposed existence of Parankylosauria suggests that most Gondwanan ankylosaurs are more closely related to each other than to those found elsewhere (Soto-Acuña et al., 2021). The only current exception is the Argentinian ankylosaur, a nodosaurid (Figure 8) that appeared in Gondwana following dispersal events from Laramidia during the Campanian–Maastrichtian, a pattern also observed among hadrosaurids and titanosaurs (Brett-Surman, 1979; Coria and Salgado, 2001; Prieto-Márquez, 2010; Arbour and Currie, 2016; Ibiricu et al., 2021). Given the mid–upper Albian ages of Australian ankylosaurs, the Gondwanan dispersal of Parankylosauria considerably predated those of the latest Cretaceous (Arbour and Currie, 2016; Kubo, 2020; Soto-Acuña et al., 2021). Interestingly, the recent identification of the putative ankylsoaur Spicomellus afer from the mid-Jurassic of Morocco (Maidment et al., 2021) hints at an initial, more ancient global diversification of Ankylosauria (Gibbons et al., 2013; Arbour and Currie, 2016; Arbour et al., 2016; Maidment et al., 2021; Soto-Acuña et al., 2021).
Conclusion
Here, we described the second Australian ankylosaur cranium and the first ankylosaurian remains from the Toolebuc Formation. Its conferral to Kunbarrasaurus is based on palatal and dental similarities. However, without extensive overlapping anatomy, an unambiguous referral is currently not possible. Nonetheless, several skull elements unknown in the holotype of K. iveresi can be inferred by SAMA P40536, notably the position and morphology of the palatines and their relation to the ectopterygoids. The future examination of SAMA P40536’s postcranial skeleton with those of K. ieversi and Minmi paravertebra will elucidate its taxonomic affinities, further testing the phylogenetic affinities of Australian and Gondwanan ankylosaurs. The exploration of palatal morphology uncovered a new phylogenetic character, and highlights the importance of this anatomical region in resolving ankylosaur phylogenetic relationships.
Data Availability Statement
Raw data and 3D models are available at MorphoSource (ark:/87602/m4/392687): https://www.morphosource.org/projects/000392635?locale=en. Supplementary Material (3D PDF) S1 and S2 are available at Figshare: https://figshare.com/articles/figure/3D_PDF_of_SAMA_P40536-_Frauenfelder_et_al_/16892908.
Author Contributions
TGF and NEC conceived and designed the study, and prepared figures or tables. TGF, PRB, and NEC wrote the manuscript. TGF, PRB, TB, and NEC analyzed the data. JJB scanned fossil material and wrote the associated methods. RDCB created the 3D PDFs. BPK collected fossil material. SW provided access to segmenting software. All authors edited and commented on the manuscript, figures, and tables.
Funding
Synchrotron scanning was facilitated by an IMBL beamtime application (15769) awarded to RDCB and TB. RDCB and TB are financed by UNE Postdoctoral Research Fellowships. This study was funded by a UNE Research Training Program Scholarship to TGF and an Australian Research Council Discovery Early Career Award (DE190101423) to NEC.
Conflict of Interest
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
Publisher’s Note
All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.
Acknowledgments
We thank Mary-Anne Binnie (SAMA) for providing access to study SAMA P40536. Christopher Goatley [University of New England (UNE)] assisted with Micro-CT scanning the in situ maxillary tooth. Anton Maksimenko provided technical assistance with the synchrotron scan data through his IMBL-Stitch software. TNT is made freely available thanks to a subsidy from the Willi Hennig Society. Special thanks to members of the Palaeoscience Research Centre at UNE for discussions related to this study, in particular Kai Allison, Marissa Betts, and John Paterson. Finally, we thank W Zheng, RT Tucker, and one anonymous reviewer for their constructive feedback. BPK acknowledges an Australian Research Council Linkage Project grant (LP0453550), which funded the excavation of SAMA P40536.
Supplementary Material
The Supplementary Material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/feart.2022.803505/full#supplementary-material
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Ankylosaurus is a genus of armored dinosaur. Its fossils have been found in geological formations dating to the very end of the Cretaceous Period, about 68–66 m...
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Not to be confused with Ankylosuchus.
Ankylosaurus[nb 1] is a genus of armored dinosaur. Its fossils have been found in geological formations dating to the very end of the Cretaceous Period, about 68–66 million years ago, in western North America, making it among the last of the non-avian dinosaurs. It was named by Barnum Brown in 1908; it is monotypic, containing only A. magniventris. The generic name means "fused" or "bent lizard", and the specific name means "great belly". A handful of specimens have been excavated to date, but a complete skeleton has not been discovered. Though other members of Ankylosauria are represented by more extensive fossil material, Ankylosaurus is often considered the archetypal member of its group, despite having some unusual features.
Quick Facts Scientific classification, Binomial name ...
Ankylosaurus
Cast of Ankylosaurus skull (AMNH 5214) in front view, Museum of the Rockies Scientific classification Domain: Eukaryota Kingdom: Animalia Phylum: Chordata Clade: Dinosauria Clade: †Ornithischia Clade: †Thyreophora Clade: †Ankylosauria Family: †Ankylosauridae Subfamily: †Ankylosaurinae Tribe: †Ankylosaurini Genus: †Ankylosaurus
Brown, 1908 Species:
†A. magniventris
Binomial name †Ankylosaurus magniventris
Brown, 1908
Close
Possibly the largest-known ankylosaurid, Ankylosaurus is estimated to have been between 6 and 8 meters (20 and 26 ft) long and to have weighed between 4.8 and 8 metric tons (5.3 and 8.8 short tons). It was quadrupedal, with a broad, robust body. It had a wide, low skull, with two horns pointing backward from the back of the head, and two horns below these that pointed backward and down. Unlike other ankylosaurs, its nostrils faced sideways rather than towards the front. The front part of the jaws was covered in a beak, with rows of small, leaf-shaped teeth farther behind it. It was covered in armor plates, or osteoderms, with bony half-rings covering the neck, and had a large club on the end of its tail. Bones in the skull and other parts of the body were fused, increasing their strength, and this feature is the source of the genus name.
Ankylosaurus is a member of the family Ankylosauridae, and its closest relatives appear to be Anodontosaurus and Euoplocephalus. Ankylosaurus is thought to have been a slow-moving animal, able to make quick movements when necessary. Its broad muzzle indicates it was a non-selective browser. Sinuses and nasal chambers in the snout may have been for heat and water balance or may have played a role in vocalization. The tail club is thought to have been used in defense against predators or in intraspecific combat. Specimens of Ankylosaurus have been found in the Hell Creek, Lance, Scollard, Frenchman, and Ferris formations, but it appears to have been rare in its environment. Although it lived alongside a nodosaurid ankylosaur, their ranges and ecological niches do not appear to have overlapped, and Ankylosaurus may have inhabited upland areas. Ankylosaurus also lived alongside dinosaurs such as Tyrannosaurus, Triceratops, and Edmontosaurus.
In 1906, an American Museum of Natural History expedition led by American paleontologist Barnum Brown discovered the type specimen of Ankylosaurus magniventris (AMNH 5895) in the Hell Creek Formation, near Gilbert Creek, Montana. The specimen (found by collector Peter Kaisen) consisted of the upper part of a skull, two teeth, part of the shoulder girdle, cervical, dorsal, and caudal vertebrae, ribs, and more than thirty osteoderms (armor plates). Brown scientifically described the animal in 1908; the generic name is derived from the Greek words αγκυλος ankulos ('bent' or 'crooked'), referring to the medical term ankylosis, the stiffness produced by the fusion of bones in the skull and body, and σαυρος sauros ('lizard'). The name can be translated as "fused lizard", "stiff lizard", or "curved lizard". The type species name, magniventris, is derived from the Latin: magnus ('great') and Latin: venter ('belly'), referring to the great width of the animal's body.[2][3][4]
The skeletal reconstruction accompanying the 1908 description restored the missing parts in a fashion similar to Stegosaurus, and Brown likened the result to the extinct armored mammal Glyptodon.[2] In contrast to modern depictions, Brown's stegosaur-like reconstruction showed robust forelimbs, a strongly arched back, a pelvis with prongs projecting forwards from the ilium and pubis, as well as a short, drooping tail without a tail club, which was unknown at the time. Brown also reconstructed the armor plates in parallel rows running down the back; this arrangement was purely hypothetical. Brown's reconstruction became highly influential, and restorations of the animal based on his diagram were published as late as the 1980s.[5][6][7] In a 1908 review of Brown's Ankylosaurus description, the American paleontologist Samuel Wendell Williston criticized the skeletal reconstruction as being based on too few remains, and claimed that Ankylosaurus was merely a synonym of the genus Stegopelta, which Williston had named in 1905. Williston also stated that a skeletal reconstruction of the related Polacanthus by Hungarian paleontologist Franz Nopcsa was a better example of how ankylosaurs would have appeared in life.[8] The claim of synonymy was not accepted by other researchers, and the two genera are now considered distinct.[9]
Brown had collected 77 osteoderms while excavating a Tyrannosaurus specimen in the Lance Formation of Wyoming in 1900. He mentioned these osteoderms (specimen AMNH 5866) in his description of Ankylosaurus but thought they belonged to the Tyrannosaurus instead. Paleontologist Henry Fairfield Osborn also expressed this view when he described the Tyrannosaurus specimen as the now synonymous genus Dynamosaurus in 1905. More recent examination has shown them to be similar to those of Ankylosaurus; it seems that Brown had compared them with some Euoplocephalus osteoderms, which had been erroneously cataloged as belonging to Ankylosaurus at the AMNH.[10][11]
In 1910, another AMNH expedition led by Brown discovered an Ankylosaurus specimen (AMNH 5214) in the Scollard Formation by the Red Deer River in Alberta, Canada. This specimen included a complete skull, mandibles, the first and only tail club known of this genus, as well as ribs, vertebrae, limb bones, and armor. In 1947 the American fossil collectors Charles M. Sternberg and T. Potter Chamney collected a skull and mandible (specimen CMN 8880, formerly NMC 8880), 1 kilometer (5⁄8 mile) north of where the 1910 specimen was found. This is the largest-known Ankylosaurus skull, but it is damaged in places. A section of caudal vertebrae (specimen CCM V03) was discovered in the 1960s in the Powder River drainage, Montana, part of the Hell Creek Formation. In addition to these five incomplete specimens, many other isolated osteoderms and teeth have been found.[12][10]
In 1990, American paleontologist Walter P. Coombs pointed out that the teeth of two skulls assigned to A. magniventris differed from those of the holotype specimen in some details, and though he expressed a "considerate temptation" to name a new species of Ankylosaurus for these, he refrained from doing so, as the range of variation in the species was not completely documented. He also raised the possibility that the two teeth associated with the holotype specimen perhaps did not belong to it, as they were found in matrix within the nasal chambers.[13] The American paleontologist Kenneth Carpenter accepted the teeth as belonging to A. magniventris in 2004, and that all the specimens belonged to the same species, noting that the teeth of other ankylosaurs are highly variable.[10]
Most of the known Ankylosaurus specimens were not scientifically described at length, though several paleontologists planned to do so until Carpenter redescribed the genus in 2004.[10] In 2017 the Canadian paleontologists Victoria M. Arbour and Jordan Mallon redescribed the genus in light of newer ankylosaur discoveries, including elements of the holotype that had not been previously mentioned in the literature (such as parts of the skull and the cervical half-rings). They concluded that though Ankylosaurus is the best-known member of its group, it was bizarre in comparison to related ankylosaurs, and therefore not representative of the group. In spite of its familiarity, it is known from far fewer remains than its closest relatives.[12]
Ankylosaurus was the largest-known ankylosaurine dinosaur and possibly the largest ankylosaurid.[12] In 2004 Carpenter estimated that the individual with the largest-known skull (specimen CMN 8880), which is 64.5 centimeters (2 ft 1+1⁄2 in) long and 74.5 cm (2 ft 5+1⁄4 in) wide, was about 6.25 m (20 ft 6 in) long and had a hip height of about 1.7 m (5 ft 7 in). The smallest-known skull (specimen AMNH 5214) is 55.5 cm (1 ft 9+3⁄4 in) long and 64.5 cm (2 ft 1+1⁄2 in) wide, and Carpenter estimated that it measured about 5.4 m (17 ft 9 in) long and about 1.4 m (4 ft 7 in) tall at the hips.[10] The English paleontologist Roger B. J. Benson and colleagues estimated the weight for AMNH 5214 at 4.78 metric tons (5.27 short tons) in 2014.[14]
In 2017, based on comparisons with more complete ankylosaurines, Arbour and Mallon estimated a length of 7.56 to 9.99 m (24 ft 9+1⁄2 in to 32 ft 9+1⁄2 in) for CMN 8880, and 6.02 to 7.95 m (19 ft 9 in to 26 ft 1 in) for AMNH 5214. Though the latter is the smallest specimen of Ankylosaurus, its skull is still larger than those of any other ankylosaurins. A few other ankylosaurs reached about 6 m (20 ft) in length. Because the vertebrae of AMNH 5214 are not significantly larger than those of other ankylosaurines, Arbour and Mallon considered their upper range estimate of nearly 10 meters (33 ft) for large Ankylosaurus too long, and suggested a length of 8 m (26 ft) instead. Arbour and Mallon estimated a weight of 4.78 t (5.27 short tons) for AMNH 5214, and tentatively estimated the weight of CMN 8880 at 7.95 t (8.76 short tons).[12]
Skull
The three known Ankylosaurus skulls differ in various details; this is thought to be the result of taphonomy (changes happening during decay and fossilization of the remains) and individual variation. The skull was low and triangular in shape, and wider than it was long; the back of the skull was broad and low. The skull had a broad beak on the premaxillae. The orbits (eye sockets) were almost round to slightly oval and did not face directly sideways because the skull tapered towards the front. The braincase was short and robust, as in other ankylosaurines. Crests above the orbits merged into the upper squamosal horns (their shape has been described as "pyramidal"), which pointed backwards to the sides from the back of the skull. The crest and horn were probably separate elements originally, as seen in the related Pinacosaurus and Euoplocephalus. Below the upper horns, jugal horns were present, which pointed backward and down. The horns may have originally been osteoderms that fused to the skull. The scale-like cranial ornamentation on the surfaces of ankylosaurs skulls is called "caputegulae", and were the result of remodeling of the skull itself. This obliterated the sutures between skull elements, which is common for adult ankylosaurs. The caputegulum pattern of the skull was variable between specimens, though some details are shared. The caputegulae are named according to their position on the skull, and those of Ankylosaurus include a relatively large, hexagonal (or diamond-shaped) nasal caputegulum at the front of the snout between the nostrils, which had a loreal caputegulum on each side, an anterior and posterior supraorbital caputegulum above each orbit, and a ridge of nuchal caputegulae at the back of the skull.[10][12][15]
The snout region of Ankylosaurus was unique among ankylosaurs, and had undergone an "extreme" transformation compared to its relatives. The snout was arched and truncated at the front, and the nostrils were elliptical and were directed downward and outward, unlike in all other known ankylosaurids where they faced obliquely forward or upward. Additionally, the nostrils were not visible from the front because the sinuses were expanded to the sides of the premaxilla bones, to a larger extent than seen in other ankylosaurs. Large loreal caputegulae—strap-like, side osteoderms of the snout—completely roofed the enlarged opening of the nostrils, giving a bulbous appearance. The nostrils also had an intranarial septum, which separated the nasal passage from the sinus. Each side of the snout had five sinuses, four of which expanded into the maxilla bone. The nasal cavities (or chambers) of Ankylosaurus were elongated and separated by a septum at the midline, which divided the inside of the snout into two mirrored halves. The nasal chambers had two openings, including the choanae (internal nostrils), and the air passage was looped.[10][12] The maxillae expanded to the sides, giving the impression of a bulge, which may have been due to the sinuses inside. The maxillae had a ridge that may have been the attachment site for fleshy cheeks; the presence of cheeks in ornithischians is controversial, but some nodosaurs had armor plates that covered the cheek region, which may have been embedded in the flesh.[10]
Specimen AMNH 5214 has 34–35 dental alveoli (tooth sockets) in the maxilla. The tooth rows in the maxillae of this specimen are about 20 centimeters (7.9 in) long. Each alveolus had a foramen (opening) near its side where a replacement tooth could be seen. Compared to other ankylosaurs, the mandible of Ankylosaurus was low in proportion to its length, and, when seen from the side, the tooth row was almost straight instead of arched. The mandibles are completely preserved only in the smallest specimen (AMNH 5214) and are about 41 centimeters (16 in) long. The incomplete mandible of the largest specimen (CMN 8880) is the same length. AMNH 5214 has 35 dental alveoli in the left dentary bone () and 36 in the right, for a total of 71. The predentary bone of the tip of the mandibles has not yet been found.[10] Like other ankylosaurs, Ankylosaurus had small, phylliform (leaf-shaped) teeth, which were compressed sideways.[13] The teeth were mostly taller than they were wide, and were very small; their size in proportion to the skull meant that the jaws of Ankylosaurus could accommodate more teeth than other ankylosaurines. The teeth of the largest Ankylosaurus skull are smaller than those of the smallest skull in the absolute sense. Some teeth from behind in the tooth row curved backwards, and tooth crowns were usually flatter on one side than the other.[10] Ankylosaurus teeth are diagnostic and can be distinguished from the teeth of other ankylosaurids based on their smooth sides. The denticles were large, their number ranging from six to eight on the front part of the tooth, and five to seven behind.[10][16]
Postcranial skeleton
The structure of much of the skeleton of Ankylosaurus, including most of the pelvis, tail, and feet, is still unknown.[10] It was quadrupedal, and its hind limbs were longer than its forelimbs.[17] In the holotype specimen, the scapula (shoulder blade) measures 61.5 cm (2 ft 1⁄4 in) long and was fused with the coracoid (a rectangular bone connected to the lower end of the scapula). It also had entheses (connective tissue) for various muscle attachments. The humerus (upper arm bone) of AMNH 5214 was short, very broad and about 54 cm (1 ft 9+1⁄2 in) long. The femur (thigh bone), also from AMNH 5214, was 67 cm (2 ft 2+1⁄2 in) long and very robust. While the feet of Ankylosaurus are incompletely known, the hindfeet probably had three toes, as is the case in advanced ankylosaurids.[10]
The cervical vertebrae had broad neural spines that increased in height towards the body. The front part of the neural spines had well-developed entheses, which was common among adult dinosaurs, and indicates the presence of large ligaments, which helped support the massive head. The dorsal vertebrae had centra (or bodies) that were short relative to their width, and their neural spines were short and narrow. The dorsal vertebrae were tightly spaced, which limited the downwards movement of the back. The neural spines had ossified (turned to bone) tendons, which also overlapped some of the vertebrae. The ribs of the last four back vertebrae were fused to the diapophyses and parapophyses (the structures that articulated the ribs with the vertebrae), and the ribcage was very broad in this part of the body. The caudal vertebrae had centra that were slightly amphicoelous, meaning they were concave on both sides.[10]
Armor
A prominent feature of Ankylosaurus was its armor, consisting of knobs and plates of bone known as osteoderms, or scutes, embedded in the skin. These have not been found in articulation, so their exact placement on the body is unknown, though inferences can be made based on related animals, and various configurations have been proposed. The osteoderms ranged from 1 centimeter (1⁄2 in) in diameter to 35.5 cm (14 in) in length, and varied in shape. The osteoderms of Ankylosaurus were generally thin walled and hollowed on the underside. Compared to Euoplocephalus, the osteoderms of Ankylosaurus were smoother. Many smaller osteoderms and ossicles probably occupied the space between the larger ones, as in other ankylosaurids. The osteoderms covering the body were very flat, though with a low keel at one margin. In contrast, the nodosaurid Edmontonia had high keels stretching from one margin to the other on the midline of its osteoderms. Ankylosaurus had some smaller osteoderms with a keel across the midline.[12][10]
Like other ankylosaurids, Ankylosaurus had cervical half-rings (armor plates on the neck), but these are known only from fragments, making their exact arrangement uncertain. Carpenter suggested that when seen from above, the plates would have been paired, creating an inverted V-shape across the neck, with the midline gap probably being filled with small ossicles (round bony scutes) to allow for movement. He believed the width of this armor belt was too wide to have fitted solely on the neck, and that it covered the base of the neck and continued onto the shoulder region. Arbour and the Canadian paleontologist Philip J. Currie disagreed with Carpenter's interpretation in 2015 and pointed out that the cervical half-ring fragments of the holotype specimen did not fit together in the way proposed by Carpenter (though this could be due to breakage). They instead suggested that the fragments represented the remains of two cervical half-rings, which formed two semi-circular plates of armor around the upper part of the neck, as in the closely related Anodontosaurus and Euoplocephalus.[10][15] Arbour and Mallon elaborated on this idea, describing the shape of these half-rings as "continuous U-shaped yokes" over the upper part of the neck, and suggested that Ankylosaurus had six keeled osteoderms with oval bases on each half-ring.[12]
The first osteoderms behind the second cervical half-ring would have been similar in shape to those in the first half-ring, and the osteoderms on the back probably decreased in diameter hindwards. The largest osteoderms were probably arranged in transverse and longitudinal rows across most of the body, with four or five transverse rows separated by creases in the skin. The osteoderms on the flanks would probably have had a more square outline than those on the back. There may have been four longitudinal rows of osteoderms on the flanks. Unlike some basal ankylosaurs and many nodosaurs, ankylosaurids do not appear to have had co-ossified pelvic shields above their hips. Some osteoderms without keels may have been placed above the hip region of Ankylosaurus, as in Euoplocephalus. Ankylosaurus may have had three or four transverse rows of circular osteoderms over the pelvic region, which were smaller than those on the rest of the body, as in Scolosaurus. Smaller, triangular osteoderms may have been present on the sides of the pelvis. Flattened, pointed plates resemble those on the sides of the tail of Saichania, and may have been distributed similarly on Ankylosaurus. Osteoderms with oval keels could have been placed on the upper side of the tail or the side of the limbs. Compressed, triangular osteoderms found with Ankylosaurus specimens may have been placed on the sides of the pelvis or the tail. Ovoid, keeled, and teardrop-shaped osteoderms are known from Ankylosaurus, and may have been placed on the forelimbs, like those known from Pinacosaurus, but it is unknown whether the hindlimbs bore osteoderms.[10][12]
The tail club (or tail knob) of Ankylosaurus was composed of two large osteoderms, with a row of small osteoderms at the midline, and two small osteoderms at the tip; these osteoderms obscured the last tail vertebra. As only the tail club of specimen AMNH 5214 is known, the range of variation between individuals is unknown. The tail club of AMNH 5214 is 60 cm (23+1⁄2 in) long, 49 cm (19+1⁄2 in) wide, and 19 cm (7+1⁄2 in) tall. The club of the largest specimen may have been 57 cm (22+1⁄2 in) wide. The tail club of Ankylosaurus was semicircular when seen from above, similar to those of Euoplocephalus and Scolosaurus but unlike the pointed club osteoderms of Anodontosaurus or the narrow, elongated club of Dyoplosaurus. The last seven tail vertebrae formed the "handle" of the tail club. These vertebrae were in contact, with no cartilage between them, and were sometimes co-ossified, which made them immobile. Ossified tendons attached to the vertebrae in front of the tail club, and these features together helped strengthen it. The interlocked zygapophyses (articular processes) and neural spines of the handle vertebrae were U-shaped when seen from above, whereas those of most other ankylosaurids are V-shaped, which may be due to the handle of Ankylosaurus being wider. The larger width may indicate that the tail of Ankylosaurus was shorter in relation to its body length than those of other ankylosaurids, or that it had the same proportions but with a smaller club.[12][10][18]
Feeding
Like other ornithischians, Ankylosaurus was herbivorous. Its wide muzzle was adapted for non-selective low-browse cropping,[10] although not to the extent seen in some related genera, especially Euoplocephalus.[12][22] Though ankylosaurs may not have fed on fibrous and woody plants, they may have had a varied diet, including tough leaves and pulpy fruits.[23] Ankylosaurus probably fed on abundant ferns and low-growing shrubs. Assuming it was endothermic, Ankylosaurus would have eaten 60 kilograms (130 pounds) of ferns per day, similar to the amount of dry vegetation a large elephant would consume. The requirements for nutrition could have been more effectively met if Ankylosaurus ate fruit, which its small, cusp-like teeth and the shape of its beak seem well adapted for, compared to for example Euoplocephalus. Certain invertebrates, which the small teeth may have been adapted for handling, could also have provided supplemental nutrition.[12]
Fossils of Ankylosaurus teeth exhibit wear on the face of the crown rather than on the tip of the crown, as in nodosaurid ankylosaurs.[10] In 1982 Carpenter ascribed to baby Ankylosaurus two very small teeth that originate from the Lance and Hell Creek Formations and measure 3.2 to 3.3 mm (1⁄8 to 17⁄128 in) in length, respectively. The smaller tooth is heavily worn, leading Carpenter to suggest that ankylosaurids in general or at least the young did not swallow their food whole but employed some sort of chewing.[16] Since adult Ankylosaurus did little chewing of its food, it would have spent less time in the day foraging than an elephant.[12] Based on the broadness of the ribcage, the digestion of unchewed food may have been facilitated by hindgut fermentation like in modern herbivorous lizards, which have several chambers in their enlarged colon.[10]
In 1969, paleontologist Georg Haas concluded that despite the large size of ankylosaur skulls, the associated musculature was relatively weak. He also thought jaw movement was limited to up and down movements. Extrapolating from this, Haas suggested that ankylosaurs ate relatively soft non-abrasive vegetation.[24] Later research on Euoplocephalus indicates that forward and sideways jaw movement was possible in these animals, the skull being able to withstand considerable forces.[25] A 2016 study of the dental occlusion (contact between the teeth) of ankylosaur specimens found that the ability for backwards (palinal) jaw movement evolved independently in different ankylosaur lineages, including Late Cretaceous North American ankylosaurids like Ankylosaurus and Euoplocephalus.[22]
The retracted position of the nostrils of Ankylosaurus were compared to those of fossorial (digging) worm lizards and blind snakes by Arbour and Mallon in 2017, and though it was probably not a burrowing animal, the snout of Ankylosaurus may indicate earth-moving behavior. These factors, as well as the low rate of tooth formation in ankylosaurs compared to other ornithischians, indicate that Ankylosaurus may have been omnivorous (eating both plant and animal matter). It may also (or alternatively) have dug in the ground for roots and tubers.[12] A 2023 study by paleontologist Antonio Ballell and colleagues found that North American ankylosaurids from the latest Cretaceous (including Ankylosaurus) had jaws with low mechanical advantage, whereas those of earlier relatives were high to moderate. These late ankylosaurids also had tooth occlusion and complex biphasal jaw mechanisms, features shared with some Late Cretaceous nodosaurids, but those instead have jaws with high mechanical advantage. This indicates that while the two groups converged in some features, the nodosaurs had higher relative bite force, which suggests diverging jaw mechanics and dietary partitioning between the two.[26]
Airspaces and senses
In 1977, paleontologist Teresa Maryańska proposed that the complex sinuses and nasal cavities of ankylosaurs may have lightened the weight of the skull, housed a nasal gland, or acted as a chamber for vocal resonance.[10][27] Carpenter rejected these hypotheses, arguing that tetrapod animals make sounds through the larynx, not the nostrils, and that reduction in weight was minimal, as the spaces only accounted for a small percent of the skull volume. He also considered a gland unlikely and noted that the sinuses may not have had any specific function.[10] It has also been suggested that the respiratory passages were used to perform a mammal-like treatment of inhaled air, based on the presence and arrangement of specialized bones.[27]
A 2011 study of the nasal passages of Euoplocephalus by paleontologist Tetsuto Miyashita and colleagues supported their function as a heat and water balancing system, noting the extensive blood vessel system and an increased surface area for the mucosa membrane (used for heat and water exchange in modern animals). The researchers also supported the idea of the loops acting as a resonance chamber, comparable to the elongated nasal passages of saiga antelope and the looping trachea of cranes and swans. Reconstructions of the inner ear suggest adaptation to hearing at low frequencies, such as the low-toned resonant sounds possibly produced by the nasal passages. They disputed the possibility that the looping is related to olfaction (sense of smell) as the olfactory region is pushed to the sides of the main airway.[28]
According to Carpenter, the shape of the nasal chambers of Ankylosaurus indicate that airflow was unidirectional (looping through the lungs during inhalation and exhalation), although it may also have been bidirectional in the posterior nasal chamber, with air directed past the olfactory lobes.[10] The enlarged olfactory region of ankylosaurids indicates a well-developed sense of smell.[28] Though hindwards retraction of the nostrils is seen in aquatic animals and animals with a proboscis, it is unlikely either possibility applies to Ankylosaurus, as the nostrils tend to be reduced or the premaxilla extended. In addition, though the widely separated nostrils may have allowed for stereo-olfaction (where each nostril senses smells from different directions), as has been proposed for the moose, little is known about this feature.[12] The position of the orbits of Ankylosaurus suggest some stereoscopic vision.[10]
Limb movements
Reconstructions of ankylosaur forelimb musculature made by Coombs in 1978 suggest that the forelimbs bore the majority of the animal's weight, and were adapted for high force delivery on the front feet, possibly for food gathering. In addition, Coombs suggested that ankylosaurs may have been capable diggers, though the hoof-like structure of the manus would have limited fossorial activity. Ankylosaurs were likely to have been slow-moving and sluggish animals,[29][30] though they may have been capable of quick movements when necessary.[17]
Growth
The squamosal horns of the largest Ankylosaurus specimen are blunter than those of the smallest specimen, which is also the case in Euoplocephalus, and this may represent ontogenetic variation (related to growth development).[12] Studies of specimens of Pinacosaurus of different ages found that during ontogenetic development, the ribs of juvenile ankylosaurs fused with their vertebrae. The forelimbs strongly increased in robustness while the hindlimbs did not become larger relative to the rest of the skeleton, further evidence that the arms bore most of the weight. In the cervical half-rings, the underlying bone band developed outgrowths connecting it with the underlying osteoderms, which simultaneously fused to each other.[31] On the skull, the middle bone plates first ossified at the snout and the rear rim, with ossification gradually extending towards the middle regions. On the rest of the body, ossification progressed from the neck backward in the direction of the tail.[32]
Defense
The osteoderms of ankylosaurids were thin in comparison to those of other ankylosaurs, and appear to have been strengthened by randomly distributed cushions of collagen fibers. Structurally similar to Sharpey's fibres, they were embedded directly into the bone tissue, a feature unique to ankylosaurids. This would have provided the ankylosaurids with an armor covering that was both lightweight and highly durable, being resistant to breakage and penetration by the teeth of predators.[33] The palpebral bones over the eyes may have provided additional protection for them.[34] Carpenter suggested in 1982 that the heavily vascularized armor may also have had a role in thermoregulation as in modern crocodilians.[35]
The tail club of Ankylosaurus seems to have been an active defensive weapon, capable of producing enough of an impact to break the bones of an assailant. The tendons of the tail were partially ossified and were not very elastic, allowing great force to be transmitted to the club when it was used as a weapon.[10] Coombs suggested in 1979 that several hindlimb muscles would have controlled the swinging of the tail, and that violent thrusts of the club would have been able to break the metatarsal bones of large theropods.[30]
A 2009 study estimated that ankylosaurids could swing their tails at 100 degrees laterally, and the mainly cancellous clubs would have had a lowered moment of inertia and been effective weapons. The study also found that while adult ankylosaurid tail clubs were capable of breaking bones, those of juveniles were not. Despite the feasibility of tail-swinging, the researchers could not determine whether ankylosaurids used their clubs for defense against potential predators, in intraspecific combat, or both.[36] Other studies have found evidence of ankylosaurids using their tail clubs for intraspecific combat. One specimen of Tarchia showed signs of injury on both the pelvic and tail area and the club was found to be asymmetrical, a sign of being worn down by the strikes.[37]
In 1993, Tony Thulborn proposed that the tail club of ankylosaurids primarily acted as a decoy for the head, as he thought the tail too short and inflexible to have an effective reach; the "dummy head" would lure a predator close to the tail, where it could be struck.[38] Carpenter has rejected this idea, as tail club shape is highly variable among ankylosaurids, even in the same genus.[10]
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https://www.wikiwand.com/en/Saichania
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Saichania is a genus of herbivorous ankylosaurid dinosaur from the Late Cretaceous period of Mongolia and China.
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https://www.wikiwand.com/en/articles/Saichania
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Size, build and distinguishing traits
Saichania was a medium-sized ankylosaur, measuring 5–7 metres (16–23 ft) in length and 1.4–2.0 metric tons (1.5–2.2 short tons) in body mass.[5][6][1] Finds of tail clubs of gigantic individuals suggest larger sizes but their reference to Saichania cannot be substantiated as the holotype, the only specimen sufficiently described, only consists of the front of the animal.[3]
Saichania shared the general ankylosaurid build, being a low-slung, broad, heavily armoured dinosaur, with short forelimbs. Even for an ankylosaurid however, Saichania is exceptionally robust, its rump strengthened by ossifications and fusions of the vertebral column, ribs, shoulder girdle and breast bones.[1]
Arbour in 2014 established a revised list of distinguishing traits. The osteoderms on the skull are bulbous. The first and second neck vertebrae are fused into a single element, a syncervical. The upper side of the humerus is very broad, equalling 70% of the total length of the bone. The rib shafts are expanded by intercostal ossifications, the cartilage connecting the ribs having been turned into bone sheets. The cervical halfrings, protecting the neck, have each an underlying continuous band of bone and the borders between the segments of these rings are covered by extra armour plates entirely hiding these connections from view.[3]
The skull of Saichania is broad, 455 millimetres long and 480 millimetres wide with the holotype.[1] The top of the snout is covered with strongly convex osteoderms. These armour tiles on the snout comprise a central large caputegula. A large "loreal" osteoderm covers much of the top edge and the side of the snout. The caputegula on the prefrontal is of moderate size and not strongly protruding sideways. The osteoderms on the upper eye socket rim are continuous, not forming two peaks. An extra osteoderm on the rear supraorbital, as in Tarchia, is lacking. The pyramid-shaped squamosal horns on the rear skull corners are broad, not narrow as with Tarchia. These horns have a uniform surface texture, not a division into a smooth and rough surface as in Zaraapelta.[3] On the cheek, large triangular quadratojugal horns are present.
Skeleton
The skull had very complex air passages. The main entrance of each external nostril consisted of a roomy "nasal vestibule". In each vestibule again two smaller entrances were present, vertically arranged. The lower hole allowed air to enter the hollow inside of the bone core of the beak. This premaxillary sinus had a little recess at the top, connected by a nerve channel to the mouth. Maryańska presumed this recess housed a Jacobson's organ, a secondary smelling organ. The main room of the premaxillary sinus was connected to behind with a sinus in the maxilla, which itself was partly divided in two by a transverse bone wall or septum. The nasal cavity was large, situated directly below the snout roof. It was divided into a left and right side by a thick vertical bone wall. It was also horizontally divided in two by high internal wings of the praemaxillae and the upper side of a crista maxilloturbinalis. This latter was a scroll-like structure, a turbinate bone serving with warm-blooded animals to condense and preserve exhaled moisture. Normally, in dinosaurs these turbinates are not ossified. Together with a crista nasoturbinalis, the crista maxilloturbinalis filled the lower half of the nasal cavity. Maryańska presumed it was connected with the underlying premaxillary sinus, allowing the animal to exhale air through the lower hole of the nasal vestibule. The upper half of the nasal cavity was the main respiratory tract, allowing air to enter via the upper hole of the nasal vestibule.[1] An unusually strongly ossified hard palate was present. The air passages may have allowed the animal to cool the air that it breathed and limit water loss. The hard palate allowed it to eat tough plants. All this suggested that it lived in a hot, arid, environment. There is even some evidence that the animal may have possessed a salt gland next to its nostrils, which would have further aided it in a desert habitat.[7]
The teeth were small and leaf-shaped. There are twenty-two of them in each maxilla, seventeen in the right and sixteen in the left lower jaw of the holotype. On the rear skull, the oval occipital condyle is obliquely pointing to below, indicating that the entire head was appending. A large hyoid bone apparatus was found, in 1977 the most complete discovered for any dinosaur. It is V-shaped with the central parts representing the basihyal and basibranchial, and the branches being the ceratobranchialia.[1] This bone apparently supported a long tongue.
The front skeleton shows some exceptional ossifications and fusions. The front neck vertebrae, the atlas and axis, are grown together. The cervical vertebrae have very long joint processes, zygapophyses, showing that thick intervertebral discs must have been present and that the neck was longer and more flexible than is often assumed. The short rib and the diapophysis of the first dorsal vertebra are fused with the coracoid, immobilising the entire shoulder girdle relative to the vertebral column. The coracoids are small but sharply curving to the inside below, almost meeting each other. From the fifth rib onwards, the rib shafts have intercostal plates on their rear edges, ossified cartilage sheets, overlapping the front edge of the next rib. The intercostal plate is positioned in a relatively high position in the fifth rib; more to the rear of the series it gradually descends towards the lower belly. These ribs also articulate at their lower ends with the breast bones, a condition which is rare in the Ornithischia. The breast bones are fully ossified and connect to form a sternal plate that is split in front and broadly forked at the rear.[1]
The humerus is very robust. Thirty centimetres long in the holotype, it has an upper side width of 212 millimetres due to a well-developed inner corner and a strong hatchet-shaped deltopectoral crest. The ulna, twenty-one centimetres in length, also is robust but has a relatively low olecranon. The metacarpus is short, in 1977 it was the shortest of any Asian ankylosaur known. The metacarpals were positioned vertically, closely connected into an arch. Below the first and second metacarpal small disc-shaped sesamoid bones were found.[1]
Body armour
The holotype preserves the front body armour in articulation. The neck is protected by two cervical halfrings, each made of six rectangular segments positioned next to each other: two at the top, two at the upper sides and two at the lower sides. Each segment has a keel parallel to the long axis of the body. The keel of the lower side segments is the largest. The segments are connected to an underlying continuous band of bone, mainly by a broad fusion at the front edge, but also by a narrow strip at the rear. The seams between the segments are covered by a rectangular zone of small oval osteoderms. Between the upper and lower side segment a larger central osteoderm is present, forming a rosette. The front halfring is smaller than the rear one.[1]
A central row of symmetrical conical osteoderms is positioned on the back. On both sides of this median series, a parallel row of large thin osteoderms is present, featuring moderately high keels, their apexes pointing to behind. The vertical sides of the rump are covered by three rows of conical osteoderms: the upper rim is equipped with large plates and apexes pointing to the rear; at the middle side a similar row is present of even larger plates; the lower edge has a row of smaller plates, their keels to the contrary directed to the front. In general, the keels are sharp and narrower plates have higher and more asymmetrically placed keels. Some osteoderms have the shape of pure cones. The larger osteoderms are also ordered in transverse rows but are not fused into bands; small ossicles connect the larger elements. On the underside of the breast, osteoderms are present also.[1]
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Kikimalou's "Scale" collection - 1/25 shelves - updated March 1 2024
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[] |
[] |
[
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] | null |
[
"Kikimalou"
] |
2024-03-02T00:41:06+11:00
|
Last year I showed you the 1/15, 1/18 and 1/4 floors of my collection. I wasn't able to show you another shelves before now, too much other things to do first
|
en
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https://illiweb.com/fa/favicon/discussion.ico
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sts-forum.forumieren.de
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https://sts-forum.forumieren.de/t25354-kikimalou-s-scale-collection-1-25-shelves-updated-march-1-2024
|
Last year I showed you the [You must be registered and logged in to see this link.], [You must be registered and logged in to see this link.] and [You must be registered and logged in to see this link.] floors of my collection. I wasn't able to show you another shelves before now, too much other things to do first and this kind of topic needs more work than a "what's new" or a walkaround.
The 1/25 (or 1/24) scale is a classic in the toy world, and was widely used officially in Germany in the second half of the 20th century.
With 99 inhabitants, it's the 5th most populated floor in my mainstream collection.
This is a pivotal scale for me, from here on and beyond, models of extinct species will occupy more space, both in number and size. For the 1/25, they only represent a quarter of the total, but that's not bad for a collector as unspecialised as me.
Let's begin our journey as usual, with Africa
South Africa: CollectA white rhino, Takara Tomy Lion and Meerkat
[You must be registered and logged in to see this image.]
Kenya savannah: Mojo White rhino calf, Eikoh White rhino, CollectA Giraffe, Colorata African wild dog and Yujin Warthog
[You must be registered and logged in to see this image.]
Liberia: Preiser Buffalo,used as a Sudan buffalo, Play Visions Spotted hyena, Starlux Goliath heron and Starlux Jentink's duiker.
[You must be registered and logged in to see this image.]
Ouganda marsh: Eikoh Shoebill and Warthog, Nayab Crocodile and Play Visions Sitatunga.
[You must be registered and logged in to see this image.]
Because I didn’t want to let the poor lemur alone, my African and Malagasy Primates tribe: Kaiyodo Black-and-white ruffed lemur and Lowland gorilla, Bandai Chimps.
[You must be registered and logged in to see this image.]
Europe: Starlux Otter, Hausser elastolin Alpine ibex, Otter, Pine matter and Brown bears, Schleich Moose.
[You must be registered and logged in to see this image.]
South-east Asia: Marx Elephant, Schleich Malayan tapir, Milelire and Safari Ltd Sumatran rhinos, Divartsity Greater mouse deer, Kaiyodo Komodo dragon and Milelire Siamang.
[You must be registered and logged in to see this image.]
China: Kaiyodo and Colorata Giant pandas, Bandai Red pandas
[You must be registered and logged in to see this image.]
India: Eikoh Black panther, Colorata Asiatic lion,Safari ltd Indian rhino, yujin Indian star tortoise, Schleich Indian rhino calf, Colorata Sloth bear
[You must be registered and logged in to see this image.]
Japan: Kaiyodo Ryukyu wild boar, Hokkaido wolf and Yezo sika deer, Ikimon Japanese hares
[You must be registered and logged in to see this image.]
South America: Kaiyodo Capybara, Jaguar and Giant anteater, Colorata Spectacled bear.
[You must be registered and logged in to see this image.]
Hausser Polar bear (who needs a repaint) and Safari Ltd walrus
[You must be registered and logged in to see this image.]
Oceania: Bandai Koalas and Western grey kangaroo pair, Ikimon Red kangaroo, Preiser Dromedary and calf, Takara Tomy Thylacine.
[You must be registered and logged in to see this image.]
Above the Pacific ocean: Colorata Sea otter and Kaiyodo Laysan Albatross.
[You must be registered and logged in to see this image.]
Below the seas: Kaiyodo Emperor penguin, Long snouted lancetfish and Blacktip reef shark, CollectA Pygmy sperm whale, Short-finned pilot whale and Blainville's beaked whale, Colorata Humphead wrasse and Yujin Loggerhead sea turtle.
[You must be registered and logged in to see this image.]
Some comparison pics maybe ?
Camels
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Deers
[You must be registered and logged in to see this image.]
Crocs
[You must be registered and logged in to see this image.]
Some "giants"
[You must be registered and logged in to see this image.]
Horned gang
[You must be registered and logged in to see this image.]
Horns and Tuscks
[You must be registered and logged in to see this image.]
Bears
[You must be registered and logged in to see this image.]
Hunters
[You must be registered and logged in to see this image.]
Rhinos
[You must be registered and logged in to see this image.]
Thyreophorans
[You must be registered and logged in to see this image.]
You know, I'm really into these topics!
It's incredible how small the figures representing animals smaller than wolves are.
For instance, a fox at this scale appears tiny, yet you can still depict martens, meerkats, and so on.
It's interesting that there isn't a single Noah's Pals which is known for working at this scale.
Also no Papo among extant animals but, these figures other major brands make at 1:24 scale, Papo tends to make them larger, namely the moose. Birds always look too big in your picture groups, I don't know if it is the way they are measured in real that promote a wrong perception or more likely it is my perception of their sizes which is adulterated.
_________________
~ Rogério [You must be registered and logged in to see this image.] [You must be registered and logged in to see this link.]
Thanks to vintage or Japanese manufacturers, you can find tiny animals for small scales, there are even smaller-scale foxes.
Make no mistake, some Papos are too big for 1/25, but others, like elephants, are too small.
As for birds, the Goliath Heron is the world's largest heron, reaching 1.50 m in height, as is the shoebill, while the Emperor Penguin can reach 1.20 m. The Laysan albatross has a wingspan of 2m. The Copepteryx titan has an estimated length of two meters.
I had quite a few Noah's Pals when I first started out, but I let them all go because I didn't like the style that much. Today, all I have still are platypus, marabou storks and skunks. I'm waiting for a pair of jackals...
As for the famous 1/25, I don't trust Noah's Pals any more than any other manufacturer. The Maiasaura that Kaiyodo concocted for us in the 90s is reputed to be 1/35, whereas it's actually 1/25 for a 9m animal.
And we all have preconceived ideas about the size of animals. For example, we imagine many dinosaurs to be extravagantly large but, for example, Nasutoceratops are actually smaller than white rhinoceros.
It's the same for me Jolie, I sometimes forget how big or small are these animals actually are. Thats' why I like very much that exercise.
Alain, Miragaia are not longer than 6m long with a lot of neck and tail in these 6m. It's not a big dino, just like the Nasutoceratops, Zhejiangosaurus and Pinacosaurus. For a long time, we were led to believe that all dinosaurs were gigantic beasts, with a few exceptions. Even though we now know that there were many small dinosaurs, we can't help imagining the Tanks and other ceratopsians to be bigger than they actually are.
Sometimes it's even the manufacturers who mislead us. For example, Kaiyodo claims that its Maiasaura is at 1/35 scale, when in fact it's at 1/25 scale for a beautiful 9m animal. At 1/35 it would measure over 12m, which was not the case.
Inostrancevia was the biggest gorgonopsian known with a total lenght between 3m and 3.5m, it dwarves even a 2.5m long Polar bear
Blaine, even though you're not sensitive to scale, you go to great efforts to inform us in your Museum
Yes, regarding the Papo figures, I was exclusively referring to models that other brands usually place near this scale, and Papo's counterparts are usually larger. But it's not a rule, as nothing is a rule when it comes to scales and major brands.
About birds, I tend to perceive them as always smaller. I remember my shock when I saw a simple white stork near me, not to mention an ostrich that escaped from the circus and stopped quite close to me. Also, during demonstrations with birds of prey, I'm always surprised to see that the wingspan of the birds clearly exceeds the size of the humans handling them. All of this makes presentations with figures at the same scale even more interesting.
_________________
~ Rogério [You must be registered and logged in to see this image.] [You must be registered and logged in to see this link.]
Kikimalou wrote:
Hunters
[You must be registered and logged in to see this image.]
Rhinos
[You must be registered and logged in to see this image.]
so many wonderful photos to study Christophe! I bet it took you a long time to lay them all out and get the angles just right for photographing?? But I bet it was fun though?
I really love your scale topic as well. I always learn so much and I am amazed at all the figures that are 1/25. I have quite a few of them as well so was surprised and delighted to see them here at this scale.
Ofcourse it's always an interesting journey to work out what will and won't fit into my scale collection and sometimes I have models that are bigger, or smaller than normal.
Still, I was surprised - is the hyena actually bigger than a wolf? I always thought wolves were massive animals! haha, maybe I've seen too many wilderness movies
And I just LOVE the photo with the 2 baby rhinos side by side! These models are really beautiful.
I think they will fit into my rhino creche as well.
Yes Annette, it takes along time to produce pics for such a topic, that's why I can't do one every week. Work, the necessities of life and family life obviously come before photo shoots.
I can only do it when I have enough free time. This week I was off work, I had planned to take care of the garden, but it rained like a cow is pissing almost every day... What luck, I was able to devote myself to this topic !
Getting the angles just right for photographing is a big part of the sport. To be able to compare the size of animals as big as or bigger than us most effectively, the camera should ideally be placed at human eye level. At 1/25 this means that the center of the lens is between 6.5 and 7cm from the ground. For smaller animals, try to position the camera at their average eye level.
The problem is that when there are a lot of animals to compare, or when they're roughly the same size, the advice I've just given no longer works, because you can't distinguish one animal from another. In this case, you need to raise the camera above the mix.
Choosing the right angle is always a compromise between these two rules.
This means that for each photo, I must try to arrange the animals as harmoniously as possible, and also try to respect rule n°1 as best I can. Sometimes I have to make several attempts. It also means that for each photo I have to adjust the height of my photographic tripod.
That's the most physical part of the job, after which all that's left for me to do is spend my time doing lab work on my computer, drinking too much coffee or tea. But I suppose, Annette, you know all that too.
About wolves and hyenas, Hokkaido wolves stood 70 to 80 cm at the withers, Spotted hyena stood 70 to 91.5 cm. What's more, the PV Hyena stands erect on its paws, while the Kaiyodo wolf bends down slightly as if on the hunt.
|
||||
622
|
dbpedia
|
2
| 29
|
https://www.frontiersin.org/articles/10.3389/feart.2022.803505/full
|
en
|
New Ankylosaurian Cranial Remains From the Lower Cretaceous (Upper Albian) Toolebuc Formation of Queensland, Australia
|
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[
"Ankylosauria",
"Kunbarrasaurus",
"Australia",
"Minmi",
"Cretaceous",
"Gondwana",
"Toolebuc formation",
"Synchrotron"
] | null |
[
"Timothy G",
"Phil R",
"Joseph J",
"Russell D. C",
"Benjamin P",
"Nicolás E"
] | null |
Australian dinosaur research has undergone a renaissance in the last 10 years, with growing knowledge of mid-Cretaceous assemblages revealing an endemic high...
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en
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Frontiers
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https://www.frontiersin.org/journals/earth-science/articles/10.3389/feart.2022.803505/full
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Introduction
Ankylosaurians are a rare component of Gondwanan Cretaceous terrestrial ecosystems with five described species: Antarctopelta oliveroi, Minmi paravertebra, Kunbarrasaurus ieversi, Spicomellus afer, and Stegouros elengassen (Molnar, 1980; Salgado and Gasparini, 2006; Leahey et al., 2015; Maidment et al., 2021; Soto-Acuña et al., 2021). A. oliveroi and M. paravertebra have been considered nomen dubia due to a perceived lack of autapomorphies (Leahey et al., 2015; Arbour and Currie, 2016). However, comparisons between M. paravertebra and K. ieversi support their differentiation and the specific validity (Leahey et al., 2015). Despite the currently limited taxonomic diversity of Gondwanan Cretaceous ankylosaurs, skeletal fragments ascribed to the clade are found throughout Argentina (Coria and Salgado, 2001; Murray et al., 2019), New Zealand (Molnar and Wiffen, 1994), and Madagascar (Russel et al., 1976), with further ichnological evidence from Bolivia (Apesteguía and Gallina, 2011; Riguetti et al., 2021), and Brazil (Francischini et al., 2018). These widespread occurrences demonstrate that ankylosaurs were a rare yet pervasive component of Gondwanan dinosaur faunas. In Australia, ankylosaurian occurrences include two named taxa, M. paravertebra (Molnar, 1980) and K. ieversi (Molnar, 1996; Leahey et al., 2015), along with footprints (Salisbury et al., 2016) and several isolated skeletal elements that span most Australian dinosaur-bearing formations (Molnar, 1996; Molnar, 2001; Barrett et al., 2010; Leahey and Salisbury, 2013; Leahey et al., 2015; Bell et al., 2018a).
Despite brief mentions in the literature for almost 40 years, ankylosaurs from the middle–upper Albian marine Toolebuc Formation of central Queensland have never previously been studied in detail (Molnar, 1996; Leahey and Salisbury, 2013; Leahey et al., 2015). Three specimens were previously reported from two localities, Julia Creek (QM F33286) and Hughenden (AM F35259, AM F119849), and all were tentatively referred to the genus Minmi (Figures 1A,B; Molnar, 1996). QM F33286 is a partially disarticulated ankylosaur comprising “thoracic and pelvic” elements associated with ventral or lateroventral dermal ossifications (Molnar, 2001; Leahey and Salisbury, 2013). A brief report mentioned that the dermal ossicles were tightly arranged as square tiles that presumably covered most of the neck and trunk (Molnar, 2001). AM F35259 includes only a series of incomplete ribs with ossicles. AM F119849 likewise comprises a single block with vertebrae, ribs, and dermal ossifications (Leahey and Salisbury, 2013).
FIGURE 1
Here we provide the first comprehensive description of ankylosaur cranial remains recovered from the Toolebuc Formation based on a previously undocumented specimen, SAMA P40536, collected by BPK in 2005 from Warra Station near Boulia in western Queensland. SAMA P40536 is preserved within a series of yellow limestone concretions typical of the Toolebuc Formation (Kellner et al., 2010) but had eroded out and were dispersed throughout the blacksoil weathering residuum. The concretions contain remnants of the skull, pelvis, and limbs associated with isolated vertebrae, ribs, and dermal armor fragments. Our initial assessment of the remains focuses on the cranium and dentition, which are important because they represent only the second example of an ankylosaur skull recovered from Australia to date.
Institutional Abbreviations
QM, Queensland Museum, Brisbane, Queensland, Australia; SAMA, South Australian Museum, Adelaide, South Australia, Australia; AM, Australian Museum, Sydney, New South Wales, Australia.
Methods for Synchrotron Scanning and 3D Modeling
Microtomographic measurements of SAMA P40536 were performed using the Imaging and Medical Beamline (IMBL) at the Australian Nuclear Science and Technology Organisation’s (ANSTO) Australian Synchrotron, Melbourne, Victoria, Australia. For this investigation, acquisition parameters included a 40.29 × 40.29 μm pixel size, monochromatic beam energy of 70 keV, a sample–detector distance of 500 mm, and the “Ruby” detector. The latter consists of a PCO. edge sCMOS camera (16-bit, 2,560 × 2,160 pixels) and a Nikon Makro Planar 100 mm lens coupled with a 20 µm thick Gadox/CsI(Tl)/CdWO4 scintillator screen. As the height and width of the specimen exceeded the detector field-of-view, the specimen was aligned axially relative to the beam, its center of rotation shifted toward one edge of the detector. Twelve successive scans were required to cover the full specimen volume, each consisting of 1,500 equally spaced angle shadow radiographs with an exposure length of 0.35 s, obtained every 0.12° as the sample was continuously rotated 180° about its vertical axis. Horizontal translation of the specimen between tomographic scans was 80 mm, and 25 mm between vertical scans. One-hundred dark (closed shutter) and beam profile (open shutter) images were obtained for calibration before and after shadow-radiograph acquisition. The total time for the scan was 140 min.
The raw 16-bit radiographic series were normalized relative to the beam calibration files and stitched with the in-house software “IMBL-Stitch” to yield a 32-bit series with a 178 × 160 mm field-of-view. Reconstruction of the 3D dataset was achieved by the filtered-back projection method using the CSIRO’s X-TRACT (Gureyev et al., 2011). The image stack (Available on MorphoSource; ark:/87602/m4/392687) was segmented, and the volume data was rendered using Mimics Innovation Suite (v. 21.4; Materialise HQ, Leuven, Belgium). As various sections are preserved as bone impressions, with the original bone lost, thin surface models (∼0.8 mm thick) were added during segmentation to illustrate the relative placement of these bones. Finally, to create positive molds of the tooth row and properly study the dental anatomy, the negative left dentary tooth row impression was molded in Pinkysil® (fast set silicone) and cast in Easycast® (Rigid Polyurethane System). Manual preparation techniques using an air scribe (Paleotools® Micro Jack 3) were used to extract a single preserved tooth crown, subsequently scanned using the General Electric (GE) Phoenix V|tome|xs microCT scanner at the University of New England, Armidale. This tooth did not show in the original synchrotron scan.
Systematic Palaeontology
Dinosauria Owen, 1842
Ornithischia Seeley, 1888
Thyreophora Nopcsa, 1915
Ankylosauria Osborn, 1923
Kunbarrasaurus Leahey et al., 2015cf. Kunbarrasaurus sp.
Referred Material—SAMA P40536, a partial skull incorporating impressions of the left and right maxillae, the vomer, both palatines, the right (and part impression of the left) ectopterygoid, a possible right mandibular ramus, impressions of both maxillary tooth rows, and eight isolated teeth (Supplementary Materials S1, S2 available at figshare; https://figshare.com/articles/figure/3D_PDF_of_SAMA_P40536-_Frauenfelder_et_al_/16892908). Associated postcranial elements include parts of the pelvis, limb elements, vertebrae, ribs, scutes, and numerous dermal ossicles. However, much of the postcranial skeleton remains encased in limestone matrix and thus requires preparation before describing it.
Horizon and locality—The limestone concretions containing SAMA P40536 had weathered out of organic-rich marine shales that are lithostratigraphically representative of the Toolebuc Formation. This unit crops out over a vast area west of the township of Boulia in southwestern Queensland (Figure 1). The Toolebuc Formation strata around Boulia have long been recognized as abundant sources of vertebrate fossils with documented finds including chimaeroids, lamniform sharks, a diverse range of actinopterygians, ceratodont dipnoans (e.g., Lees, 1986; Rozefelds, 1992; Kemp, 1993; Bartholomai, 2004; Kear, 2007; Wilson et al., 2011; Bartholomai, 2012; Bartholomai, 2015a; Bartholomai, 2015b), marine reptiles incorporating elasmosaurid, polycotylid, and pliosaurid plesiosaurs, ophthalmosaurid ichthyosaurs and protostegid turtles (e.g., Kear, 2003; Kear, 2006; Kear and Lee, 2006; Zammit et al., 2010; Kear and Hamilton-Bruce, 2011; Kear, 2016; Kear et al., 2018), ornithocheiroid pterosaurs (Molnar and Thulborn, 1980; Molnar, 1987; Fletcher and Salisbury, 2010; Kellner et al., 2010, 2011; Pentland and Poropat, 2019), and enantiornithine birds (Molnar, 1986; Chiappe, 1996; Kurochkin and Molnar, 1997; Kear et al., 2003). These assemblages are associated with abundant pelagic cephalopod (ammonites, belemnites, and coleoids) and benthic invertebrate remains (as summarized in Henderson et al., 2000) that indicate a shallow off-shore marine setting subject to poorly oxygenated bottom water conditions (Henderson, 2004; Jiang et al., 2018). Chronostratigraphically, the Toolebuc Formation is correlated with the latest middle–upper Albian Coptospora paradoxa spore/pollen and Pseudoceratium ludbrookiae dinoflagellate zones (Figure 1C; McMinn and Burger, 1986; Moore et al., 1986).
Description
The skull of SAMA P40536 is incomplete and preserves a portion of the palatal region (Figure 2). The choanal width is 82 mm, similar to proposed values of 72 mm for K. ieversi (measured from Leahey et al., 2015 p. 22), suggesting a potential skull length of ∼270 mm and skull width of ∼280 mm. The holotype of K. ieversi was assumed to be near-mature to mature at the time of death based on its small size and lack of cranial fusion (Molnar, 1996; Leahey et al., 2015). Given the similarity in size, SAMA P40536 may have been of equivalent age.
FIGURE 2
Although not described herein, a large bony fragment is preserved on the ventral and right lateral surfaces, with the lateral extent unknown due to erosion. The bone is mediolaterally compressed anteriorly and forms a ventrally projecting hook-like process. We tentatively identify this fragment as belonging to the dentary due to its position, ventral to the maxillary tooth row. The paired choanae of SAMA P40536 are separated medially by the posterior portion of the vomer (Figure 2). They are crescent-shaped, curving away from the midline posterolaterally, and are slightly wider than they are long. Anteriorly, the vomer, maxillary tooth roots, and maxillary bone impressions form the boundary of both choanae. Typically, the secondary palate separates the choana from the maxillary tooth row (Bourke et al., 2018); however, here, these elements form the choanal boundary due to mediolateral compression of the maxillary tooth row. The palatines form the posterior boundary of the choanae, medially, along with the ectopterygoid, laterally (Figure 2). The contribution of ectopterygoid to the choanal boundary suggests that a “posteroventral” secondary palate, which generally braces the palatine against the medial surface of the maxilla, may not have been present (Vickaryous et al., 2004; but see Bourke et al., 2018 for a new interpretation of this palatal structure). The choanae are posteriorly positioned, with their preserved anterior extent roughly in line with the middle of the maxillary tooth row. This condition is similar to that seen in K. ieversi but differs from that of ankylosaurids and nodosaurids (Lee, 1996; Godefroit et al., 1999; Carpenter, 2004; Kilbourne and Carpenter, 2005; Leahey et al., 2015; Kinneer et al., 2016).
Facial Bones
Maxilla—The right and left ventromedial surfaces of the maxilla are preserved as impressions on the block; a small bony fragment of the left maxilla is also present along the anterior choanal margin (Figures 2B,C). The maxillary impressions form an arcuate contact with the ectopterygoid posteriorly and medially (Figure 3D), together forming the lateral and posterolaterally margins of the choanae, respectively. Anteriorly, the impressions enclose the posterior portion of the exposed maxillary tooth roots. Right, and left maxillary tooth rows are partially preserved as impressions of their medial surfaces (Figure 3C). The left maxillary tooth row is 82 mm in length, whilst the right is 90 mm. The tooth rows are straight for approximately two-thirds of their length, diverging posteriorly. The final third of the tooth row curves laterally, producing an overall curved tooth row in palatal view. An impression of the alveolar margin occurs on the left side of the block; however, it is not preserved on the right. Approximately 20 tooth impressions are preserved on the right tooth row, with replacement teeth visible dorsal to the impressions of the erupted teeth (Figure 3C). There are at least 15 preserved alveolar positions on the left tooth row. As both tooth rows are preserved as incomplete impressions, the total number of teeth within the maxillae is a minimum count.
FIGURE 3
Palatal Bones
Vomer—The vomer is an anteroposteriorly elongate and mediolaterally compressed bone; becoming dorsoventrally tall, posteriorly (Figure 4). Several portions of the nasal septum are missing, but this is likely due to poor mineralization (Witmer and Ridgley, 2008; Leahey et al., 2015). As a result, the dorsal and ventral extents of the vomer are unclear. The ventral nasal keel is partially preserved (Figure 4C). As observed in the CT scans, the anterior half of the vomer forms an elongate triangular process that tapers to a point anteriorly (Figure 4A). This process is dorsoventrally compressed (Figure 4C) with an inverted triangular cross-section and, unlike the posterior half, does not divide the nasal passage. At roughly its mid-point and the mediolateral choanal corner, the vomer mediolaterally expands and is bulbous in dorsal view (Figure 2C). A posterodorsally oriented groove is present on the left lateral surface of the expanded region. This groove is not present on the right lateral surface and is a notable asymmetrical feature (Figures 4C,D). Posterior to the posterodorsal groove, the vomer expands laterally, resulting in an hourglass shape in dorsal view (Figures 2B, 4A).
FIGURE 4
Palatines—The palatines form the posteromedial margins of the choanae and are preserved mostly as impressions, with the right preserving a fragment of bone (Figures 2C, 3B). The palatine is a thin, anteroventrally/posterodorsally angled element, similar to other ankylosaurs, such as Ankylosaurus magniventris and Gargoyleosaurus parkpinorum (Carpenter, 2004; Kilbourne and Carpenter, 2005). Together, the palatines form a U-shaped wall in posterior view that would have separated the palatal and orbital regions, similar to Pawpawsaurus campbelli (Figures 2C, 3B; Lee, 1996). Medially, the palatines contact the vomer. Examining the CT data and physical specimen, we cannot confirm if the palatines and vomer are fused (Figures 2B,C). The palatines extend posterior to the vomer, suggesting that they contacted each other for part of their length; however, the medial edges of the palatines are missing. Along the posterior margin of the choanae, the palatines contact the ectopterygoid along a straight suture that is angled ventrolaterally in posterior view, similar to Ankylosaurus (Figure 3B; Carpenter, 2004).
Ectopterygoid—A partial ectopterygoid is preserved as both bone and impression on the right lateral surface of the skull block, and the left ectopterygoid is partially preserved as an impression (Figures 2C, 3B). The ectopterygoid is sub-triangular in mediolateral view (Figures 5A,B), curved medially, with a concave dorsal margin forming the choanal posterolateral edge (Figures 2B, 5E). The ectopterygoid contacts the maxilla anteriorly, but the contact would have continued laterally to the ectopterygoid along a scarf joint (Figure 6). The ectopterygoid contacts the palatine medially, along the choanal posterior margin, and forms a straight joint angled ventrolaterally (Figure 6). The posteroventral corner is drawn out into a wedged-shaped process (as preserved) buttressed medially along its length (Figure 5).
FIGURE 5
FIGURE 6
Dentition
Eight isolated teeth are identified from the CT scans: one in situ left maxillary tooth crown and seven isolated teeth with roots of unknown position found “floating” within the matrix (Table 1; Supplementary Materials S1, S2). Of these, the in situ crown provides the best resolution and the basis for the following description (Figures 7H–J). The crown is asymmetrical in labial view and is labiolingually compressed (Figures 7I,J). The apex corresponds to the primary ridge/denticle (sensu Bell et al., 2018b) and is distally offset. The primary denticle is marginally larger than the remaining denticles and bears a shallow, mesiodistally oriented, and grooved wear facet on the apex. Two denticles are present distal to the primary denticle and three mesially. Denticles are apically pointed and curved away from the central tooth axis, giving the crown a fan-shaped appearance. Apicobasal grooves (or sulci) between denticles extend basally but do not appear to have reached the cingulum (Figure 7; the cingulum itself is only preserved on the more complete isolated teeth; see above). Individual denticles are difficult to identify on the isolated teeth due to the low resolution of the CT scans; however, small bumps on the crowns suggest five to seven denticles (including the primary denticle) were present per tooth (Figures 7A,E).
TABLE 1
FIGURE 7
The low denticle count is similar to K. ieversi and other nodosaurids but differs from the approximately 8–12 denticles per tooth seen in the upper Strzelecki group teeth (Molnar, 1996; Barrett et al., 2010; Leahey et al., 2015). The in situ crown is broken basally, and the presence of cingula are unknown; however, cingula are present on five isolated teeth. The lingual cingulum is prominent, forming a bulbous shelf, whereas the labial cingulum is less prominent and positioned apically (Figures 7A–F). Five teeth preserve large roots, two to three times taller than the crowns (Table 1), separated by a constriction. Roots are straight, bullet-shaped, and circular in cross-section, similar to other Australian ankylosaurian teeth (Figure 7G; Molnar, 1996; Barrett et al., 2010; Leahey and Salisbury, 2013). Two isolated teeth have partially resorbed roots, as the labial surface is either partially or completely missing (Figure 7B), suggesting root resorption proceeded in a labial–lingual direction. All tooth impressions from the right maxilla have prominent lingual cingula. Consistent with the in situ maxillary crown, tooth impressions have anywhere from five to seven denticles. Crowns of the 16 mesial-most teeth are fan-shaped, whereas the four distal-most teeth are more pointed (Figure 3C). The preserved crown impressions range from 4 to 6 mm wide and 5–8 mm long.
Phylogenetic Analysis
Methods
SAMA P40536 was scored into two phylogenetic datasets (Supplementary Material S3): Arbour and Currie (2016) (Modified from Thompson et al., 2012; Arbour and Currie, 2013; Arbour et al., 2014a; Arbour et al., 2014b) and Soto-Acuña et al. (2021) (Modified from Arbour and Currie, 2016; Arbour et al., 2016). The Arbour and Currie (2016) matrix consists of 178 characters and 45 ingroup taxa, including SAMA P40536 (scores provided in Table 2) and the non-ankylosaurian outgroup taxa Lesothosaurus diagnosticus, Sceliosaurus harrisonii, and Huayangosaurus taibaii. In this matrix, we updated the terminal taxon designation from Minmi sp. to Kunbarrasaurus ieversi, and “Zhejiangosaurus lishuiensis” replaced the incorrectly identified “Zhejiangosaurus luoyangensis” (Lü et al., 2007; Leahey et al., 2015). The Soto-Acuña et al. (2021) matrix consists of 190 characters and 66 ingroup taxa, including SAMA P40536 (scores provided in Table 2) and the non-ankylosaurian outgroup taxa Lesothosaurus diagnosticus, Scelidosaurus harrisonii, and members of Stegosauria (Huayangosaurus taibaii, Paranthodon africanus, and Stegosaurus stenops). Here, we updated the terminal taxon designation from Sauropelta edwardsi to Sauropelta edwardsorum, and “Argentinian ankylosaur” replaced the “Salitral Moreno ankylosaur” for consistency between matrices.
TABLE 2
To incorporate additional palatal variations noted in SAMA P40536, K. ieversi, and other ankylosaurians, we added a new character (178 in Arbour and Currie, 2016; and 190 in Soto-Acuña et al., 2021) that describes the position of the choanae within the palate relative to the maxillary tooth row: choanae with their anterior margins inline or within the anterior third of the maxillary tooth row (178/190:0); choanae posteriorly situated with their anterior margins at least mid-way along the tooth row (178/190:1). Ankylosaurians were scored from the literature (Eaton Jr, 1960; Sereno and Zhimin, 1992; Lee, 1996; Godefroit et al., 1999; Carpenter et al., 2001a, 2008, 2011; Vickaryous et al., 2001; Hill et al., 2003; Carpenter, 2004; Kilbourne and Carpenter, 2005; Parsons and Parsons, 2009; Arbour and Currie, 2013; Arbour et al., 2014a; Leahey et al., 2015; Kinneer et al., 2016; Arbour and Evans, 2017; Yang et al., 2017; Bourke et al., 2018; Paulina-Carabajal et al., 2018; Wiersma and Irmis, 2018; Norman, 2020; Park et al., 2020); however, we could not adequately code the outgroup taxon Huayangosaurus taibaii because the condition of its choanae is unknown. Character polarity was therefore determined from Hesperosaurus mjosi (Maidment et al., 2018) and a 3D cranial model of Stegosaurus armatus (specimen number; UMNH VPC 44, sketchfab. com/ivlpaleontology), which preserve the choanae approximately in line with the anterior-most maxillary tooth (Sereno and Zhimin, 1992; Carpenter et al., 2001b; Chengkai et al., 2007; Mateus et al., 2009). The anterior extent of the choanae in Lesothosaurus diagnosticus is likewise not directly observable. Nonetheless, the maxillae are mediolaterally compressed and unlikely to have formed a bony palate as in many ankylosaurs (Porro et al., 2015). Consequently, we interpret the anterior margins of the choana as probably being anteriorly placed. Finally, the choanae of Scelidosaurus harrisonii end anteriorly relative to the anterior-most maxillary tooth (Norman, 2020). Character 31 was recoded as “?” for K. ieversi in both matrices because the placement of the ectopterygoid cannot be adequately interpreted from the physical specimen or 3D tomographic renderings (Leahey et al., 2015). Moreover, given the absence of a “caudoventral” secondary palate in SAMA P40536, it is doubtful that such as structure was present in K. ieversi.
The updated character-taxon matrices were compiled using Mesquite version 3.61 (Maddison and Maddison, 2019) and analyzed in TNT version 1.5 (Goloboff and Catalano, 2016). All characters were assumed unordered and of equal weight, with Lesothosaurus diagnosticus designated the most distant outgroup taxon in both matrices. Both matrices were subjected to a phylogenetic analysis involving 1,000 replicates of a “traditional” search using random addition sequence starting trees and the Tree Bisection Reconnection (TBR) branch swapping algorithm. Following the initial search, we performed another round of branch swapping on the set of most parsimonious trees (MPTs) using TBR to more fully explore the tree space. We used iterative PCR to identify wildcard taxa that could be pruned to improve resolution in the strict consensus of the final set of MPTs (Pol and Escapa, 2009), while retaining SAMA P40536. Nodal supports for the resulting reduced strict consensus tree from each matrix were calculated from 1,000 bootstrap resampling replicates using the same initial “traditional” tree search strategy. Clade frequencies were summarised using the Groups present/Contradicted (GC) metric.
Results
The phylogenetic analysis of Arbour and Currie (2016) returned 1,630 MPTs of 423 steps [Consistency Index (CI): 0.536, Retention Index (RI): 0.704, Rescaled Consistency Index (RC): 0.377] from the initial tree search, and over 10,000 additional trees of equal length after another round of branch swapping. Sixteen wildcard taxa were identified for removal by iterative PCR: Aletopelta coombsi, Antarctopelta oliveroi, Bissektipelta archibaldi, Dongyangopelta yangyanensis, Glyptodontopelta mimus, Gobisaurus domoculus, Liaoningosaurus paradoxus, Minmi paravertebra, Pawpawsaurus campbelli, Sauroplites scutiger, Scolosaurus cutleri, Shamosaurus scutatus, Stegopelta landerensis, Taohelong jinchengensis, Zhejiangosaurus lishuiensis, and Zipaleta sanjuanensis. The reduced strict consensus tree is almost completely resolved (Figure 8A); Ankylosauridae is fully resolved, whilst Nodosauridae contains one polytomy (Sauropelta edwardsorum, Tianchisaurus nedegoapeferima, and the Argentinian ankylosaur). SAMA P40536 was recovered as the sister of K. ieversi, which together form the sister clade to Mymoorapelta maysi + Ankylosauridae + Nodosauridae (Figure 8A). Kunbarrasaurus ieversi and SAMA P40536 share two autapomorphies: the maxillary tooth row is medially convex (28:1), and the choanae are posteriorly placed approximately mid-way along the maxillary tooth row (178:1). As M. paravertebra was removed by IterPCR, the relationship with SAMA P40536 is currently unknown; however, when the analysis was run with M. paravertebra included, the tree collapsed into a large polytomy (results not included).
FIGURE 8
The initial search of Soto-Acuña et al. (2021) returned 20 MPTs of 693 steps (CI: 0.359, RI: 0.653, RC: 0.234). More than 10,000 trees of equal length were found after another round of branch swapping. Seven wildcard taxa were identified for removal by iterative PCR: Ahshislepelta minor, Denversaurus schlessmani, Donyangopelta yangyanensis, Hylaeosaurus armatus, Sauroplites scutiger, Taohelong jinchengensis, and Zhejiangosaurus lishuiensis. Both Ankylosauridae and Nodosauridae are well resolved in the resulting reduced strict consensus tree (Figure 8B), each containing one polytomy (Ziapelta sanjuanensis and Anodontosaurus lambei, and all three Struthiosaurus, respectively). Parankylosauria is monophyletic but unresolved, recovered as the sister-taxon to Ankylosauridae, Nodosauridae, and a clade formed by Cedarpelta bilbeyhallorum, Chuanqilong chaoyangensis, and Liaoningosaurus paradoxus. Parankylosauria contains Stegouros elengassen, K. ieversi, and A. oliveroi, as in Soto-Acuña et al. (2021), along with SAMA P40536.
Discussion
Comparisons with Kunbarrasaurus ieversi
SAMA P40536 shares five features with Kunbarrasaurus ieversi: a sinuous maxillary tooth row, posteriorly placed choanae, asymmetric tooth crowns, and tooth crown striations that are both confluent with the denticles and extend to the cingulum (Table 2 for clarification on matrix assignment). In addition, SAMA P40536 and K. ieversi share teeth with low denticle counts, cylindrical tooth roots, and both lingual and labial cingula (Molnar, 1996). Unfortunately, none of the K. ieversi autapomorphies identified on the holotype (QM F18101; Leahey et al., 2015) are preserved in SAMA P40536, precluding unambiguous referral. Nevertheless, their dental and choanal similarities are sufficient to recover both specimens as sister-taxa (Figure 8A) and within Parankylosauria with the added tooth characters (Figure 8B). Combined with their spatiotemporal proximity (Figure 9), we provisionally refer SAMA P40536 to cf. Kunbarrasaurus sp. pending further preparation and description of the postcranial skeletons.
FIGURE 9
Although our taxonomic assignment is tentative, the palatal osteology of SAMA P40536 fills anatomical gaps not observable in QM F18101 (Leahey et al., 2015). For example, the palatines of SAMA P40536 are vertically positioned and separate the palatal and orbital regions. Although the posterior extent of the palatines is not preserved, vertical sutures along the posterior margins of the choanae represent the contacts between the palatines and the ectopterygoids (Figures 3A,B). This arrangement indicates that the ectopterygoid forms the posterolateral margin of the choana, as in Edmontonia longiceps (Vickaryous et al., 2004). Furthermore, the palatines did not contact the maxillae along the choanal margin as reconstructed by (Leahey et al., 2015). The ectopterygoid contacts are unknown in K. ieversi due to poor preservation and encasing matrix (Leahey et al., 2015) but differ from the interpretation of (Leahey et al., 2015, p. 22, Figure 6), who placed the ectopterygoid more posteriorly between the maxilla and pterygoids.
Implications of Choanal Variation in Ankylosaurs
The palatal choanae form part of the nasal passages and are the boundary between the internal cranial nasal passages and the buccal region (Bourke et al., 2018). Therefore, it may be expected that choanal variations reflect the complexities of ankylosaurian nasal passages, such as mineralised soft tissue creating convoluted nasal passages and the paranasal sinus system formed by an extensive set of air sacs surrounding the nasal airways. (Brown and Kaisen, 1908; Vickaryous et al., 2004; Vickaryous, 2006; Witmer and Ridgley, 2008; Leahey et al., 2015; Paulina-Carabajal et al., 2016; Bourke et al., 2018). Surprisingly, the palate remains an anatomically under-sampled component of ankylosaur phylogenies. Indeed, only five of the 178 and 190 characters employed in both matrices capture palatal morphologies. We found notable variation in the relative placement of the choana within the palate. The majority of ankylosaurians display choanae that span most of the palatal region, and the anterior choanal margins are either in line with the anterior-most maxillary tooth (e.g., Pawpawsaurus campbelli; Paulina-Carabajal et al., 2016 p. 5) or within the anterior third of the tooth row (e.g., Ankylosaurus magniventris; Carpenter, 2004). As we used the literature to code most ankylosaurian taxa, we simplified the previous conditions into a single character relative to the outgroup (Sereno and Zhimin, 1992; Porro et al., 2015; Maidment et al., 2018; Norman, 2020), indicating that anteriorly placed choanae are primitive for ankylosaurs. By contrast, SAMA P40536 and K. ieversi exhibit a derived condition whereby the choanae are relatively posterior within the palate. The other parankylosaurian Stegouros ellengassen may also exhibit this derived condition, as the secondary palate that marks the anterior extent of the choanae extends posteriorly to approximately the mid-point of the preserved tooth row (Soto-Acuña et al., 2021). Unfortunately, the maxillary tooth row is incomplete posteriorly and so we cannot confirm its condition at this time, and it was coded as “?”. The derived condition stabilizes SAMA P40536 within both trees but otherwise does not dramatically affect the phylogenetic placements of other ankylosaurians. Without this character, the trees collapse into a large polytomy with virtually no basal resolution (results not provided). It is worth noting that if future postcranial observations of SAMA P40536 support an assignment to K. ieversi, this character may represent a new autapomorphy for K. ieversi.
Presently, only the ankylosaurids Cedarpelta bilbeyhallorum, and Gobisaurus domoculus (Vickaryous et al., 2001; Carpenter et al., 2008) exhibit posteriorly positioned choanae comparable to K. ieversi (Leahey et al., 2015) and SAMA P40536. The ankylosaurid Akainacephalus johnsoni and the nodosaurid Panoplosaurus mirus show the most extreme condition, whereby the anterior choanal margins are approximately in line with the posterior-most maxillary tooth (Bourke et al., 2018; Wiersma and Irmis, 2018). These occurrences suggest that posteriorly located choanae evolved independently at least four times: once before the Nodosauridae + Ankylosauridae split, once in nodosaurids, and at least twice in ankylosaurids. Note that there is currently no resolution on the relationships of C. bilbeyhallorum and G. domoculus in Arbour and Currie (2016) and that our safe taxonomic reduction analysis of their matrix excluded G. domoculus. However, our analysis of the Soto-Acuña et al. (2021) matrix places C. bilbeyhallorum outside of Ankylosauridae + Nodosauridae (Figure 8B), which would incur a more complex evolutionary path for the character. However, we refrain from over interpreting this tree given its labile nature (See next Section).
The phylogenetic placement of SAMA P40536 as an early-branching ankylosaurian has implications for interpreting choanal and palatal variations in ankylosaurs. The bones forming the choanal margins are often difficult to discern due to a tightly sutured or fused nature. The preserved palatal bones of SAMA P40536 are not fused, and contacts are visible. The choanal margins are formed by four bones, the maxilla, vomer, ectopterygoid, and palatines (Figure 2). Given that the ectopterygoid contributes to the choanal margin, the palatines were excluded from contacting the maxilla, suggesting that SAMA P40536 did not have a “posteroventral” secondary palate (Vickaryous et al., 2004) or lamina transversa (Bourke et al., 2018); these contacts are unknown in K. ieversi (Leahey et al., 2015). Therefore, our results indicate that the ankylosaurian lamina transversa evolved later in ankylosaur evolution, perhaps due to modifications to the nasal passages (Bourke et al., 2018). Functionally, the lamina transversa separates the olfactory and nasal regions and is associated with a heightened sense of smell (Bourke et al., 2018). Therefore, its absence in SAMA P40536 suggests that a more basic sense of smell was primitive for ankylosaurians.
Implications for Australian Ankylosaur Diversity
Australia has the largest abundance of Gondwanan ankylosaurs and species-level diversity documented from the mid-Cretaceous (Figure 9; Molnar, 2001; Barrett et al., 2010; Leahey and Salisbury, 2013; Leahey et al., 2015; Salisbury et al., 2016; Bell et al., 2018a). Occurrences include ankylosaur footprints from the Valanginian–Barremian Broome Sandstone of Western Australia (Salisbury et al., 2016), isolated skeletal elements from the uppermost Barremian–lower Aptian “Wonthaggi formation” strata of the upper Strzelecki Group in Victoria, Minmi paravertebra from the lower Aptian Bungil Formation of Queensland, Kunbarrasaurus ieversi from the upper Albian Allaru Mudstone of Queensland, and other indeterminate bones and teeth from the uppermost Albian–lower Cenomanian Mackunda and Griman Creek formations of Queensland and New South Wales, respectively (Molnar, 2001; Barrett et al., 2010; Leahey and Salisbury, 2013; Leahey et al., 2015; Bell et al., 2018a). Historically, all Australian ankylosaur fossils were referred to the genus Minmi (Molnar, 1980, 1996); however, the re-classification of K. ieversi (previously Minmi sp.; Leahey et al., 2015) as a separate genus and species implicates greater intra-clade diversity. Our attribution of SAMA P40536 to cf. Kunbarrasaurus sp. further extends the stratigraphical range of this taxon into the lower–upper Albian and evinces a novel sampling occurrence some ∼550 km to the SW of other earlier discoveries.
Previous phylogenies treated K. ieversi (QM F18101) and M. paravertebra (QM F10329) as a generic hypodigm, recovering it as either a sister to all other ankylosaurians (Kirkland, 1998; Carpenter, 2001) or ankylosaurids (Sereno, 1999; Hill et al., 2003; Vickaryous et al., 2004; Ősi, 2005; Burns et al., 2011; Thompson et al., 2012). However, K. ieversi is now considered an early-branching ankylosaurian outside of both Ankylosauridae and Nodosauridae (Arbour and Currie, 2016). Recently, the discovery of Stegouros ellengassen introduced a new phylogenetic hypothesis, whereby K. ieversi, S. ellengassen, and Antarctopelta oliveroi form a Gondwanan ankylosaur clade, Parankylosauria, which would have diverged before the Ankylosauridae + Nodosauridae split (Soto-Acuña et al., 2021). In testing the phylogenetic position of SAMA P40536, we were unable to reproduce the same tree from Soto-Acuña et al. (2021), even when the aforementioned specimen was removed. Our replication attempt produced 20 MPTs and contained several differences in the strict consensus, such as a polytomy outside of Ankylosauridae + Nodosauridae containing Chuanqilong chaoyangensis, Lianingosaurus paradoxus and Cedarpelta bilbeyhallorum, and slightly better resolution within Ankylosauridae and Nodosauridae (Supplementary Material S3; Supplementary Figure S3.2). Furthermore, an additional round of branch swapping on the initial 20 MPTs produced over 10,000 MPTs with considerably reduced resolution in the strict consensus. For instance, there was no clear split between Ankylosauridae and Nodosauridae and most ingroup taxa were reduced to a polytomy (Supplementary Material S3; Supplementary Figure S3.3). The reason for the difference in phylogenetic hypothesis observed in our replication of Soto-Acuña et al. (2021) is unclear; however, it likely stems from the overall poorly supported nature of ankylosaur phylogenetics. Fortunately, these broader differences had no effect on the phylogenetic placement of SAMA P40536 and Parankylosauria was nonetheless recovered in all analyses, supporting the existence of an early-branching, exclusively Gondwanan ankylosaurian clade. Given the placement of SAMA P40536 as the sister taxon of K. ieversi in the Arbour and Currie (2016) phylogeny, it is not surprising that it is also occurs within Parankylosauria. All four terminal units share four dental features: asymmetric tooth crowns, striations confluent with the denticles and extend to the cingula, and cingulum present on either maxillary or dentary teeth. Of note, the first three features are also present in the Argentinian ankylosaur (Coria and Salgado, 2001), although current hypotheses place it deep within Nodosauridae (Figure 8).
Previous studies hypothesized that Australian ankylosaurians evolved independently from the Late Cretaceous ankylosaurs found elsewhere in Gondwana (i.e., A. oliveroi and the Argentinian ankylosaur), which were previously considered to be more closely related to Laurasian nodosaurids (Figure 8A; Coria and Salgado, 2001; Salgado and Gasparini, 2006; Arbour and Currie, 2016; Arbour et al., 2016). However, the proposed existence of Parankylosauria suggests that most Gondwanan ankylosaurs are more closely related to each other than to those found elsewhere (Soto-Acuña et al., 2021). The only current exception is the Argentinian ankylosaur, a nodosaurid (Figure 8) that appeared in Gondwana following dispersal events from Laramidia during the Campanian–Maastrichtian, a pattern also observed among hadrosaurids and titanosaurs (Brett-Surman, 1979; Coria and Salgado, 2001; Prieto-Márquez, 2010; Arbour and Currie, 2016; Ibiricu et al., 2021). Given the mid–upper Albian ages of Australian ankylosaurs, the Gondwanan dispersal of Parankylosauria considerably predated those of the latest Cretaceous (Arbour and Currie, 2016; Kubo, 2020; Soto-Acuña et al., 2021). Interestingly, the recent identification of the putative ankylsoaur Spicomellus afer from the mid-Jurassic of Morocco (Maidment et al., 2021) hints at an initial, more ancient global diversification of Ankylosauria (Gibbons et al., 2013; Arbour and Currie, 2016; Arbour et al., 2016; Maidment et al., 2021; Soto-Acuña et al., 2021).
Conclusion
Here, we described the second Australian ankylosaur cranium and the first ankylosaurian remains from the Toolebuc Formation. Its conferral to Kunbarrasaurus is based on palatal and dental similarities. However, without extensive overlapping anatomy, an unambiguous referral is currently not possible. Nonetheless, several skull elements unknown in the holotype of K. iveresi can be inferred by SAMA P40536, notably the position and morphology of the palatines and their relation to the ectopterygoids. The future examination of SAMA P40536’s postcranial skeleton with those of K. ieversi and Minmi paravertebra will elucidate its taxonomic affinities, further testing the phylogenetic affinities of Australian and Gondwanan ankylosaurs. The exploration of palatal morphology uncovered a new phylogenetic character, and highlights the importance of this anatomical region in resolving ankylosaur phylogenetic relationships.
Data Availability Statement
Raw data and 3D models are available at MorphoSource (ark:/87602/m4/392687): https://www.morphosource.org/projects/000392635?locale=en. Supplementary Material (3D PDF) S1 and S2 are available at Figshare: https://figshare.com/articles/figure/3D_PDF_of_SAMA_P40536-_Frauenfelder_et_al_/16892908.
Author Contributions
TGF and NEC conceived and designed the study, and prepared figures or tables. TGF, PRB, and NEC wrote the manuscript. TGF, PRB, TB, and NEC analyzed the data. JJB scanned fossil material and wrote the associated methods. RDCB created the 3D PDFs. BPK collected fossil material. SW provided access to segmenting software. All authors edited and commented on the manuscript, figures, and tables.
Funding
Synchrotron scanning was facilitated by an IMBL beamtime application (15769) awarded to RDCB and TB. RDCB and TB are financed by UNE Postdoctoral Research Fellowships. This study was funded by a UNE Research Training Program Scholarship to TGF and an Australian Research Council Discovery Early Career Award (DE190101423) to NEC.
Conflict of Interest
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
Publisher’s Note
All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.
Acknowledgments
We thank Mary-Anne Binnie (SAMA) for providing access to study SAMA P40536. Christopher Goatley [University of New England (UNE)] assisted with Micro-CT scanning the in situ maxillary tooth. Anton Maksimenko provided technical assistance with the synchrotron scan data through his IMBL-Stitch software. TNT is made freely available thanks to a subsidy from the Willi Hennig Society. Special thanks to members of the Palaeoscience Research Centre at UNE for discussions related to this study, in particular Kai Allison, Marissa Betts, and John Paterson. Finally, we thank W Zheng, RT Tucker, and one anonymous reviewer for their constructive feedback. BPK acknowledges an Australian Research Council Linkage Project grant (LP0453550), which funded the excavation of SAMA P40536.
Supplementary Material
The Supplementary Material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/feart.2022.803505/full#supplementary-material
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2013年第87卷第3期文章目次
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摘要:At 08:02 on April 20, 2013, a Ms7.0 earthquake occurred in Lushan, Ya’an, in the Longmenshan fault zone, Sichuan. The epicenter was located between Taiping Town and Shuangshi Town, Lushan County and the maximum earthquake intensity at the epicenter reached class IX. Field investigations in the epicenter area found that, although buildings were seriously damaged, no obvious surface rupture structure was produced, only some ground fissures and sand blows and water ejection phenomena being seen. An integrated analysis of high-resolution remote sensing image interpretation, mainshock and aftershock distribution, and focal mechanism solutions indicated that this earthquake was an independent rupturing event in the southwestern segment of the Longmenshan fault zone, belonging to the thrust-type earthquake. Ruptures occurred along the south-central segment of the Shuangshi-Dachuan fault and the principal rupture plane dipped SW at 33–43°. It is inferred that the Lushan earthquake might be related to the ramp activity of the basal detachment zone (13–19 km) of the Longmenshan fault zone. Historically, there occurred at least two Ms6-6.5 earthquakes along the Shuangshi-Dachuan fault zone; thus it is thought that the Lushan earthquake, different from the Wenchuan earthquake, was a characteristic one in the southwestern segment of the Longmenshan fault zone. In-situ stress measurements indicated the Lushan earthquake was the result of stress release of the southwestern segment of the Longmenshan fault zone after the Wenchuan earthquake. This paper analyzes the tectonic setting of the seismogenic structure of this earthquake.
摘要:Geohazards induced by the Lushan Ms 7.0 earthquake on April 20, 2013 mainly have four types: collapse, landslide, slope debris flow, and sand-soil liquefaction. These geohazards mainly occurred near the epicenter, on steep slopes or below cliffs in high mountain and deep valley areas, and at or near fault ends. They have no obvious relationships to active faults, but their relationships to the weathering degree and structures of rock and rock mass are obvious. Compared with the Wenchuan Ms 8.0 earthquake on May 12, 2008, the Lushan earthquake is relatively little in the impact force and the throwing amount. All of these should be related to the magnitude of this earthquake, not very large but not very little. This character of the Lushan earthquake would make some processes uncompleted so as to bring about some concealed geohazards. Finally, in order to deal with challenges presented by such conceal geohazards, some brief recommendations are put forward.
摘要:Dongyangopelta yangyanensis gen. et sp. nov. from the Chaochuan Formation (Albian - Cenomanian) of Dongyang, Zhejiang Province, China is characterized: the convex anterior surface of the first presacral rod centrum strongly inflates laterally and slightly curves posteriorly; the fused pelvic shield composes of larger pebble-shaped bosses, defined by smaller tubercles or flat stretches of bone; most osteoderms are heavily roughened with notches and grooves for dermal attachment along the edge; domed triradiate osteoderm is present; sigmoid curvature of the dorsal surface of the ilium is present; the preacetabular process curves lateroventrally at the anterior end and has a shallow groove in the edge of the lateral and anterior ends and strong lateromedial expansion of the distal femur. The femoral head is well separated from the greater trochanter, indicating that Dongyangopelta is a nodosaurid ankylosaur, the second from southeast China. Phylogenetic analysis also positions this taxon in the Nodosauridae clade. Dongyangopelta differs from Zhejiangosaurus in the characters of presacral rod, ilium, and femur. Dongyangopelta represents the first ankylosaur outside North America and Europe that definitively possesses a pelvic shield with fused armor.
摘要:Based on the systematic study of two fossil skeleton specimens collected from the top of the third member of the marine Lower Triassic Jialingjiang Formation of Yuanan, Hubei Province, South China, a new Early Triassic primitive ichthyosaur Chaohusaurus zhangjiawanensis sp. nov. is reported and described. The beds yielding the type material are correlated with the Neospathodus homeri–N. triangularis Conodont zone. The new taxon is most similar to Chaohusaurus geishanensis Young and Dong, 1972 in the shape and configuration of the scapula, forefin, pelvic girdle and hindfin, presacral vertebral count and well-developed caudal peak, but distinguished from the latter by its larger size, the position of the pineal foramen in the centre of the parietal, the occurrence of a larger calcaneum in hindfin and the first sacral rib with distal expansion. The new species exhibits common features of primitive ichthyosaurs such as: (1) the prefrontal has an antero-dorsal shelf projecting towards the orbit; (2) the upper temporal fenestra is small; (3) the postorbital and the squamosal meet laterally to the upper temporal fenestra; and (4) cylindrical centra. However, more derived ichthyosaur characters are seen with the frontal separated from the orbital dorsal margin by the contact of the prefrontal and postfrontal, which offer new clues to the early radiation of ichthyosaurs.
摘要:Lindera is a large genus of graceful, pleasantly scented and common native trees and shrubs of southern China and neighboring regions of SE Asia. There is a well-documented Cenozoic fossil record not only in these regions but also from elsewhere. A new fossil leaf record has been found in diatomite beds from the Upper Pliocene Mangbang Formation of Tuantian, Tengchong County, Yunnan. The leaves are identified and assigned to Lindera acuminatissima K. Q. Dao et B. N. Sun sp. nov., by comparing their leaf architecture and epidermal characteristics with those of 51 extant Lauraceae species and with 15 known fossil Lindera taxa. The specimens have well-preserved cuticles, with typical leaf architecture and epidermal characteristics of the Lauraceae, including entire leaf margin, intramarginal veins, basal ternate acrodromous primary veins, one-cell trichome base, paracytic stomatal apparatus, sunken guard cells, subsidiary hardly staining cells and presence of oil cells. These characteristics are consistent with Lindera sect. Daphnidium but are different from reported fossil and extant species of Lindera. The cuticles of Lindera are fragile and delicate with only three Lindera fossils reported based on this tissue. In terms of paleobiogeography, the fossil record indicates that Lindera is distributed in high- to mid-latitude regions of the Late Cretaceous to Paleocene northern hemisphere. Coincidentally, the records of Lindera located on both sides of the Bering Land Bridge possibly support the hypothesis that ancient plants extended via transcontinental exchanges through the Bering Corridor. In the Eocene, ancient Lindera spread to Europe through the Northern Degeer Route and the Southern Thulian Route. At the same time, ancient Lindera spread into Central Asia. Climatic changes and tectonization since the Neogene prevented the propagation of Lindera throughout Asia, North America and Europe, and hence the distribution areas have just regressed to the low-latitude regions in Asia and North America. From the Paleogene to the Neogene, Lindera has changed its distribution by surviving extreme climate changes. Quaternary glaciations ultimately led to Lindera becoming extinct in Europe. The new record from Tengchong, Yunnan, with its lower latitude located in tropical and subtropical regions, indicates that Lindera has lived in those regions since the late Pliocene.
摘要:Based on high-resolution remote sensing image interpretation, digital elevation model 3-D analysis, field geologic field investigation, trenching engineering, and ground-penetrating radar, synthetic research on the evolution of the Yuguang Basin South Margin Fault (YBSMF) in northwest Beijing was carried out. We found that the propagation and growth of faults most often occurred often at two locations: the fault overlapping zone and the uneven or rough fault segment. Through detailed observation and analysis of all cropouts of faults along the YBSMF from zone a to zone i, we identified three major factors that dominate or affect fault propagation and growth. First, the irregularity of fault geometry determine the propagation and growth of the fault, and therefore, the faults always propagate and grow at such irregular fault segments. The fault finally cuts off and eliminates its irregularity, making the fault geometry and fault plane smoother than before, which contributes to the slipping movement of the half-graben block in the basin. Second, the scale of the irregularity of the fault geometry affects the result of fault propagation and growth, that is, the degree of the cutting off of fault irregularity. The degree of cutting off decreases as irregularity scale increases. Third, the maximum possible slip displacement of the fault segment influences the duration of fault propagation and growth. The duration at the central segments with a large slip displacement is longer than that at the end segments with a smaller slippage value.
摘要:Many equiaxial dome-like structures developed in the north segment of the Xuefengshan orocline, Central China are obviously inconcordant with the NE-trending linear structures in this area, which contain important records for understanding the structural framework and evolution of this belt. In this paper, taking one of the typical dome-like structures in the Xuefengshan orcline (e.g. Moping dome-like structure) as an example, based on its structural framework interpratatoin, superposed deformation analysis and paleo-stress fields reconstruction, we propose the Moping dome-like structure is composed of two populations of different-striking thrust-fold structures, ~E-trending and NE-striking structures, indicative of two-stages shortening, ~N- and NW-striking, respectively. Together with the geochronological analysis, we suggest the first stage of shortening occurred in Late Triassic to Early Jurassic, due to the Indosinian intercollisional orogeny of the Yangtze Block and the North China Block. The second occurred during Late Jurassic?Early Cretaceous owing to Yanshanian intracontinental orogeny, leading to the intensive superposition of the NE-trending structures onto the ~E-trending structures, and the final ocurrence of the Moping dome. Thus, our study indicates the Xuefengshan arc-shape belt also experienced two-phase deformation, and resulted from the superposition of NE?SW structures onto ~E–W structures in Late Jurassic?Early Cretaceous, which could provide new structural evidence for probing the Mesozoic tectonic framework and evolution of the Xuefengshan orocline.
摘要:The Late Cretaceous ü?kap?l? Granitoid including mafic microgranular enclaves intruded into metapelitic and metabasic rocks, and overlain unconformably by Neogene ignimbrites in the Ni?de area of Turkey. It is mostly granite and minor granodiorite in composition, whereas its enclaves are dominantly gabbro with a few diorites in composition. The ü?kap?l? Granitoid is composed mainly of quartz, K-feldspar, plagioclase, biotite, muscovite and minor amphibole while its enclaves contain mostly plagioclase, amphibole, minor pyroxene and biotite. The ü?kap?l? Granitoid has calcalkaline and peraluminous (A/CNK= 1.0–1.3) geochemical characteristics. It is characterized by high LILE/HFSE and LREE/HREE ratios ((La/Lu)N= 3–33), and has negative Ba, Ta, Nb and Eu anomalies, resembling those of collision granitoids. The ü?kap?l? Granitoid has relatively high 87Sr/86Sr(i) ratios (0.711189–0.716061) and low εNd(t) values (-5.13 to -7.13), confirming crustal melting. In contrast, the enclaves are tholeiitic and metaluminous, and slightly enriched in LILEs (K, Rb) and Th, and have negative Ta, Nb and Ti anomalies; propose that they were derived from a subduction-modified mantle source. Based on mineral and whole rock chemistry data, the ü?kap?l? granitoid is H-(hybrid) type, post-collision granitoid developed by mixing/mingling processes between crustal melts and mantle-derived mafic magmas.
摘要:Authigenic gypsum crystals, along with pyrite and carbonate mineralization, predominantly calcites were noticed in distinct intervals in a 32 m long piston core, collected in the gas hydrate-bearing sediments in the northern portion of the Krishna-Godavari basin, eastern continental margin of India at a water depth of 1691 m. X-ray diffraction and energy dispersive spectrum studies confirm presence of pyrite, gypsum, calcite, and other mineral aggregates. The occurrence of gypsum in such deep sea environment is intriguing, because gypsum is a classical evaporite mineral and is under saturated with respect to sea water. Sedimentological, geochemical evidences point to diagenetic formation of the gypsum due to oxidation of sulphide minerals (i.e. pyrite). Euhedral, transparent gypsum crystals, with pyrite inclusions are cemented with authigenic carbonates, possibly indicating that they were formed authigenically in situ in the gas hydrate-influenced environment due to late burial diagenesis involving sulphate reduction and anaerobic oxidation of methane (AOM). Therefore, the authigenic gypsums found in sediments of the Krishna-Godavari and Mahanadi offshore regions could be seen as one of the parameters to imply the presence of high methane flux possibly from gas hydrate at depth.
摘要:Chemoautotrophic organisms have once been excluded from the development of universally applicable CO2 fixation technology due to its low production yields of biomass. In this study, we used Acidithiobacillus ferrooxidans (A.f.) as a model chemoautotrophic microorganism to test the hypothesis that exogenetic photoelectrons from semiconducting mineral photocatalysis can enable the regeneration of Fe2+ that could be then used by A.f. and support its growth. In a simulated electrochemical system, where exogenetic electrons were provided by an electrochemical approach, an accelerated growth rate of A.f. was observed as compared with that in traditional batch cultivation. In a coupled system, where light-irradiated natural rutile provided the primary electron source to feed A.f., the bacterial growth rate as well as the subsequent CO2 fixation rate was demonstrated to be in a light-dependent manner. The sustaining flow of photogenerated electrons from semiconducting mineral to bacteria provided an inexhaustible electron source for chemoautotrophic bacteria growth and CO2 fixation. This finding might contribute to the development of novel effective CO2 fixation technology.
摘要:The Shenhu gas hydrate drilling area is located in the central Baiyun sag, Zhu II depression, Pearl River Mouth basin, northern South China Sea. The gas compositions contained in the hydrate-bearing zones is dominated by methane with content up to 99.89% and 99.91%. The carbon isotope of the methane (δ13C1) are ?56.7‰ and ?60.9‰, and its hydrogen isotope (δD) are ?199‰ and ?180‰, respectively, indicating the methane from the microbial reduction of CO2. Based on the data of measured seafloor temperature and geothermal gradient, the gas formed hydrate reservoirs are from depths 24–1699 m below the seafloor, and main gas-generation zone is present at the depth interval of 416–1165 m. Gas-bearing zones include the Hanjiang Formation, Yuehai Formation, Wanshan Formation and Quaternary sediments. We infer that the microbial gas migrated laterally or vertically along faults (especially interlayer faults), slump structures, small-scale diapiric structures, regional sand beds and sedimentary boundaries to the hydrate stability zone, and formed natural gas hydrates in the upper Yuehai Formation and lower Wanshan Formation, probably with contribution of a little thermogenic gas from the deep sedments during this process.
摘要:It is important to determine the properties of the tectonics in Cambrian period for the sake of prospecting deep hydrocarbon in the near future in the southern Ordos Kratogen of North China. Authors chose the marginal areas of the southern Ordos basin as the object of research, avoided the effects of both the Qinling Orogenic Belts (QOB) and Weihe River Graben (WRG) whose geological structures are too complicated. By surveying typical Cambrian outcrops and profiles in the basin edges and based on the cores of 57 wells which penetrated the Cambrian in the basin, combined with the seismic profiles, the field gammaray measuement results and the carbon isotope analysis, Authors conclude that the southern margin of the Ordos Kratogen during Cambrian was a passive continental margin which resulted from sea–floor spreading of the Ancient Qinling Ocean. Epicontinental sea carbonate sediments formed in the south Ordos continental margin during Cambrian, and were predominant as tidal flat and o?litic shoal. Both transgression–regression process and the change in palaeostructure have the obvious cyclicity. Using the junction between the late Nangao age of Qiandong epoch and the early Duyun age of Qiandong epoch as a boundary, each had a full transgression cycle at the upper and lower stages. The early cycle is characterized by high energy clastic littoral facies while the late cycle is characterized by carbonate ramp on which clear water and muddy water developed alternately changing to carbonate platform last. During the early stages, An aulacogen was formed in the middle section of the southern margin. The southern Ordos margin was uplifted and denudated by the Huaiyuan Movement which occurred from the late Furongian age to the middle Flolan age and the history of the passive continental margin ended and entering into a new tectonic cycle. The unconformity surface caused by the Huaiyuan Movement, along with its neighborhood areas where dissolved pores and cavities are developed, may be another important district for good hydrocarbon reservoirs (excluding the unconformity surface on the top of the Ordovician in the Ordos basin).
摘要:BSR (Bottom Simulating Reflector) occurs widely in the strata since the late Miocene in the deep-water area of the northern continental slope of South China Sea (SCS). It is an important seismic reference mark which identifies the gas hydrate and its distribution influenced by the tectonic movements. Single-point basin modeling was conducted using 473 points in the study area. To discuss the relationships between the tectonic subsidence and BSR, the volume and rate of tectonic subsidence in each geological time have been simulated. The results show that there are three tectonic accelerate subsidence processes in the study area since the late Miocene, especially since 1.8Ma the tectonic subsidence accelerates more apparently. Since the Late Miocene to Pleistocene, the rate of tectonic subsidence in deep-water underwent a transformation from weak to strong. The ratio of tectonic subsidence to the total subsidence was relatively high (65-70%). Through the superposition of the BSR developed areas and the contours of tectonic subsidence in this area, it was discovered that more than 80% of BSR tend to be distributed at the slope break or depression-uplift structural transfer zone and the average tectonic subsidence rate ranges from 70 m/Ma to 125 m/Ma.
摘要:In order to understand the origin and flow of formation water and to evaluate the hydrocarbon accumulation and preservation conditions, the properties of formation water chemistry and dynamics of the Zhenwu area in the southern Gaoyou Sag, North Jiangsu Basin, China, have been investigated. The results show that Xuzhuang oilfield is infiltrated discontinuously by meteoric water under gravity, which consequently leads to the desalination of formation water. Formation water in the Zhenwu and Caozhuang oilfields is less influenced by meteoric water infiltration, and the origin is interpreted to be connate water. Hydrocarbon migration, accumulation and preservation are closely related to the hydrodynamic field of formation water. Formation water concentrates gradually during the process of centrifugal flow released by mudstone compaction and the centripetal flow of meteoric water infiltration, leading to the high salinity of the central part. The geological conditions of the southern fault–terrace belt are poor for hydrocarbon accumulation and preservation as meteoric water infiltration, leaching and oxidation, while the central part, i.e., northern Zhenwu and Caozhuang oilfields is beneficial for an abundance of hydrocarbon accumulation. Most of the large scale oil–gas fields locate herein.
摘要:Eastern Iran has great potential for the discovery of different types of mineralization. The study area encompasses Tertiary magmatism in the northern Lut block located in northern Khur, South Khorasan, eastern Iran and is mostly covered by volcanic rocks, which are intruded by porphyritic subvolcanic intrusions in some places. Application of the spectral angle mapper (SAM) technique to Advanced Spaceborne Thermal Emission and Reflection Radiometer (ASTER) images detected sericitic, argillic, and propylitic alterations, silicification, and secondary iron oxides. The alteration is linear and associated within vein-type mineralization. Twelve prospective areas are selected for detailed exploration and based on our processing results, in addition to NW–SE faults, which are associated with Cu mineralization indications, NE–SW faults are also shown to be important. Based on the presence of subvolcanic rocks and numerous Cu ± Pb-Zn vein-type mineralizations, extensive alteration, high anomaly of Cu and Zn (up to 100 ppm), the age (43.6 to 31.4 Ma) and the initial 87Sr/86Sr ratio (0.7047 to 0.7065) of the igneous rocks, and the metallogenic epoch of the Lut block (middle Eocene–lower Oligocene) for the formation of porphyry Cu and epithermal deposits, the studied area shows great potential for porphyry copper deposits.
摘要:This paper discusses the enrichment and depletion regularities for porphyry copper-molybdenum ore deposits in different regions and varied deposit genetic types in the same area, taking three porphyry copper-molybdenum ore deposits (i.e., the Chengmenshan in Jiangxi, Wunugetushan in Inner Mongolia, Baishantang in Gansu) and two copper deposits in Gansu Province (the Huitongshan skarn deposit and Gongpoquan composite deposit) as case studies. The results show that porphyry Cu-Mo deposits or skarn copper deposits include both enrichment of the ore-forming elements and associated elements, and depletion of some lithophile dispersed elements, rare earth elements (REE) and some major elements. And the depleted elements vary with deposits, having generality and their own features. On a deposit scale, the positive anomalies of enriched elements and negative anomalies of depleted elements follow in a sequence to comprise regular anomaly models of spatial structures. The exploration in the Tongchang deposit in Jiangxi and Huitongshan deposit in Gansu suggests that anomaly models play a key role in the identification of mineral occurrences and deposits compared to one single enriched element anomaly. And the anomaly models exert a critical effect on the optimization of prospecting targets and their potential evaluation.
摘要:An anisotropic geomechanical model for jointed rock mass is presented. Simultaneously with deriving the orthotropic anisotropy elastic parameters along the positive axis, the equivalent compliance matrix for the deflection axis orthotropic anisotropy was derived through a three-dimensional coordinate transformation. In addition, Singh's analysis of the stress concentration effects of intermittent joints was adopted, based on two groups of intermittent joints and a set of cross-cutting joints in the jointed rock mass. The stress concentration effects caused by intermittent joints and the coupling effect of cross-cutting joints along the deflection-axis are also considered. The proposed anisotropic mechanics parameters method is applied to determine the deformation parameters of jointed granite at the Taishan Nuclear Power Station. Combined with the deterministic mechanical parameters of rock blocks and joints, the deformation parameters and their variability in jointed rock masses are estimated quantitatively. The computed results show that jointed granite at the Taishan Nuclear Power Station exhibits typical anisotropic mechanical characteristics; the elastic moduli in the two horizontal directions were similar, but the elastic modulus in the vertical direction was much greater. Jointed rock elastic moduli in the two horizontal and vertical directions were respectively about 24% and 37% of the core of rock, showing weakly orthotropic anisotropy; the ratio of elastic moduli in the vertical and horizontal directions was 1.53, clearly indicating the transversely isotropic rock mass mechanical characteristics. The method can be popularized to solve other rock mechanics problems in nuclear power engineering.
摘要:Identifying the provenance of aeolian sediments in the Hunshandake Sandy Land is of great importance for understanding the formation of the dune fields in the mid-latitudes and for deciphering information about desert’s responses to global change. By determining the major and trace elements concentrations of aeolian sands in three grain size fractions from the central and western parts of the Hunshandake Sandy Land, we systematically study the provenance and the depositional history of aeolian sands in this desert environment. Our results show that aeolian sands from the Hunshandake Sandy Land are enriched in SiO2 and are depleted in many other elements compared to those of the Upper Continent Crust (UCC). Variations of the immobile elements ratios like Zr/Hf, La/Yb, Th/Nb, La/Nb, LaN/YbN, GdN/YbN are relatively large in the coarse and medium fractions but minor in the fine fractions. Eu anomalies are quite different in the coarse fractions, but mostly positive in the medium fractions and all negative in the fine fractions. Decreasing tendency of Zr concentrations from the west to the east in the Hunshandake Sandy Land is evident in the coarse sands but rather weak in the fine grain size fractions. Our geochemical data indicate that the sources for the coarse and medium fractions of aeolian sands are diverse, influenced by local geology and geomorphology, while the fine sand fractions are more homogenous due to intensive mixture mainly by aeolian processes. Various ratios of immobile elements suggest that these sands should be sourced primarily from the surrounding mountains by fluvial/alluvial processes rather than from any remote territories. Aeolian sands with Ce negative anomalies are widely distributed in the Hunshandake Sandy Land, indicating that aquatic environments have occurred extensively prior to the occurrence of the dune field.
摘要:The knowledge of Martian salts has gone through substantial changes during the past decades. In the 70th of last century, Viking landers have noticed the existence of salts on Mars. Several salt species have been suggested from then on, such as sulfates and chlorides. However, their origin was a mystery due to the lack of observations. The recent explorations and related studies at the beginning of this century revealed that the crustal composition of Mars is similar to that of Earth, and it was hypothesized that almost one third of Martian surface was covered by oceans and lakes in the early stage of Mars. The huge water bodies may have dissolved a large quantity of ions from Martian primary rocks during the whole Noachian and Hesperian epoch. After the enormous drought event happened during the late Hesperian and the early Amazonian, these dissolved ions have formed huge salts deposits and most of them were preserved on Mars until today. To date, carbonates, sulfates, chlorides have all been detected by orbital remote sensing and by landers and rovers. However, the salt mineral assemblages on Mars seems to have some differences from those on Earth, e.g., rich in sulfates and lack of massive carbonates. To explain this difference, we propose that most of the surface carbonates precipitated from the ancient oceans may have been dissolved by the later ubiquitous acidic fluids originated from the global volcanism in the Hesperian era, and formed the enormous sulfate deposits as detected, and this hypothesis seems to be supported by the evidence that most of the sulfate deposits distribute around the Tharsis volcanic province while the survived carbonates located far from it. This process can release most of the carbon on Mars to the atmosphere in the form of CO2 and then be erased by the late heavy bombardments, which might have profound influence on the climate change happened in the Hesperian age. The positive correlation between the GRS results of the potassium distributions and the distribution of chlorides on Mars, together with the high Br concentration measured from the evaporate sediments at two Mars exploration rover landing sites, indicate that the brines in the regions where the chlorides deposited may have reached the stage for potassium salts deposition, thus we propose for the first time that potassium salts deposits might be prevalent in these regions.
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Greetings!
This page is dedicated to keeping you updated with a list of all known genera of Mesozoic dinosaurs, arranged according to the groups to which they belong. This list is provided as an Adobe PDF format, which is readable by most web browser programs and printable on most computers. If you need to get a copy of Adobe Acrobat Reader, you can go here to download a copy of the software.
I also include an expanded version of the introduction to the dinosaur genus list. This includes material that was going to be in the published version of the book, but had to be cut out to save on space.
I will try to update this list when the opportunity arises, hopefully once a season. The first update was 31 July 2008. That update included the addition of 108 new genera and 55 new classification categories. If you are interested, here are the new additions:
New genera added for 31 July 2008 are: 26 genera awaiting official names, plus Achillesaurus, Albertonykus, Alethoalaornis, Aniksosaurus, Aristosaurus, Asylosaurus, Australodocus, Balochisaurus, Berberosaurus, Brohisaurus, Cedrorestes, Cerasinops, Cerebavis, Dakotadon, Dalingheornis, Dashanpusaurus, Diceratus, Didactylornis, Dollodon, Dongbeititan, Dongyangosaurus, Dracovenator, Dromomeron, Elsornis, Enantiophoenix, Eocarcharia, Eoconfuciusornis, Eocursor, Eomamenchisaurus, Fusuisaurus, Futalognkosaurus, Gigantoraptor, Gigantspinosaurus, Glacialisaurus, Huanghetitan, Khetranisaurus, Koutalisaurus, Lamplughsaura, Liaoningornis, Lophostropheus, Loricatosaurus, Luanchuanraptor, Macrogryphosaurus, Mahakala, Mantellisaurus, Marisaurus, Martinavis, Maxakalisaurus, Microceratus, Muyelensaurus, Nanningosaurus, Nanosaurus, Nopcsaspondylus, Orkoraptor, Ornithotarsus, Oryctodromeus, Othnielosaurus, Pakisaurus, Paluxysaurus, Pantydraco, Paraprotopteryx, Pengornis, Pradhania, Qingxiusaurus, Sacisaurus, Sahaliyania, Shanag, Sinocalliopteryx, Sulaimanisaurus, Suzhousaurus, Theiophytalia, Turiasaurus, Uberabatitan, Urbacodon, Velafrons, Wulagasaurus, Xenoposeidon, Xuanhuaceratops, Yamaceratops, Zhejiangosaurus, Zhongornis, Zhongyuansaurus, and Zhuchengosaurus
New genera added for 13 January 2011 are: 6 genera awaiting an official name, plus Aardonyx, Abydosaurus, Adeopapposaurus, Aerosteon, Ajkaceratops, Alamitornis, Albalophosaurus, Anatosaurus, Anchiornis, Angulomastacator, Arkharavia, Arenysaurus, Asilisaurus, Astinganosaurus, Australovenator, Austrocheirus, Austroraptor, Balaur, Banji, Baotianmansaurus, Barilium, Barrosasaurus, Bauxitornis, Beishanlong, Bishtahieversor, Blasisaurus, Bolong, Ceratonykus, Chromogisaurus, Chuxiongosaurus, Coahuilaceratops, Concavenator, Cruxicheiros, Daxiatitan, Demandasaurus, Diabloceratops, Diamantinasaurus, Duriatitan, Duriavenator, Dyoplosauurus, Dysalotosaurus, Elbretornis, Elrhazosaurus, Eodromaeus, Epidexipteryx, Flexomornis, Fruitadens, Fukuititan, Geminiraptor, Glishades, Haplocheirus, Helioceratops, Hesperonychus, Hippodraco, Hollanda, Huoshanornis, Hypselospinus, Ignavusaurus, Iguanacolossus, Intiornis, Jeyawati, Jianchangornis, Jintasaurus, Kayentavenator, Kemkemia, Kileskus, Kinnareemimus, Kol, Koreaceratops, Koreanosaurus, Kosmoceratops, Kukufeldia, Leshanosaurus, Lesnesovia, Limusaurus, Linheraptor, Liubangosaurus, Longicrusavis, Luoyanggia, Malarguesaurus, Machairasaurus, Medusaceratops, Minotaurasaurus, Miragaia, Naramguenatitan, Ojoceratops, Owenodon, Paludititan, Panamericansaurus, Panphagia, Peloroplites, Pitekunsaurus, Pneumatoraptor, Proplanicoxa, Qiaowanglong, Qiupalong, Rahiolisaurus, Rapaxavis, Raptorex, Rubeosaurus, Ruyangosaurus, Sanjaunsaurus, Sarahsaurus, Seitaad, Sellacoxa, Shanweiniao, Shaochilong, Shenshiornis, Shenqiornis, Shidaisaurus, Similicaudipteryx, Sinoceratops, Sinotyrannus, Skorpiovenator, Spinophorosaurus, Tastavinsaurus, Tatankacephalus, Tatankaceratops, Tawa, Tethyshadros, Texacephale, Tianyulong, Tianyuraptor, Titanoceratops, Tongasaurus, Traukutitan, Utahceratops, Vagaceratops, Wintonotitan, Xianshanosaurus, Xiongguanlong, Xixiankykus, Xixiasaurus, Zanabazar, Zhongjiangornis, Zhuchengceratops, and Zuolong ("General Tso's dinosaur").
Genera removed for 6 January 2011: Diceratus (now in Triceratops) and Homalocephale (now in Prenocephale).
New genera added for 10 January 2012 are: eight genera awaiting an official name, plus Acristavus, Ahshishapelta, Albinykus, Amtocephale, Angolatitan, Arcusaurus, Atacamatitan, Bohaiornis, Bonapartenykus, Brontomerus, Cathayornis, Daemonosaurus, Delapparentia, Diodorus, Drusilasaura, Epichirostenotes, Gracilornis, Gryphognathus, Haya, Huaixasaurus, Leonerasaurus, Leyesaurus, Linhenykus, Linhevenator, Manidens, Mystiornis, Nambalia, Jaklapallisaurus, Ojoraptorsaurus, Osmakasaurus, Oxalaia, Pampadromaeus, Pamparaptor, Parahonshanornis, Petrobrasaurus, Propanoplosaurus, Qiliania, Ratchasimasaurus, Spinops, Talos, Tapuiasaurus, Teratophoneus, Unescoceratops, Uteodon, Veterupristisaurus, Xiaotingia, Xuwulong, Yueosaurus, Zuchengtyrannus
Genera removed for 31 December 2011: Bugensaura (now in Thescelosaurus (Chapter 30); Dollodon and Proplanicoxa (now in Mantellisaurus) and Kukufeldia and Sellicoxa (now in Barilium) (Chapter 31); Anatotitan (now in "Anatosaurus") (Chapter 32)
New genera added for 19 January 2015 are: Acheroraptor, Ajancingenia, Alnashetri, Anzu, Aorun, Arcovenator, Aurornis, Bicentenaria, Camarillasaurus, Changyuraptor, Datanglong, Eoabelisaurus, Eosinopteryx, Fosterovenator, Gobivenator, Gryphoceratops (replaces "Gryphognathus"), Hexing, Ichthyovenator, Ignotosaurus, Jianchangosaurus, Jiangxiasaurus, Juratyrant, Leptorhynchos, Lutungutali, Lythronax, Martharaptor, Nankangia, Nanuqsaurus, Ningyuansaurus, Nyasasaurus, Ostafrikasaurus, Panguraptor, Pectinodon, Philovenator, Qianzhousaurus, Siats, Sauroniops, Tachiraptor, Wulatelong, Yulong, Yurgovuchia, Yutyrannus
Genera removed for 19 January 2015: Crosbysaurus, Galtonia, Gryphognathus (replaced with Gryphoceratops), Krzyzanowskisaurus, Lucianosaurus, Pekinosaurus, Protecovasaurus, Razanandrongobe, Sigilmassasaurus (now in Spinosaurus), Tecovasaurus, Trialestes, Wellnhoferia (now in Archaeopteryx)
New classification categories added for 31 July 2008 are (in order of appearance in the book and table): Lagerpetonids, Silesaurs (Chapter 11); Dilophosaurids and relatives (Chapter 13); "Megalosaurs", Megaraptors (Chapter 14); Primitive Oviraptorids, Elmisaurines, "Ingeniines" (Chapter 19); Sapeornithids, Confuciusornithids, Primitive Enantiornithines, Primitive Euenantiornithines, Avisaurids, Gobipterygids, Longipterygids, Primitive Euornithines, Yanornithiforms, Advanced Euornithines (Chapter 21); Primitive Sauropodomorphs, Plateosaurids, Riojasaurids, Massospondylids, Near-Sauropods (Chapter 22); Primitive Sauropods, Vulcanodontids, Primitive Eusauropods, Primitive Cetiosaurids, Mamenchisaurines, Turiasaurs, Primitive Neosauropods (Chapter 23); Apatosaurines, Diplodocines (Chapter 24); Argyrosaurids, Aeolosaurids, Lognkosaurs, Antarctosaurids, Nemegtosaurids (Chapter 25); Primitive Stegosaurs, Stegosaurids (Chapter 28); Primitive Neornithischians, Zephyrosaurs(Chapter 30); Dryosaurids, Camptosaurids, Primitive Styracosternans (Chapter 31); Primitive Lambeosaurines, Parasaurolophinins, Corythosaurinins, Primitive Hadrosaurines, Gryposaurinins, Saurolophinins, Edmontosaurinins (Chapter 32); Primitive Pachycephalosaurs, Pachycephalosaurids (Chapter 33); Chaoyangsaurids and Other Primitive Ceratopsians, Psittacosaurids (Chapter 34)
New classification categories added for 6 January 2011 are (in order of appearance in the book and table): Silesaurids (replaces silesaurs) (Chapter 11); Herrerasaurs (replaces "Herrerasaurids"), Primitive Theropods (Chapter 12); Elaphrosaurs, Ceratosaurids (Chapter 13); Primitive Megalosauroids, Megalosaurids (Chapter 14); Primitive Carcharodontosaurs, Primitive Neovenatorids, Megaraptorans (Chapter 15); Coelurids (Chapter 16); Proceratosaurids, Near-Tyrannosaurids (replaces "Primitive Tyrannosaurids") (Chapter 17); Primitive Alvarezsauroids, Parvicursorines (replaces "Mononykines") (Chapter 18); Saurornitholestines (Chapter 20); Archaeopterygids, Scansoriopterygids, Omnivoropterygids (replaces "Sapeornithids") (Chapter 21); Guaibasaurids (Chapter 22); Huayangaosauridae, Primitive Stegosaurids, Dacentrurines, Stegosaurines (Chapter 28); Thescelosaurids (Chapter 30); Primitive Hadrosaurians (replaces "Primitive Hadrosauroids"), Maiasaurinins (Chapter 32); Bagaceratopsids (Chapter 34); Primitive Ceratopsids, Chasmosaurines (replaces "Ceratopsines") (Chapter 35)
New classification categories added for 12 December 2012 are (in order of appearance in the book and table): Elmisaurids (replaces "Elmisaurines"), Oviraptorids (replaces "Oviraptorines") (Chapter 19); Primitive Ankylosaurids and Ankylosaurines (former just "Ankylosaurids") (Chapter 29); Changchungsaurs (previously included in "Primitive Ornithopods") (Chapter 30); Primitive Saurolophines and Brachylophosaurinins (the latter replaces "Maiasaurinins") (Chapter 32)
New classification categories added for 19 January 2015 are (in order of appearance in the book and table): Primitive Neotheropods, Primitive Abelisaurids, Majungasaurines, Primitive Brachyrostrans, Carnotaurinins (the last four replace "Abelisaurids") (Chapter 13); Piatnitzkysaurids (replaces part of "Primitive Megalosauroids"), Primitive Megalosaurids, Afrovenatorines, Megalosaurines (the last three replace "Megalosaurids"), Primitive Spinosaurids, Baryonychines, Spinosaurines (the last three replace "Spinosaurids") (Chapter 14); Primitive Metriacanthosaurids, Metriacanthosaurines (the last two replace "Sinraptorids"), Primitive Carchaodontosaurids, Gigantosaurinins (the last two replace "Carcharodontosaurids"), Primitive Megaraptorans, Megaraptorids (the last two replace "Megaraptors") (Chapter 15); Deinocheirids (Chapter 18); Caudipterids (Chapter 19); Primitive Eumaniraptorans, Anchiornithines, Jinfengopterygines, Advanced Troodontids (the last three replace "Troodontids") (Chapter 20)
Classification categories removed for 2 January 2015 are (in order of original appearance in the book and table): Dilophosaurids (now found to be a collection of primitive theropods from different parts of the family tree) (Chapter 13); Primitive Caenagnathoids (Chapter 19); Archaeopterygids (Chapter 21);
These new additions are highlighted on the list with an "*" if it is a new genus, with a "^" it is a new name for a genus without a genus name in the original list; and a "**" if it is a new classification category.
Expanded Introductory Information:
In the following list, I've arranged all the known genera of Mesozoic dinosaurs according to the groups to which they belong. A genus, as you may recall, (if not, see chapter 7) is the one-word name that we typically use when talking about dinosaurs. Tyrannosaurus, Triceratops, and Ankylosaurus are all examples of a genus. Each genus is a group of one or more species. Most dinosaur genera are known from only a single species, but a few (such as Psittacosaurus, Apatosaurus, and Edmontosaurus) are known from several different species. Each group of genera is shown in the same order as their respective chapters in this book.
Some things to note: I have not included genera of Cenozoic dinosaurs--that is, modern birds and the other birds that lived after 66 million years ago--in this list. There are simply too many of them! And since dinosaurs are being discovered and named at a rate of about two new genera per month, this list will lack a few recent names. (In an illustrated book such as this, there is a minimum time of approximately three months between when the text is released to print and when finished books are available in bookstores and libraries.) Also, in some cases, the placement of a particular genus in a group is very uncertain. This can happen when a genus is known only from a very incomplete fossil, or when it has a confusing mixture of features. I've excluded dinosaur genus names that are based on material so fragmentary that it is very difficult to say what groups they belong to.
Some dinosaurs are currently without proper names. For example, there is a dinosaur that was once called "Ingenia." Unfortunately, there is also a nematode worm called Ingenia, and it was named first! So by the proper rules of naming, the worm keeps the name and the dinosaur needs a new one. In 2013, the "Ingenia" dinosaur was given the new genus name Ajancingenia.
Also, there are dinosaur fossils that were once considered new species in previously named genera, but which turn out not to really belong to those, or to any other already named, genera. These will eventually get their own genus name when newer studies are completed. Because some of these species are interesting (maybe they are the only member of their group from a particular time or region; maybe they have some peculiar feature), I've put them in the appendix, too. These unnamed (or at least un-genus-named) dinosaurs are being studied, and as those names are finally given I'll make sure they get included in future versions of this list.
Also, paleontologists and biologists who work on modern animals, I might add, sometimes disagree on whether two particular species belong to the same genus or to two different genera (see chapter 7). I've tried to indicate some of these differences of opinion in this list, which is based on my interpretations of the best classifications.
For each genus, I list the name and what it means.
I also give the dinosaurs age: both the geologic epoch it comes from and approximately when (in millions of years ago) it lived. Unfortunately, our understanding of how old a dinosaur was is only as good as our understanding of the age of the rocks it was found in. The ages of some rocks are pretty well known: for these dinosaurs, we have very narrow time ranges. For others, though, the geologic ages are much less certain, so I list much longer possible ages for the oldest and youngest that dinosaur might be. Future studies should narrow those ranges down. In the 2015 updates the range of some of these ages have become very narrow. This is because some long on-going projects to precisely plot where each fossil specimen was found in certain geologic units--and when in time those particular layers were deposited--have finally been published. This much greater precision is giving us a better idea of which dinosaurs lived at the same time as each other. It also suggests that MOST dinosaur genera probably lasted for only a million years or so; if we knew the geology better, and had more samples to examine, we could probably narrow down the age ranges of most of the dinosaurs on this list to just a million years.
I give the length for these dinosaurs, based on the largest specimens. (Of course, for dinosaurs that are only known from babies, those "largest specimens" are a LOT smaller than the adult would be!) Since most dinosaurs are known from incomplete fossils, these measurements are often just guesses. Particularly wild guesses are marked with a question mark. (After all, since many dinosaurs are mostly tail and neck, and since tails and necks vary widely in some groups, it is pretty tough to make even a reasonable guess.) And since some dinosaurs are known from just a few tail bones or the like, there is no way to be accurate for these lengths. In these cases, I have just put a question mark. Keep these facts in mind. Also, keep in mind that the largest individuals may not be a typical individual. After all, consider really tall basketball players or really massive (American) football linesmen: they show that there are people who are much larger than the average individual, but still part of the normal variation of our species. The same was true of dinosaur species: not all Tyrannosaurus rex adult males would have been exactly the same size!
Keep in mind also that many dinosaurs are known from only a few individuals, so are unlikely to be large individuals. And even more importantly, they might not even be adults! In fact, even in some of the best known dinosaurs (Allosaurus and Apatosaurus are two good examples) we do not have definite adult individuals known, so the actual largest size of these genera is not yet known.
Weights are even tougher to determine. A baby dinosaur of just a few pounds could grow up to be a dinosaur weighing dozens of tons. So where I can, I give the weight of the biggest individuals. Instead of given exact numbers (which sound pretty accurate, but are really just guesses), I list a modern animal of around the same size as that dinosaur. Here is the list of modern animals, the weight they represent, and other modern animals in that size range:
Modern Animal Weight Range Other Modern Examples Sparrow Less than 2 oz (57 g) House Mouse; Finch Pigeon 2-16 oz (58-453 g) Blue Jay; Robin; Rat Chicken 1-5 lbs (0.45-2.27 kg) Crow; Hawk; Seagull Turkey 5-20 lbs (2.27-9.1 kg) House Cat; Goose; Raccoon Beaver 20-50 lbs (9.1-22.7 kg) Lynx; Jackal Wolf 50-100 lbs (22.7-45 kg) Baboon; Goat Sheep 100-200 lbs (45-91 kg) Leopard Lion 200-500 lbs (91-227 kg) Tiger Grizzly Bear 500-1000 lbs (227-454 kg) Zebra Horse 1000-2000 lbs (454-907 kg) Bison; Kodiak Bear Rhino 1-4 tons (0.9-3.6 t) Hippo; Giraffe Elephant 4-8 tons (3.6-7.2 t) Killer Whale
For dinosaurs of 8-16 tons, I've listed them as "two elephants"; for 16-24 tons, "three elephants"; and so on. For comparison's sake, the largest blue whale ever recorded (the largest animal known) was 209 tons and would be listed as "27 elephants" according to this scale.
As with length, though, there are plenty of dinosaurs that are known only from very fragmentary fossils. For ones where I could guess the weight even approximately, I've indicated that with a question mark next to the weight; for those that are just too hard to figure out, I've just put a question mark.
Estimating the masses of dinosaurs and other animals from fossils is exceedingly difficult. There are many complicating factors: how much fat they had (was it a good feeding season or a bad one?); how much muscle they had; etc.; etc. Different paleontologists have used different techniques. Some use laser scans of nearly complete skeletons, then create computer models based on the skeletons to get the volume. Others use measurements of individual bones, and used equations based on the size of more complete skeletons to estimate the size of bipedal and quadrupedal dinosaurs. But these estimates are just that--estimates--and so it is hard for us at present to figure out how accurate they are (that is, how close the calculations are to the actual masses.)
Also listed is the place where each dinosaur has been discovered. Of course they lived other places, too. In fact, you can pretty much guarantee that if a dinosaur species is known from fossils in (for example) Montana in the northern part of the western U.S., and New Mexico in the southern part of the western U.S., it almost certainly lived in the states in between. We just haven't found the fossils of it from there yet.
Finally, I mention some additional comments or fun facts about each dinosaur genus.
Here is the most recently finished Updated Genus List (18 January 2015).
I really want to give my thanks to Mr. Fred Barmwater of Highlands Ranch, Colorado. He helped in compiling the old data in a way that was easier to update. If you happen to see Mr. Barmwater while at the Denver Museum of Nature and Science, or anywhere, please thank him for helping me out! Additional proof-reading help has been provided by "Albertonykus", Pete Bucholz, Barbara Peterson, Christian Schley, Adam "Oxalaia" Schmoetzer, and Hans-Dieter Sues.
because dinosaurs have so many fans, the dinosaur pages on Wikipedia are often very up-to-date. (But make sure to check out the reference sources for any Wikipedia page, to see if they are using the latest information.)
BACK to main page.
Last modified 17 December 2019
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Skull of Cedarpelta bilbeyhallorum, on display at the USU Eastern Prehistoric Museum, Price, Utah. Scientific classification Domain: Eukaryota Kingdom: Animalia Phylum: Chordata Clade: Dinosauria Clade: †Ornithischia Clade: †Thyreophora Clade: †Ankylosauria Family: †Ankylosauridae Genus: †Cedarpelta
Carpenter et al., 2001 Species:
†C. bilbeyhallorum
Binomial name †Cedarpelta bilbeyhallorum
Carpenter et al., 2001
Cedarpelta is an extinct genus of basal ankylosaurid dinosaur from Utah that lived during the Late Cretaceous period (Cenomanian to lower Turonian stage, 98.2 to 93 Ma) in what is now the Mussentuchit Member of the Cedar Mountain Formation. The type and only species, Cedarpelta bilbeyhallorum, is known from multiple specimens including partial skulls and postcranial material. It was named in 2001 by Kenneth Carpenter, James Kirkland, Don Burge, and John Bird. Cedarpelta has an estimated length of 7 metres (23 feet) and weight of 5 tonnes (11,023 lbs). The skull of Cedarpelta lacks extensive cranial ornamentation and is one of the only known ankylosaurs with individual skull bones that are not completely fused together.
Discovery and naming
[edit]
The partial remains of an ankylosaur were discovered by Evan Hall and Sue Ann Bilbey at the CEM site near the Price River in Carbon County, Utah while they were visiting an excavation in the surrounding area.[1] The site was originally interpreted as being from the top of the Ruby Ranch Member of the Cedar Mountain Formation,[1] but was later interpreted as being from the bottom of the Mussentuchit Member.[2] The age of the layer was originally thought to have been 104.46 ± 0.95 Ma,[3] but more recent estimates date it to 98.2 ± 0.6 to 93 Ma.[4] In 1998, the discovery was reported by Kenneth Carpenter and James Kirkland.[5] In 2001, it was subsequently described, along with other material, by Kenneth Carpenter, James Kirkland, Don Burge, and John Bird. The holotype specimen, CEUM 12360, consists of a partial skull that is missing the snout and lower jaws. Numerous osteoderms, postcranial material and a disarticulated skull were designated as paratype specimens. Both holotype and paratype specimens represent at least three individuals and are currently housed at the College of Eastern Utah, Prehistoric Museum, Utah.[5][1]
The generic name, Cedarpelta, is derived from the Cedar Mountain Formation and the Greek word "pelte" (small shield). The specific name, bilbeyhallorum, honours Sue Ann Bilbey and Evan Hall, who discovered the remains of Cedarpelta.[1]
In 2008, additional specimens were referred to Cedarpelta from the Price River II Quarry, which is about 24.5 km southeast of Price River, Utah and at the base of the Mussentuchit Member. The quarry also produced specimens pertaining to four individuals of a brachiosaurid, an iguanodontian, a turtle, a pterosaur, and specimens of the nodosaurid Peloroplites. The referred material includes: CEUM 10396, a cervical vertebra; CEUM 10412, CEUM 10404, caudal vertebrae; CEUM 10371, a coracoid; CEUM 10256, CEUM 11629, humeri; CEUM 10266, an ischium; CEUM 11334, a femur; and CEUM 11640, a tibia.[2]
Description
[edit]
Carepnter et al. (2001) originally gave Cedarpelta an estimated length of 7.5-8.5 metres (24.6-27.9 feet). However, Gregory S. Paul gave a lower estimate of 7 metres (23 feet) and a weight of 5 tonnes (11,023 lbs), while Thomas Holtz gave a higher estimation at 9 meters suggesting that it was rivalling Ankylosaurus.[6][7][8]
Carpenter et al. (2001) established several distinguishing traits of Cedarpelta. The body of the praemaxilla, the front snout bone, is short in front of its nasal branch. The outer sides of the two praemaxillae run more parallel compared to the snouts of later forms which are strongly diverging to behind. The cutting edge of the bone core of the upper beak is limited to the front of the praemaxilla. Each praemaxilla has six (conical) teeth. The quadrate, and with it the entire back of the skull, is inclined to the front. The head of the quadrate is not fused with the paroccipital process, contrary to the situation in Shamosaurus. The neck of the occipital condyle is long and sticking out to behind, like with nodosaurids, not obliquely to below as in typical ankylosaurids. The tubera basilaria, appending processes of the rear lower braincase, form a large wedge directed to below. The pterygoid is elongated from the front to the rear and has a saddle-shaped process on its outer edge oriented to behind and sideways. The coronoid process of the rear lower jaw has an oval process at the inside. The straight ischium has a knob-shaped boss at the inside near the pubic pedicle.[1]
Cedarpelta shows a mix of basal and derived traits. The presence of premaxillary teeth is a plesiomorphic character because it is inherited from earlier Ornithischia. In contrast, closure of the opening on the side of the skull behind the orbit, the lateral temporal fenestra, is an advanced, derived (apomorphic) character only known in ankylosaurid ankylosaurians.[1]
Two skulls are known, and the skull length for Cedarpelta is estimated to have been roughly 60 centimetres (24 in). One of the Cedarpelta skulls was found disarticulated, a first for an ankylosaur skull, allowing paleontologists a unique opportunity to examine the individual bones instead of being limited to an ossified unit. The skull is relatively elongated and does not show a strongly appending beak. Of the conical premaxillary teeth, the first is the largest. The maxilla bears eighteen teeth. The eye socket is surrounded by the lacrimal, a single supraorbital and a large postorbital, excluding the prefrontal and the jugal from the orbital rim. The postcranial skeleton was in 2001 not described in any detail.[1]
The skulls, though of large and thus not juvenile individuals, do not show a distinctive pattern of fused caputegulae, head tiles. This inspired Carpenter to propose an alternative hypothesis of ankylosaur skull osteoderm formation. Formerly, it had been assumed that such armour plates were either formed by direct skin ossification into distinct scutes which later fused to the skull (the more popular theory), or by a reaction of the skull bones to the pattern of overlying scales. The lack of a clear pattern in Cedarpelta suggested to Carpenter that the ossification took place in an intermediate layer between the scales and the skull roof itself, which he surmised to have been the periosteum.[1]
Classification
[edit]
Carpenter (2001) placed Cedarpelta within the family Ankylosauridae and offered two interpretations of its position. The first was that it could be the basalmost known ankylosaurid, i.e. the first discovered branch to split off from the ankylosaurid stem line. This would be in line with its plesiomorphic traits and the fact that the in 2001 supposed Barremian age made it one of the oldest known ankylosaurids. The second was that it formed an early ankylosaurid branch, or clade, Shamosaurinae together with Gobisaurus of north-central China and the eponymous Shamosaurus of Mongolia.[9] Thompson et al. (2012),[10] Chen et al. (2013),[11] Yang et al. (2013),[12] Han et al. (2014),[13] Arbour & Currie (2015),[14] Arbour et al. (2016),[15] Arbour & Evans (2017),[16] Yang et al. (2017),[17] Zheng et al. (2018),[18] Rivera-Sylva et al. (2018),[19] Park et al. (2019)[20] and Frauenfelder et al. (2022)[21] have all found Cedarpelta to be within Ankylosauridae, as either within a polytomy with Liaoningosaurus, Aletopelta, Chuanqilong, Gobisaurus and Shamosaurus or as sister taxon to Chuanqilong. The results of Arbour & Currie (2015) are reproduced below.
Vickaryous et al. (2004) interpreted Cedarpelta as the basalmost member of the family Nodosauridae, positioned even below the nodosaurids Pawpawsaurus, Silvisaurus, and Sauropelta.[22] Wiersma & Irmis (2018) also interpreted Cedarpelta as a nodosaurid.[23] The results of Vickaryous et al. (2004) are reproduced below.
See also
[edit]
Dinosaurs portal
Timeline of ankylosaur research
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Nodosauridae is a family of ankylosaurian dinosaurs, from the Late Jurassic to the Late Cretaceous Period of what are now North America, Europe, Asia, and Antarctica.
Characteristics
Diagnostic characteristics for the Nodosauridae include supraorbital boss rounded protuberance, occipital condyle derived from only the basioccipital, and ornamentation present on the premaxilla. There is a fourth ambiguous characteristic called the acromion, which is a knob-like process. All nodosaurids, like other ankylosaurs, may be described as medium-sized to large, heavily built quadrupedal herbivorous dinosaurs, possessing small denticulate teeth and parasagittal rows of osteoderms (a type of armour) on the dorsolateral surfaces of the body.
Classification
The family Nodosauridae was erected by Othniel Charles Marsh in 1890, and anchored on the genus Nodosaurus.[1][2]
The clade Nodosauridae was first defined by Paul Sereno in 1998 as "all ankylosaurs closer to Panoplosaurus than to Ankylosaurus," a definition followed by Vickaryous, Maryanska, and Weishampel in 2004. Vickaryous et al. considered two genera of nodosaurids to be of uncertain placement (incertae sedis): Struthiosaurus and Animantarx, and considered the most primitive member of the Nodosauridae to be Cedarpelta.[3] The cladogram below follows the most resolved topology from a 2011 analysis by paleontologists Richard S. Thompson, Jolyon C. Parish, Susannah C. R. Maidment and Paul M. Barrett.[4] The placement of Polacanthinae follows its original definition by Kenneth Carpenter in 2001.[5]
Nodosauridae
Antarctopelta
Mymoorapelta
Hylaeosaurus
Anoplosaurus
Tatankacephalus
Horshamosaurus
Polacanthinae
Gargoyleosaurus
Hoplitosaurus
Gastonia
Peloroplites
Polacanthus
Struthiosaurus
Zhejiangosaurus
Hungarosaurus
Animantarx
Niobrarasaurus
Nodosaurus
Pawpawsaurus
Sauropelta
Silvisaurus
Stegopelta
Texasetes
Edmontonia
Panoplosaurus
Timeline
Biogeography
The near simultaneous appearance of nodosaurids in both North America and Europe is worthy of consideration. Europelta is the oldest nodosaurid from Europe, it is derived from the lower Albian Escucha Formation. The oldest western North American nodosaurid is Sauropelta, from the lower Albian Little Sheep Mudstone Member of the Cloverly Formation, at an age of 108.5±0.2 million years. Eastern North American fossils seem older. Teeth of Priconodon crassus from the Arundel Clay of the Potomac Group of Maryland, which dates near the AptianâAlbian boundary. The Propanoplosaurus hatchling from the base of the underlying Patuxent Formation, dating to the upper Aptian, is the oldest known nodosaurid.[1]
Polacanthids are known from pre-Aptian fauna from both Europe and North America. The timing of the appearance of nodosaurids on both continents indicates that the origins of the clade preceded the isolation of North America and Europe, pushing the group's date of evolution back to at least the "middle" Aptian. The separation of Nodosauridae into European Struthiosaurinae and North American Nodosaurinae by the end of the Aptian provides a revised date for the isolation of the continents from each other by rising sealevels.[1]
Below is a table showing the age difference between continents. North American nodosaurids are teal, European nodosaurids are green, European polacanthids are blue, and North American polacanthids are brown. Other nodosaurids or polacanthids are black. This table supports the observations by Kirkland et al. (2013).[1]
James Kirkland et al. considers Mymoorapelta, Gargoyleosaurus, Hylaeosaurus, Polacanthus, Hoplitosaurus and Gastonia to be Polacanthids, outside of Nodosauridae.[1]
See also
Timeline of ankylosaur research
References
Carpenter, K. (2001). "Phylogenetic analysis of the Ankylosauria." In Carpenter, K., (ed.) 2001: The Armored Dinosaurs. Indiana University Press, Bloomington & Indianapolis, 2001, pp. xv-526
Osi, Attila (2005). Hungarosaurus tormai, a new ankylosaur (Dinosauria) from the Upper Cretaceous of Hungary. Journal of Vertebrate Paleontology 25(2):370-383, June 2003.
Wikimedia Commons has media related to Nodosauridae.
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Alvarezsaurs in the Lori matrix
This time our topology is-
I provide a new definition for Alvarezsauroidea that adds Therizinosaurus as an external specifier since I find it most parsimonious for Therizinosauria to be its sister group, and uses Alvarezsaurus as the internal specifier unlike Sereno's that uses Shuvuuia- (Alvarezsaurus calvoi< - Ornithomimus velox, Therizinosaurus cheloniformis, Passer domesticus) . Alvarezsauroids have had a controversial phylogenetic placement, with the Lori matrix recovering them as basal maniraptorans sister to therizinosaurs. Yet they can be outside therizinosaurs plus pennaraptorans in 3 steps, become avemetatarsalians in 4 steps (can bring therizinosaurs or not), non-maniraptoriforms in 6 steps (they bring therizinosaurs), closer to pennaraptorans than therizinosaurs in 6 steps, paravians in 11 steps (therizinosaurs move with), closer to Compsognathus than to birds in 15 steps, closer to birds than deinonychosaurs in 27 steps, and closer to Archaeopteryx and other birds than to dromaeosaurids and troodontids in 30 steps.
Fukuivenator is an odd taxon, recovered here as the basalmost alvarezsauroid. But it can be a therizinosaurian in only two steps, and outside Maniraptoriformes in 4 steps (it emerges in Coeluridae). One thing I don't think it is is a dromaeosaurid, as that takes 27 more steps, and getting it into Paraves or Pennaraptora requires 11 and 7 steps respectively. Still, I wouldn't be surprised to see this taxon work its way around the base of Maniraptoriformes once an osteology comes out.
Shuvuuia deserti IGM 100/975 axial elements in ventral view and pelvis in dorsal view (courtesy AMNH).
Nqwebasaurus was recently redescribed by Sereno (2017), which I incorporated into its scorings. Choiniere et al. (2012) recovered it in Ornithomimosauria, but note most of the characters they list to support that are also said to be present in alvarezsauroids. Even they could place it in Alvarezsauroidea with only 4 steps. The Lori matrix needs 6 steps to place it in Ornithomimosauria, which I think is higher partially due to it finding Pelecanimimus to be an alvarezsauroid too. So similarities between the two like their teeth being in a common groove and maxillary teeth being confined to the anterior third of the bone are no longer ornithomimosaur-like. As recently noted by Cerroni et al. (2019), this makes more sense biogeographically as well. Oh, and note that the Lori matrix found Afromimus to be a ceratosaur as in that paper. In any case, Nqwebasaurus takes 10 steps to move to Compsognathidae, and 7 steps to move sister to Pennaraptora.
As for Pelecanimimus itself, it seems plausibly alvarezsauroid if you think about it. The skull is famously similar to Shuvuuia, the posterior tympanic recess is in the otic recess, ossified sterna are otherwise unknown for ornithomimosaurs, the long manual digit I was always out of place compared to Harpymimus, and Europe makes more sense for otherwise Gondwanan clades in the Cretaceous. Now if only someone would release Perez-Moreno's thesis describing it in detail...
Shuvuuia deserti IGM 100/975 pectoral and forelimb elements. Note the tiny phalanx from digit II or III at the bottom (courtesy AMNH).
Patagonykus and Bonapartenykus are usually closer to parvicursorines than Alvarezsaurus and Achillesaurus, but the Lori matrix found them just outside Alvarezsauridae instead. Interestingly, Xu et al. (2018) recovered the same results. It takes 3 steps to move Patagonykus closer to parvicursorines, and 4 steps to join Alvarezsaurus and Patagonykus to the exclusion of parvicursorines as in Alifanov and Barsbold (2009). Xu et al. recover these in 5 and 7 steps respectively, and the most recent version of Longrich and Currie's alvarezsaurid matrix (Lu et al., 2018) recovers a basal Patagonykus and a basal Parvicursorinae in 3 steps each.
One odd result is that the newly described Xiyunykus and Bannykus fall in Patagonykinae too. Yet only 2 steps move them outside the Patagonykus plus Parvicursorinae clade, where they form a clade. Another step breaks that up to place Xiyunykus more basal as in Xu et al.. Them being basal certainly fits better stratigraphically, and Xu et al. use several characters designed for alvarezsauroids that the Lori matrix didn't include yet. Hopefully full osteologies will be published as well.
Mononykus olecranus cast YPM 56693 (of holotype) pes in plantar view (courtesy of Senter).
A patagonykine Achillesaurus as suggested by Agnolin et al. (2012) takes 7 additional steps in the Lori matrix where it instead emerges just closer to parvicursorines than Alvarezsaurus. On the other hand, only a single step joins it with Alvarezsaurus as in Longrich and Currie (2009) and only 2 steps makes it just further from parvicursorines than Alvarezsaurus as in Xu et al. (2018).
Alnashetri is known from type hindlimb material, but now also from MPCA 377, a nearly complete specimen with interesting characters like flat and unfused sternal plates. Makovicky et al. (2016) used this data to recover it as the sister group to Alvarezsauridae, and while the few published details left it more derived in the Lori tree, it can go to a more basal position with only two steps. It should be interesting to compare to e.g. Bannykus once it is published.
Mononykus olecranus cast YPM 56693 (of holotype) (courtesy of Senter).
The arctometatarsal clade has a unique topology, but no other analysis has included nearly as many characters or all of these taxa, with Lu et al. omitting Albinykus and Ceratonykus among non-fragmentary specimens, and Xu et al. omitting the more recently described Qiupanykus. Enforcing the Lori topology in Lu et al.'s matrix is only 5 steps longer, and doing so in Xu et al.'s matrix is only 6 steps longer. On the other hand, Xu et al.'s topology is so unresolved at this level, the only difference in mine is placing the Albinykus plus Xixianykus clade basally near Albertonykus, which takes 5 steps to do in the Lori matrix.
It should be noted that Lu et al.'s illustrated topology (their Figure 3) is not their matrix's real result, as they did not fully analyze tree space. Instead of 20 trees, there are 214 trees. These differ in that Albertonykus, YPM 1049 and undescribed 41HIII-0104 can fall out anywhere more derived than Patagonykus, and that Parvicursor, the Tugriken Shireh taxon, Shuvuuia and Mononykus form an unresolved polytomy. This leaves Linhenykus, Qiupanykus and Xixianykus unresolved between that polytomy and Patagonykus, which is perfectly compatible with the Lori topology. This may also show that the small alvarezsauroid-specific matrix of Longrich and Currie is insufficient given all the new taxa described since 2009. YPM 1049 was far too fragmentary to include (distal metatarsal III) but I tried testing undescribed Quipa specimen 41HIII-0104. Didn't make it into the publication, but here's its scorings-
'41HIII0104' ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ???????-?? ?????????1 ?10??????? ?????????? ?????????? ???0?????? ????1????? ???1?????{01} ?????????? ?????????? ?????????? ????????0? ????????3? ?????????? ?????????? ?????????? ?????????? ?????????1 ?????????1 1????????? 1????????? ?????????? ?????????? ?????????? ???1?????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ????????0? ?????????? ?????????? ?????????? ?????????? ?????????? ???1?????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????{123}???? ?????????? ?????????? ?????????? ?????????? ?????????? ??1???0???
Tugriken Shireh parvicursorine (IGM 100/99) vertebrae and ilia in ventral view, forelimb and fibula in lower right (courtesy AMNH).
Interestingly, Agnolin et al., Xu et al. and the Lori analysis all recovered Albinykus sister to Xixianykus outside Parvicursorinae. Wonder if that's a real signal? Unfortunately, the only attempt to name this clade was Agnolin et al. who also recovered Ceratonykus in there and called it Ceratonykini. Xu et al. place Ceratonykus closer to parvicursorines, while I found it more basal than either, sister to Qiupanykus which neither of the other studies used. Forcing Ceratonykus sister to Albonykus plus Xixianykus takes 3 more steps in the Lori matrix. Forcing Ceratonykus sister to Mononykus as in its original description (with or without Qiupanykus) takes 5 more steps. As stated in the paper, we were the first analysis to include Hateg tibiotarsi Bradycneme and Heptasteornis. While the former can fall into many positions in Maniraptora, the latter was resolved as an alvarezsaurid as proposed by Naish and Dyke (2004). Note this used only the tibiotarsus and not alvarezsaurid-like distal femur FGGUB R.1957. A single step moves Heptasteornis to Troodontidae.
We also provide an updated definition for Parvicursorinae (Mononykus olecranus + Parvicursor remotus), like Choiniere et al.'s (2010) but using species. One accident of our definitional and discovery history is that all these newer arctometatarsal alvarezsaurids (Xixianykus, Albertonykus, Albinykus, Linhenykus, Qiupanykus, Ceratonykus, etc.) emerge outside the originally discovered and defined Parvicursorinae. We could really use some clade defining taxa closer to Mononykus than Patagonykus, Alvarezsaurus or Achillesaurus. In any case, I got a lot of experience with parvicursorine specimens, examining Shuvuuia and the Tugriken Shireh specimen IGM 100/99 in person, and having photos of high quality casts of Mononykus thanks to Senter. I found the Tugriken Shireh taxon closer to Shuvuuia, but moving it closer to Parvicursor as in Longrich and Currie is just 1 step longer.
Tugriken Shireh parvicursorine (IGM 100/99) vertebrae and ilia in dorsal view, forelimb and fibula in lower right (courtesy AMNH).
Next time, therizinosaurs...
References- Naish and Dyke, 2004. Heptasteornis was no ornithomimid, troodontid, dromaeosaurid or owl: The first alvarezsaurid (Dinosauria: Theropoda) from Europe. Neus Jahrbuch für Geologie und Paläontologie. 7, 385-401.
Alifanov and Barsbold, 2009. Ceratonykus oculatus gen. et sp. nov., a new dinosaur (?Theropoda, Alvarezsauria) from the Late Cretaceous of Mongolia. Paleontological Journal. 43(1), 94-106.
Longrich and Currie, 2009. Albertonykus borealis, a new alvarezsaur (Dinosauria: Theropoda) from the Early Maastrichtian of Alberta, Canada: Implications for the systematics and ecology of the Alvarezsauridae. Cretaceous Research. 30(1), 239-252.
Choiniere, Xu, Clark, Forster, Guo and Han, 2010. A basal alvarezsauroid theropod from the early Late Jurassic of Xinjiang, China. Science. 327, 571-574.
Agnolin, Powell, Novas and Kundrat, 2012. New alvarezsaurid (Dinosauria, Theropoda) from uppermost Cretaceous of north-western Patagonia with associated eggs. Cretaceous Research. 35, 33-56.
Makovicky, Apesteguia and Gianechini, 2016. A new, almost complete specimen of Alnashetri cerropoliciensis(Dinosauria: Theropoda) impacts our understanding of alvarezsauroid evolution. XXX Jornadas Argentinas de Paleontologia de Vertebrados. Libro de resumenes, 74.
Sereno, 2017. Early Cretaceous ornithomimosaurs (Dinosauria: Coelurosauria) from Africa. Ameghiniana. 54, 576-616.
Lu, Xu, Chang, Jia, Zhang, Gao, Zhang, Zhang and Ding, 2018. A new alvarezsaurid dinosaur from the Late Cretaceous Qiupa Formation of Luanchuan, Henan Province, central China. China Geology. 1, 28-35.
Xu, Choiniere, Tan, Benson, Clark, Sullivan, Zhao, Han, Ma, He, Wang, Xing and Tan, 2018. Two Early Cretaceous fossils document transitional stages in alvarezsaurian dinosaur evolution. Current Biology. 28, 1-8. DOI: 10.1016/j.cub.2018.07.057
Cerroni, Agnolin, Egli and Novas, 2019. The phylogenetic position of Afromimus tenerensis Sereno, 2017 and its paleobiogeographical implications. Journal of African Earth Sciences. DOI: 10.1016/j.jafrearsci.2019.103572
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
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Therizinosaurs in the Lori matrix
Next up are therizinosaurs. These are one of the best analyzed clades because I incorporated all of Zanno's (2010) characters, which is by far the largest and most recent analysis of the group until the Lori paper was published. The topology is-
Falcarius is the most basal taxon shown of course, but Martharaptor was pruned a posteriori and can fall out anywhere in Therizinosauria outside the Alxasaurus plus Segnosaurus clade. I tried including Thecocoelurus, but the Lori matrix is pretty terrible when it comes to scoring single vertebrae-
Thecocoelurus ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ???????(01)0? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ???1?????? ?????????? ?????2???? ?????????? ?????????? ?????????? ?????????? ???????0?? ?????????? ?????????? ?????????? ?????1???? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ??0??????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ??????????
Jianchangosaurus fell out in the same place as its original description, with Cau's (2018) placement in Alvarezsauroidea taking 18 more steps so is very unlikely. Mine is the only published matrix besides Senter (2011) and its derivatives to use information from the second Beipaiosaurus specimen, and incorporated photos from Zanno and the new paper on the holotype skull elements too.
Beipiaosaurus inexpectus holotype (IVPP V11559) cervical vertebra in dorsal view (courtesy of Zanno).
Zanno also provided photos of Alxasaurus and Enigmosaurus, and its depressing how much of the former is lost. Enigmosaurus rather famously was shown by Zanno to not resemble Barsbold's original illustration that was the only reference picture known for over two decades. Placing Enigmosaurus closer to Segnosaurus than Neimongosaurus or Erliansaurus as in Zanno's tree takes 4 more steps. Forcing Enigmosaurus and Erlikosaurus to be sister taxa to simulate the synonymy mentioned by Barsbold (1983) takes 4 steps, so seems unlikely. The duo moves between Nanshiungosaurus and the Segnosaurus plus Nothronychus clade. We were the first analysis to include "Chilantaisaurus" zheziangensis, which emerged in a polytomy with Alxasaurus, Enigmosaurus and therizinosaurids.
Alxasaurus elesitaiensis holotype (IVPP V88402a) chevrons in right lateral view (natural order reversed) (courtesy of Zanno).
As was the case with Archaeornithomimus? bissektensis, we didn't include the possible chimaera of Bissekty Therizinosauria as an OTU, unlike Sues and Averianov (2015). But if you do want to experiment with it, here's the scorings. It emerges in a polytomy in the Suzhousaurus plus Therizinosaurus clade of therizinosaurids. Btw, Archaeornithomimus? bissektensis does fall out most parsimoniously sister to A. asiaticus when all Bissekty material is used.
'Bissekty-Therizinosauroidea' ????1??0?? ????1????? ?????1???? ?1???????? ?????{01}1000 ?01??????? 00???????? 2??21?0??? ???????00{123} {12}00(01)2011?? ??0???0(01)00 ??11????{12}? ?{01}0?100??? ?????????? ?1{01}?0????? ??0?0???{012}0 000(01)?010?? ?????????? ?????????? ?????0{12}?00 1?0???0??0 ??0??{01}0??? 0?1??????? ????0?1?0? {01}0?1???{01}?? ??0??????? ?????????? ?????????0 ???001??00 ??00?01?1? 10???????? ?????????1 ???0?????? ???1?????? ??????0??1 0???1(12)???? ????0???1? ????0?011? ???0????0? ???0{12}0???? ????????0? 0?0????1?? ?????????? ??010101?1 ??0?0(01)???? ?????{01}010? ???100???? ????????11 1010?????? ??0??0???? ?????????? ?????????? ??????0??? ??00?????? ?00??????? ?????????? ???????0-- -??00??010 ?????????? ??????-??? ???-010??1 0100????00 10?1?00??? ???0?00??? ?????????? ????0????? ???000???? ?0???-???? ?????????? ?????001??
Enigmosaurus mongoliensis holotype (IGM 100/84) synsacrum and ilium in ventral view (courtesy of Zanno).
Next is Therizinosauridae itself, which we refined Zhang et al.'s (2001) definition of to include type species. Therizinosaurids first split into a clade of Erliansaurus, Neimongosaurus, Suzhousaurus and Therizinosaurus. Forcing the former two to be outside a clade of Suzhousaurus, Therizinosaurus and the taxa below, as in Zanno's tree, takes 5 more steps. Notably, we did not include the hindlimb IGM 100/45 in the Therizinosaurus OTU since there's no overlap and its not even particularly large. But here's the Therizinosaurus OTU including the hindlimb. Using this version of Therizinosaurus leaves the tree basically the same but destabilizes it somewhat in that Therizinosaurus and Erlikosaurus can now go in multiple positions within Therizinosauridae, and the Nanchao embryos are in a trichotomy with the Suzhousaurus clade and the Nothronychus clade. Is this an indication the hindlimb produces homoplasy and so might not belong to Therizinosaurus?
Therizinosaurus ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ????????01 0110021000 01?0001100 00001110?? ?????????? ?????????? ?????????? ???0???010 1?0000000? 0?1??1???? ?????????? ?????????? 000??0???? ??????3??1 ?????????0 ???00????? 0?1?0?1??? ?????0???? ?????????1 ?????????? ?????0???? ?????????? ?????????? 1011010101 100100??1? ????{12}?00?? 00?12110?? 101??????? 0????0?111 00?010???? ????????1? ?????????? ????100111 1101110??? ?????????? ????????1? ?1???????? ?????????1 ???0?????? ?000?????? ?????0???1 ?????????? ?????????? ?????????? ?????????? ?????0???? ??????-00? ?00-000001 0?0100??0? 10000???0{01} ?{01}?0000??0 {01}0???????? ????????00 ?00?????0- -??11-???? ?????????? ???1?0???0
Segnosaurus galbinensis paratype (IGM 100/83) cervical neural arch in right lateral view (courtesy of Zanno).
Now comes the Nanchao therizinosaur embryos, those described by Kundrat et al. inside dendroolithid eggs. While including such young specimens might be seen as risky, my ontogenetically conservative scoring method with state N seems to have worked fine here. They fall out where you'd expect a Santonian-Campanian therizinosaur to do so. Following that is Nanshiungosaurus brevispinus, which Senter et al. (2012) recovered as the next most derived therizinosaur after Alxasaurus. Forcing it into this basal position takes 4 steps. Nanshiungosaurus? bohlini was included but pruned a posteriori since it can go anywhere in the Segnosaurus plus Nothronychus clade. Forcing Nanshiungosaurus monophyly is just a single step longer though, while forcing bohlini to be sister to the contemporaneous Suzhousaurus takes 2 steps.
Segnosaurus itself (which Zanno also provided photos of) pairs with ex-Alectrosaurus forelimb AMNH 6368, which has only previously been analyzed by Zanno (2006) where it pairs with Erliansaurus. Forcing that here compared to other taxa she included results in trees 3 steps longer. Erlikosaurus groups with the Nothronychus species in a trichotomy where it can be sister to either species. Forcing Nothronychus monophyly takes only a single step, but note that no proposed Nothronychus characters involve elements that can be compared to Erlikosaurus (humerus and pes). Forcing Erlikosaurus to group with Therizinosaurus as in Senter et al. requires only a single step, with Erlikosaurus moving to the Therizinosaurus clade.
Next time, oviraptorosaurs...
References- Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Transactions of the Joint Soviet-Mongolian Palaeontological Expedition. 19, 117 pp.
Zhang, Xu, Sereno, Kwang and Tan, 2001. A long-necked therizinosauroid dinosaur from the Upper Cretaceous Iren Dabasu Formation of Nei Mongol, People’s Republic of China. Vertebrata PalAsiatica. 39(4), 282-290.
Zanno, 2006. The pectoral girle and forelimb of the primitive therizinosauroid Falcarius utahensis (Theropoda, Maniraptora): Analyzing evolutionary trends within Therizinosauroidea. Journal of Vertebrate Paleontology. 26(3), 636-650.
Zanno, 2010. A taxonomic and phylogenetic re-evaluation of Therizinosauria (Dinosauria: Maniraptora). Journal of Systematic Palaeontology. 8(4), 503-543.
Senter, 2011. Using creation science to demonstrate evolution 2: Morphological continuity within Dinosauria. Journal of Evolutionary Biology. 24(10), 2197-2216.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid tail. PLoS ONE. 7(5), e36790.
Sues and Averianov, 2015. Therizinosauroidea (Dinosauria: Theropoda) from the Upper Cretaceous of Uzbekistan. Cretaceous Research. 59, 155-178.
Cau, 2018. The assembly of the avian body plan: A 160-million-year long process. Bollettino della Società Paleontologica Italiana. 57(1), 1-25.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
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Happy New Year 2020
Hi all. A Theropod Database update is online, with the main additions being troodontid information and info from the Hayashibara Museum of Natural Sciences Research Bulletins 1-3. I love these publications and wish more like them existed for other collections. They detail the expeditions into Mongolia with exact discovery dates and field numbers for taxa like Nomingia, Elsornis and Aepyornithomimus, and tons of still undescribed specimens. It's amazing just how many ornithomimosaurs are known from the Bayanshiree Formation for instance, when only the Garudimimus holotype has been described. There are over twenty more including the sort-of-described "Gallimimus""mongoliensis" specimen IGM 100/14. So often for new taxa, especially those from the Jehol biota, no information is provided in the description as to when the specimen was discovered. I get that many are found by non-professionals and given to museums, but at least say "the specimen was given to the museum on x-x-xx by someone who said it was excavated around year y." Next up, halszkaraptorine and dromaeosaurid updates...
undescribed ?Gobivenator skull (HMNS coll.; field number 940801 TS-I WTB) (after Tsogtbaatar and Chinzorig, 2010).
Reference- Tsogtbaatar and Chinzorig, 2010. Fossil specimens prepared in Mongolian Paleontological Center: 2002–2008. Hayashibara Museum of Natural Sciences Research Bulletin. 3, 155-166.
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Details on Teinurosaurus and random musings
Hi all. When updating The Theropod Database I noticed my entry for Teinurosaurus is pathetically bad- wrong authors, wrong age, wrong size, and generally missing the complicated history of this innocuous vertebra. How embarrassing! So here's the revised version that will be uploaded-
Teinurosaurus Nopcsa, 1928
= Saurornithoides Nopsca, 1928 (preoccupied Osborn, 1924)
= Caudocoelus Huene, 1932
T. sauvagei (Huene, 1932) Olshevsky, 1978
= Caudocoelus sauvagei Huene, 1932
Tithonian, Late Jurassic
Mont-Lambert Formation, Hauts-de-France, France
Holotype- (BHN2R 240; = Boulogne Museum 500) incomplete distal caudal vertebra (75 mm)
Diagnosis- Provisionally indeterminate relative to Kaijiangosaurus, Tanycolagreus and Ornitholestes.
Other diagnoses- (after Huene, 1932; compared to Elaphrosaurus) centrum wider; narrower ventral surface; ventral median groove wider; transversely narrower prezygapophyses.
While Huene attmpted to distinguish Teinurosaurus from Elaphrosaurus, only the wider median ventral groove is apparent in existing photos of the former. This is compared to the one distal caudal of the latter figured in ventral view, but as Kobayashi reports grooves become distally narrower in Harpymimus while Ostrom reports they become distally wider in Deinonychus, groove width is not considered taxonomically distinctive at our current level of understanding. Indeed, this lack of data is most relevent to both diagnosing and identifying Teinurosaurus. Very few taxa have detailed descriptions of distal caudal vertebrae or more than lateral views figured, let alone indications of variation within the distal caudal series. So the facts that Fukuiraptor and Deinonychus share ventrally concave central articulations with Teinurosaurus in their single anteriorly/posteriorly figured distal caudal vertebra, or that Afromimus, "Grusimimus" and Falcariusalso have have wide ventral grooves in their few ventrally figured distal caudals, are not considered taxonomically important.
Comments- Sauvage (1897-1898; in a section written in January 1898) first mentioned a distal caudal vertebra he referred to the ornithischian Iguanodon prestwichii (now recognized as the basal styracosternan Cumnoria prestwichii) - "We are disposed to regard as belonging to the same species the caudal vertebra of a remote region, the part which we figure under n ° 7, 8" [translated]. Note Galton (1982) was incorrect in claiming Sauvage reported on this specimen in his 1897 paper (written December 6), which includes a section on prestwichiinearly identical to the 1897-1898 one but which lacks the paragraph describing this vertebra. This could provide a specific date of December 1897 to January 1898 for the discovery and/or recognition of the specimen. Huene (1932) correctly noted Sauvage mislabeled plate VII figure 8 as dorsal view, when it is in ventral view as understood by the text. Compared to Cumnoria, the caudal is more elongate (length 3.93 times posterior height compared to 2.54 times at most), has a ventral median groove instead of a keel, and the prezygapophyseal base in 71% of the anterior central height compared to ~30-40%, all typical of avepods. Nopcsa (1928) recognized its theropod nature and in his list of reptile genera meant to use a footnote to propose Teinurosaurusas a "new name for the piece described and figured by Sauvage (Direct. Traveaux Geol. Portugal Lisbonne 1897-1898, plate VII, Fig. 7-10) as late caudal of Iguanodon Prestwichi." Teinurosaurusis listed as an aublysodontine megalosaurid (not as an ornithomimine, contra Galton), roughly equivalent to modern Eutyrannosauria. However due to a typographical error, the footnote's superscript 1 was placed after Saurornithoides instead of Teinurosaurus. Sauvage (1929) corrected this in an addendum- "footnote 1 does not refer to Saurornithoides (line 19 from below) but to Teinurosaurus(last line of text)." Unfortunately, Huene missed the addendum, and thus wrote "Nopcsa recognized in 1927 (43, p. 183) that this was a coelurosaur and intended to give it a name, but used one already used by Osborn, namely "Saurornithoides" (91, 1924, p. 3- 7). For this reason, a new name had to be given here" [translated]. Huene's proposed new name was Caudocoelus sauvagei, placed in Coeluridae and "somewhat reminiscent of Elaphrosaurus." Huene is also perhaps the first of several authors to place the specimen in the Kimmeridgian, when it is actually from the Tithonian (Buffetaut and Martin, 1993; as Portlandian). Galton wrote "Lapparent and Lavocat (1955: 801) gave a line drawing of the vertebra after Sauavage (1898) and included it in the section on Elaphrosaurus" and that the specimen "was referred to Elaphrosaurusby Lapparent and Lavocat (1955)." This was perhaps done because Huene explicitly compared the two, ironically making it the only taxon distinguished from Teinurosaurus at the time. Most of Huene's characters cannot be checked in the few published photos of Teinurosaurus, but the ventral median sulcus is indeed much wider than Elaphrosaurus. Ostrom (1969) was the first author to detail Nopcsa's (1929) addendum, stating "Nopcsa's name Teinurosaurus has clear piority over Huene's Caudocoelus, but since Nopcsa failed to provbide a specific name, Teinurosaurus is not valid." Olshevsky (1978) solved this by writing "Teinurosaurus has clear priority over Caudocoelus, as noted in Ostrom 1969, and it is certainly a valid generic name. The species Caudocoelus sauvagei is proposed here as the type species of the genus Teinurosaurus, resulting in the new combination Teinurosaurus sauvagei(von Huene 1932) as the proper name of the type specimen." He also claimed "the specimen itself, unfortunately, was destroyed during World War II and thus must remain a nomen dubium." This was repeated by Galton, but as Buffetaut et al. (1991) wrote- "Contrary to a widespread opinion (expressed, for instance, by Lapparent, 1967), the vertebra in question has survived two world wars and years of neglect, like a large part of the other fossil reptile remains in the collections of the Boulogne Natural History Museum (see Vadet and Rose, 1986)." Olshevsky noted Steel misunderstood Nopsca in a different way, believing Teinurosaurus instead of Aublysodon was a "name, proposed by Cope in 1869 ... used instead of Deinodon", as stated under superscript 2. Galton did have the first modern opinion on Teinurosaurus' affinities, stating "In addition to Elaphrosaurus, elongate prezygapophyses occur in the allosaurid Allosaurus and the dromaeosaurid Deinonychus, so this caudal vertebra can only be identified as theropod, family incertae sedis." Buffetaut and Martin (1993) agreed, saying "no really distinctive characters that would allow a familial assignment can be observed." Ford (2005 online) gave the type repository as "Dortigen Museum", but this is a misunderstanding based on Huene's "Boulogne-sur-mer (Nr. 500 im dortigen Museum)", which roughly translated is "Boulogne-sur-mer region (No. 500 in the museum there)", referring to the Boulogne Museum where it has always been held. It was originally number 500, but was recatalogued at some point.
Sauvage lists the vertebra's length as 75 mm and his plate at natural size would have it be 79 mm, Huene lists it as 11 cm (110 mm) and his figure at 1:2 size would have it be 152 mm. Galton's drawing with supposed 5 cm scale would have it be 235 mm, while Buffetaut and Martin's plate with scale would leave it at 74 mm. As Huene's and Galton's figures are taken from Sauvage's original plate and the newest and unique photo matches Sauvage's reported length almost exactly, 75 mm is taken as the correct length.
Relationships- While prior authors haven't specified Teinurosaurus' relationships past Theropoda (besides Lapparent and Lavocat's apparent synonymy with Elaphrosaurus), there are several ways to narrow down its identity. Only neotheropods are known from the Late Jurassic onward, so coelophysoid-grade taxa are excluded. Some theropod clades were too small to have a 75 mm caudal, including most non-tyrannosauroid coelurosaurs besides ornithomimosaurs, therizinosaurs and eudromaeosaurs. The former two are unknown from the Jurassic, and additionally paravians like eudromaeosaurs lack any neural spine by the time the centrum gets as elongate as Teinurosaurus (e.g. by caudal 12 in Deinonychus at elongation index of 2.4). Teinurosaurus has an elongation index (centrum length/height) of 3.9, which also excludes Ceratosauridae, Beipiaosaurus+ therizinosauroids and oviraptorosaurs. Prezygapophyses basal depth is significantly less in ceratosaurids, megalosaurids, carnosaurs except Neovenator, compsognathids, Fukuivenator and Falcarius. Remaining taxa are elaphrosaur-grade ceratosaurs, piatnitzkysaurids, Neovenator and basal tyrannosauroids.
References- Sauvage, 1897. Notes sur les Reptiles Fossiles (1). Bulletin de la Société géologique de France. 3(25), 864-875.
Sauvage, 1897-1898. Vertebres Fossiles du Portugual, Contributions a l'etude des poissions et des reptiles du Jurassique et du Cretaceous. Direction des Travaux Geologiques Portugal. 1-46.
Osborn, 1924. Three new Theropoda, Protoceratops zone, central Mongolia. American Museum Novitates. 144, 1-12.
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Nopcsa, 1929. Addendum "The genera of reptiles". Palaeobiologica. 2, 201.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1), 361 pp.
Lapparent and Lavocat, 1955. Dinosauriens. In Piveteau (ed.). Traite de Paleontologie. Masson et Cie. 5, 785-962.
Lapparent, 1967. Les dinosaures de France. Sciences. 51, 4-19.
Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Peabody Museum of Natural History Bulletin. 30, 1-165.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia of Paleoherpetology. Gustav Fischer Verlag. 87 pp.
Olshevsky, 1978. The archosaurian taxa (excluding the Crocodylia). Mesozoic Meanderings. 1, 50 pp.
Galton, 1982. Elaphrosaurus, an ornithomimid dinosaur from the Upper Jurassic of North America and Africa. Paläontologische Zeitschrift. 56, 265-275.
Vadet and Rose, 1986. Catalogue commente des types et figures de dinosauriens, ichthyosauriens, sauropterygiens, pterosauriens et cheloninens du Musée d'Histoire Naturelle de Boulogne-sur-Mer. In E. Buffetaut, Rose and Vadet (eds.). Vértébrés Fossiles du Boulonnais. Mémoires de la Société Académique du Boulonnais. 1(2), 85-97.
Rose, 1987. Redecouverte d'une vertebre caudale reptilienne (Archosauriens) de status controverse et provenant des terrains jurassiques superieurs du Boulonnais. Bulletin de la Société académique du Boulonnais. 1(5), 150-153.
Buffetaut, Cuny and le Loeuff, 1991. French Dinosaurs: The best record in Europe? Modern Geology. 16(1-2), 17-42.
Buffetaut and Martin, 1993. Late Jurassic dinosaurs from the Boulonnais (northern France): A review. Revue de Paléobiologie. 7(vol. spéc.), 17-28.
Ford, 2005 online. http://www.paleofile.com/Dinosaurs/Theropods/Teinurosaurus.asp
And before we go, here are a couple more tidbits I've noticed in the upcoming update...
- That theropod tail preserved in Burmese amber (DIP-V-15103) described by Xing et al. (2016) was only placed as specifically as a non-pygostylian maniraptoriform. But as the deposits are Gondwanan (e.g. Poinar, 2018), the range of potential Cenomanian theropods is better understood. And only one group has caudal centra over three times longer than tall- unenlagiines. I bet DIP-V-15103 is our first sample of preserved plumage in an unenlagiine, which makes you wonder if the weird alternating barb placement was a feature that evolved on Gondwana, and if so did Rahonavis' remiges exhibit it too?
- Does anyone realize both "Tralkasaurus" (Cerroni et al., 2019) and "Thanos" (Delcourt and Iori, 2018) are nomina nuda? Neither are in an official volume yet, though "Tralkasaurus" is scheduled for March and "Thanos" will probably make it this year if the average papers per volume of Historical Biology holds up. "Tralkasaurus" has an empty space in its "Zoobank registration:" section, while the "Thanos" paper doesn't mention ZooBank at all, and neither show up in ZooBank searches. Also, one of "Thanos"' supposed autapomorphies is a deep prezygapophyseal spinodiapophyseal fossa, which does not exist in abelisaurs as it would require a spinodiapophyseal lamina. The labeled structure seems internal, probably the centroprezygapophyseal fossa or prezygapophyseal centrodiapophyseal fossa based on CT-scanned noasaurid cervical DGM929-R. That leaves axial pleurocoel size and distance from each other, and ventral keel strength as suggested characters. Which can only be compared to Carnotaurus among brachyrostrans. Hmmm...
References- Xing, McKellar, Xu, Li, Bai, Persons IV, Miyashita, Benton, Zhang, Wolfe, Yi, Tseng, Ran and Currie, 2016. A feathered dinosaur tail with primitive plumage trapped in Mid-Cretaceous amber. Current Biology. 26(24), 3352-3360.
Delcourt and Iori, 2018. A new Abelisauridae (Dinosauria: Theropoda) from São José do Rio Preto Formation, Upper Cretaceous of Brazil and comments on the Bauru Group fauna. Historical Biology. DOI: 10.1080/08912963.2018.1546700
Poinar, 2018. Burmese amber: Evidence of Gondwanan origin and Cretaceous dispersion. Historical Biology. DOI: 10.1080/08912963.2018.1446531
Cerroni, Motta, Agnolín, Aranciaga Rolando, Brissón Egli and Novas, 2019. A new abelisaurid from the Huincul Formation (Cenomanian-Turonian; Upper Cretaceous) of Río Negro province, Argentina. Journal of South American Earth Sciences. 98, 102445.
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Oculudentavis is not a theropod
Hi all. This week we got the announcement of a tiny theropod skull in Myanmar amber, which was bound to happen eventually as amazing finds from that deposit keep being published. Alas, whatever Oculudentavis is, it's not a theropod.
Oculudentavis skull (after Xing et al., 2020).
Just look at it. No antorbital fenestra, incomplete ventral bar to the laterotemporal fenestra, huge posttemporal fenestrae, teeth that extend posteriorly far under the orbit...
All of which might be coincidental, but then look at the mandible.
Oculudentavis mandible (after Xing et al., 2020).
That spike-like coronoid process is classic lepidosaur, plus the dentary is way too long compared to the post-dentary elements, then the description says "The tooth geometry appears to be acrodont to pleurodont; no grooves or sockets are discernable." And of course "the scleral ring is very large and is formed by elongated spoon-shaped ossicles; a morphology similar to this is otherwise known only in lizards (for example, Lacerta viridis)."
Add to this the size of this partially fused specimen being smaller than any extant bird (14 mm), and no feather remains, and why is this a theropod again? The endocast is big, but why not a clade of brainier lizards or late surviving megalancosaurs by the Cenomanian?
The authors add it to Jingmai's bird analysis where it ends in a huge polytomy closer to Aves than Archaeopteryx, but outside fake Ornithuromorpha. That's often what happens when a taxon is wrongly placed in a clade. Note the figured placement between Archaeopteryx and Jeholornis is only found using implied weights. At least add it to e.g. Nesbitt's or Ezcurra's archosauromorph analyses, or Cau's theropod analyses before assuming it's a bird.
Thanks to Ruben Molina Perez for suggesting this issue in the first place.
Reference- Xing, O'Connor, Schmitz, Chiappe, McKellar, Yi and Li, 2020. Hummingbird-sized dinosaur from the Cretaceous period of Myanmar. Nature. 579, 245-249.
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What is Oculudentavis if it's not a theropod?
In my last post, I argued the recently described Oculudentavis (Xing et al., 2020) is not a theropod. So what is it? To answer that question, I entered it into Simoes et al.'s (2018) sauropsid analysis which emphasizes basal lepidosauromorphs and comes out with basal gekkos and nested iguanians even using just morphological characters. To test Jingmai's avialan hypothesis, I also added Archaeopteryx to the matrix. The result is 384 MPTs of 2337 steps each.
Strict consensus of 384 MPTs of Simoes et al.'s (2018) analysis after adding Oculudentavis and Archaeopteryx. Compare to Extended Data Figure 3 of Simoes et al..
As you can see, Oculudentavis resolves as a stem-squamate in a trichotomy with Huehuecuetzpalli and squamates, while Archaeopteryx is an archosauromorph sister to Erythrosuchus. And this matrix didn't score for scleral ossicle shape, posttemporal fenestra size or maxillary tooth row length. After scoring Oculudentavis, its teeth are clearly not acrodont, it seems to have a ventral parietal fossa and lacks an ossified laterosphenoid. The authors could have made it easier to evaluate by separating the cranial elements in the 3D pdf file. As it is, a lot of palatal and braincase info is uncertain. But Huehuecuetzpalli is Albian compared to Oculudentavis' Cenomanian, and has a skull length of 32 mm (19 mm in the juvenile) versus 14 mm in Oculudentavis.
Huehuecuetzpalli skull (top; after Reynoso, 1998), Oculudentavis skull and separate mandible (middle; after Xing et al., 2020), and Archaeopteryx skull (after Rauhut, 2014).
References- Reynoso, 1998. Huehuecuetzpalli mixtecus gen. et sp. nov: A basal squamate (Reptilia) from the Early Cretaceous of Tepexi de Rodriguez, central Mexico. Philosophical Transactions of the Royal Society B: Biological Sciences. 353, 477-500.
Rauhut, 2014. New observations on the skull of Archaeopteryx. Paläontologische Zeitschrift. 88(2), 211-221.
Simōes, Caldwell, Talanda, Bernardi, Palci, Vernygora, Bernardini, Mancini and Nydam, 2018. The origin of squamates revealed by a Middle Triassic lizard from the Italian Alps. Nature. 557(7707), 706-709.
Xing, O'Connor, Schmitz, Chiappe, McKellar, Yi and Li, 2020. Hummingbird-sized dinosaur from the Cretaceous period of Myanmar. Nature. 579, 245-249.
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It's finally January 1, 200n and Phylonyms is published!
Ah the PhyloCode, the so-called future of biological nomenclature whose release has always kept on slipping ever more distantly into the future. After 20 years of waiting, we now have Phylonyms: A Companion to the PhyloCode, by de Queiroz et al. (2020), "a turning point in the history of phylogenetic nomenclature" according to its introduction. As the book states "Phylonyms serves as the starting point for phylogenetic nomenclature governed by the PhyloCode. According to the preamble, “This code will take effect on the publication of Phylonyms: A Companion to the PhyloCode, and it is not retroactive.” Thus, names and definitions published here have precedence over any competing names and definitions published either before (or after) the publication of Phylonyms." So for anyone invested in standardized phylogenetic nomenclature, this is it. Nothing better is coming down the pipeline in our lifetimes, so let's see what we're stuck with.
First of all, it's expensive. You can get an ebook for $222 on Amazon or a hardcover sometime after June 9th for $234. I found an electronic version for $169 plus tax on VitalSource, but you have to use their reader. It's 1323 pages though, so isn't a bad deal. That's less than five $37.95 Cretaceous Research pdfs, and I figure this is one of those historical volumes it's good to have, like Sibley and Ahlquist's bird phylogeny book.
The format is an encyclopedia-style list of clades in phylogenetic order with Registration Number, Definition, Etymology, Reference Phylogeny, Composition, Apomorphies, Synonyms, Comments and Literature Cited. Rather like my Theropod Database, so no complaints there. Well, one complaint is really more to do with the PhyloCode itself where they decided to abbreviate definitions with the non-standard del/nabla triangle symbol ∇. If you want people to start using your format, you might want to choose symbols that exist on a standard keyboard. Alt+2207 is supposed to generate it in Windows, but results in ƒ here in Blogger. Anyone know the correct Unicode numbers?
On to the substance, where Phylonyms covers all life. Dinosaurs are the last section of the book, and non-avian dinosaurs get all of four definitions-
Dinosauria R. Owen 1842 [M. C. Langer, F. E. Novas, J. S. Bittencourt, M. D. Ezcurra, and J. A. Gauthier], converted clade name
Registration Number: 194
Definition: The smallest clade containing Iguanodon bernissartensis Boulenger in Beneden 1881 (Ornithischia/Euornithopoda) Megalosaurus bucklandii Mantell 1827 (Theropoda/Megalosauroidea) and Cetiosaurus oxoniensis Phillips 1871 (Sauropodomorpha).
I'm glad we've standardized which theropod, ornithischian and sauropodomorph are used (or so I thought, see below), but otherwise there's not much to say. The caveats around which apomorphies are also found in Nyasasaurus and at least some silesaurs illustrate why apomorphy-based definitions are bad. The reference phylogeny for this and Saurischia is Lloyd et al.'s (2008) supertree, which is quite outdated and has a lot of artifacts from being a supertree.
Saurischia H. G. Seeley 1888 [J. A. Gauthier, M. C. Langer, F. E. Novas, J. Bittencourt, and M. D. Ezcurra], converted clade name
Registration Number: 195
Definition: The largest clade containing Allosaurus fragilis Marsh 1877 (Theropoda/Carnosauria) and Camarasaurus supremus Cope 1877 (Sauropodomorpha), but not Stegosaurus stenops Marsh 1887 (Ornithischia/Stegosauridae).
It's rather odd the same authors didn't choose the same specifiers for each dinosaurian clade as they did in the previous definition, leaving us without a neat node-stem triplet. Instead they went with the Kischlatian approach of using taxa"mentioned and figured as examples of their respective groups by Seeley (1888)." This is funny because I don't think this rationale is ever suggested in the PhyloCode, whereas Dinosauria and Saurischia are actually the official examples used for Recommendation 11F encouraging node-stem triplets ("If it is important to establish two names as applying to sister clades regardless of the phylogeny, reciprocal maximum-clade definitions should be used in which the single internal specifier of one is the single external specifier of the other, and vice versa"). Specifically- "If one wishes to define the names Saurischia and Ornithischia such that they will always refer to sister clades, Saurischia might be defined as the largest clade containing Megalosaurus bucklandiiMantell 1827 but not Iguanodon bernissartensisBoulenger in Beneden 1881, and Ornithischia would be defined as the largest clade containing Iguanodon bernissartensis but not Megalosaurus bucklandii. To stabilize the name Dinosauria as referring to the clade comprising Saurischia and Ornithischia, Dinosauria should be defined as the smallest clade containing Megalosaurus bucklandii and Iguanodon bernissartensis."
Ornithoscelida and its consequences are mentioned, but I'm glad more time is not taken up with it as I expect the hypothesis to fall away as Baron's phylogenetic mistakes are not followed by future authors.
Sauropodomorpha F. R. von Huene 1932 [M. Fabbri, E. Tschopp, B. McPhee, S. Nesbitt, D. Pol, and M. Langer], converted clade name
Registration Number: 295
Definition: The largest clade containing Saltasaurus loricatus Bonaparte and Powell 1980 (Sauropodomorpha) but not Allosaurus fragilis Marsh 1877 (Theropoda) and Iguanodon bernissartensis Boulenger in Beneden 1881 (Ornithischia).
I dislike the use of Saltasaurus as the internal specifier, which is a holdover of Sereno's weird use of deeply nested OTUs when others would be more historically relevant and/or eponymous. Fabbri et al. defend the choice because "Fossil specimens referred to Saltasaurus loricatus are abundant, the species is well known, and its phylogenetic position is consistent among phylogenetic analyses", but this would be even more true for e.g. Camarasaurus supremus used in Saurischia's definition. The other specifiers are a mix of those in Dinosauria's and Saurischia's definition, so there's absolutely no consistency. The reference phylogeny is Otero et al.'s (2015) Sefapanosaurus description using Yates' matrix, so is fine.
There's a rare error in the comments for this entry. Fabbri et al. state "Segnosaurus galbinensis from the Cretaceous was briefly thought to be a relatively early diverging sauropodomorph (Paul, 1984; Gauthier, 1986; Olshevsky, 1991). More material referable to that species and the discovery of closely related taxa later showed that Segnosaurus galbinensis is part of the Therizinosauria", but material of S. galbinensis besides that initially recovered in the 1970s is not known.
Theropoda O. C. Marsh 1881 [D. Naish, A. Cau, T. R. Holtz, Jr., M. Fabbri, and J. A. Gauthier], converted clade name
Registration Number: 216
Definition: The largest clade containing Allosaurus fragilis Marsh 1877 (Theropoda) but neither Plateosaurus engelhardti Meyer 1837 (Sauropodomorpha) nor Heterodontosaurus tucki Crompton and Charig 1962 (Ornithischia).
Here we've chosen two completely different specifiers for Sauropodomorpha and Ornithischia, so again we have no consistency. The reference phylogeny is Cau (2018), which is ideal.
What about the rest? It's a HUGE volume, and obviously most of Pan-Biota is outside my area of expertise. One obvious issue is the wildly varying coverage of different clades. Apparently nobody could be bothered with the vast majority of vertebrates (euteleosts) or animals (insects, except one definition for Trichoptera), and Molluska doesn't even get a definition. But we do get several entries for edrioasterid taxa down to subfamily-level, generally obscure Paleozoic echinoderms. Closer to dinosaurs, there's nothing at all for pan-crocs, but we get an entry for Pterosauromorpha for which only Scleromochlus is given as a plausible non-pterosaurian example (perhaps wrongly- Bennett, 2020).
Then there are the apomorphy-based definitions which will cause headaches in the future. Look at Apo-Chiroptera- "Definition: The clade for which the unique modifications of the hand, forearm, humerus, scapula, hip, and ankle (see Diagnostic Apomorphies) associated with flapping flight, as inherited by Vespertilio murinus Linnaeus 1758, are apomorphies." Then you go down to the nine listed sets of Diagnostic Apomorphies like "Modification of the scapula: Scapular spine originates at the posterior edge of the glenoid fossa. Long axis of scapular spine offset 20–30 degrees from axis of rotation of the humeral head. Scapular spine reduced in height—acromion process appears more strongly arched and less well supported than in other mammals. Presence of at least two facets in infraspinous fossa." These are all going spread out as more stem bats are discovered, and indeed the authors already note "Simmons and Geisler (1998) included the absence of claws on wing digits III-V with this suite of modifications; however, the presence of claws on all the wing digits of Onychonycteris suggests that claws were present primitively in Apo-Chiroptera."
Ungulata is defined by Archibald as "The least inclusive crown clade containing Bos primigenius Bojanus 1827 (= Bos taurus Linnaeus 1758) (Artiodactyla) and Equus ferus Boddaert 1785 (= Equus caballus Linnaeus 1758) (Perissodactyla), provided that this clade does not include Felis silvestris Schreber 1777 (= Felis catus Linnaeus 1758) (Carnivora), Manis pentadactyla Linnaeus 1758 (Pholidota), Vespertilio murinus Linnaeus 1758 (Chiroptera), or Erinaceus europaeus Linnaeus 1758 (Lipotyphla)." But this doesn't exist in molecular studies, including those of ultraconserved elements, which consistently place carnivorans, pangolins and bats closer to perrisodactyls. So this is likely to be a historical footnote, as well established molecular relationships end up trumping morphological relationships in every example I know of.
Finally, we get Pan-Lepidosauria for the total group of lepidosaurs, which has been Lepidosauromorpha for over thirty years. Yet Archosauromorpha is retained as "The least inclusive clade containing Gallus (originally Phasianus) gallus (Aves) (Linnaeus 1758), Alligator (originally Crocdilus) mississippiensis (Daudin 1802) (Crocodylia), Mesosuchus browni Watson 1912 (Rhynchosauria), Trilophosaurus buettneri Case 1928 (Trilophosauridae), Prolacerta broomi Parrington 1935 (Prolacertiformes), and Protorosaurus speneri von Meyer 1830 (Protorosauria)" even though Pan-Archosauria is also used for the total group of archosaurs, traditionally the definition of Archosauromorpha. I agree our new Archosauromorpha deserved a name for being a generally recognized group, whereas whether e.g. choristoderes or sauropterygians fell out closer to lizards or birds is highly unstable. But I would have rather kept the tradition of -omorpha for the stem clades and gave this a new name.
Overall, I'm not very impressed for something 20 years in the making that intends to be so important. How do you contradict your own example for choosing specifiers in four papers, where two share the same author list, the other two share another author (Fabbri), and each of those shares an author with both of the first two (Langer and Gauthier)? And one of those is an editor for the volume. Nothing could be negotiated in over two decades? But it's what we have to work with now, and in the name of consistancy I'll adopt the definitions proposed. Now to see what happens when RegNum goes online.
References- Lloyd, Davis, Pisani, Tarver, Ruta, Sakamoto, Hone, Jennings and Benton, 2008. Dinosaurs and the Cretaceous terrestrial revolution. Proceedings of the Royal Society B. 275, 2483-2490.
Otero, Krupandan, Pol, Chinsamy and Choiniere, 2015. A new basal sauropodiform from South Africa and the phylogenetic relationships of basal sauropodomorphs. Zoological Journal of the Linnean Society. 174, 589-634.
Cau, 2018. The assembly of the avian body plan: A 160-million-year long process. Bollettino della Società Paleontologica Italiana. 57(1), 1-25.
Bennett, 2020. Reassessment of the Triassic archosauriform Scleromochlus taylori: Neither runner nor biped, but hopper. PeerJ. 8:e8418.
de Queiroz, Cantiono and Gauthier, 2020. Phylonyms: A Companion to the PhyloCode, 1st Edition. Taylor & Francis Group. 1323 pp.
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The Unecessary Death of Steneosaurus
Not a dinosaur, but a new paper on the classic crocodylomorph Steneosaurus exemplifies a troubling trend in recent vertebrate taxonomy. Johnson et al. (2020) reexamine the original material of Steneosaurus, an aquatic croc from the Jurassic of France. It hadn't been seriously looked at since the 1860s, so this is one of my favorite kinds of paleontology papers- restudying a fragmentary old specimen in a modern light. What do they find?
We first get a detailed recount of its history, with two decades as Cuvier's "tête à museau plus allongé" (= head with a more elongated snout; I have to praise the authors for translating all the French to English, even in our spoiled era of Google Translate it saves time), before it was named Steneosaurus rostro-major by Geoffroy Saint-Hilaire in 1825. Eudes Deslongchamps and son tackled it in the 1860s, where they viewed the specimen as too poorly preserved and so "stated that the taxon to represent the genus Steneosaurus should be either ‘Steneosaurus’ megistorhynchus Eudes-Deslongchamps, 1866, or ‘Steneosaurus’ edwardsi Eudes-Deslongchamps, 1868c." Ha! You don't get to just take somebody's genus and affix your new species as its type. They were the last to examine the specimen in detail however, making that a pretty bad note to end on.
Johnson et al. then reexamine the type snout of Steneosaurus, correcting the species name by eliminating the hyphen, officially making it the lectotype, noting Steel had determined the posterior skull to be Metriorhynchus, and illustrating and redescribing the specimen. Excellent work and very well done. After eliminating Mycterosuchus nasutus, 'Steneosaurus' leedsi, 'S.' heberti and Lemmysuchus and other machimosaurins based on numerous dissimilar characters, the authors come to the contemporaneous 'Steneosaurus' edwardsi.
"As mentioned before, this was a second species that Eudes-Deslongchamps (1867–69) considered identical to S. rostromajor. These two taxa share a combination of features including:
1. A subcircular, moderately interdigitating premaxilla-maxilla suture.
2. Maxillae ornamented with irregular grooves.
3. A shallower mediolateral compression of the posterior maxillae, as opposed to ‘S.’ heberti (MNHN.F 1890-13).
4. Horizontally flat posterior premaxilla in lateral view.
5. Deep anterior and mid-maxillary reception pits that gradually become shallower towards the posterior maxilla.
6. Subcircular to circular alveoli that remain relatively the same size throughout the maxilla.
7. Teeth with well-pronounced enamel ridges at the base."
Well how cool is that? They put in the hard work, found the matching more complete specimens, and now we have Steneosaurus edwardsi as a junior synonym of S. rostromajor, giving us a good look at what Steneosaurus really was after two hundred years.
Lectotype of Steneosaurus rostromajor (MNHN.RJN 134c-d) in dorsal (A, B) and ventral (C, D) views. (after Johnson et al., 2020).
But no.
Johnson et al. immediately say "it is important to note that many of these characters may, in fact, be related to sexual dimorphism, ontogeny and intraspecific variation." True, but that could be said for basically every character supposed to diagnose Mesozoic croc genera, or theropod genera, pterosaur genera, etc.. Unless you have some specific example like 'enamel ridges have been shown to develop with age and both S. rostromajor and S. edwardsi are larger than S? leedsi or S? heberti with weak ridges', then it's just hand-waving. And no, Johnson et al. never develop such an argument for one of those characters, let alone all seven.
Next, we get "In addition to the sexual dimorphism/ontogeny problem, one of the critical issues about MNHN.RJN 134c-d is that it is poorly preserved." Sure, but you were still able to perform many comparisons. Again, the authors never say any of their seven characters are taphonomic, so it's another objection without substance.
Yet the worst rationale for rejecting Steneosaurus is "in reality, the name Steneosaurus is extremely impractical. It was used for many metriorhynchid specimens (e.g. ‘Steneosaurus’ gracilis, ‘Steneosaurus’ palpebrosus and ‘Steneosaurus’ manselii) during much of the 19th century, largely in part due to Cuvier’s metriorhynchid skull region (MNHN. RJN 134a-b) being attributed to the teleosauroid rostral section (MNHN.RJN 134c-d). Indeed, the concise, classical definition of ‘Steneosaurus’ as we interpret it today was not given until the work of both Eudes-Deslongchampses (1868c, 1867–69)"
Substitute Megalosaurus in there to see how ridiculous it is. That has had over 45 species assigned to it, and was named in the 1820s but didn't have a modern concept associated with it until the 1980s. When Johnson et al. lament that for Steneosaurus "rather than comparing characters outright, comparison is by process of elimination (or the question of ‘what features does this specimen lack?’)", that perfectly describes the Megalosaurus paralectotype dentary.
"After the Eudes-Deslongchampses’ treatment, what was left was an undiagnostic, chimeric type specimen for S. rostromajor (MNHN.RJN 134) and the genus Steneosaurus was redefined using a new type species that was not accepted by some researchers. In addition, since the Eudes-Deslongchampses, there has been no attempt to rectify this taxonomic nightmare;"
You just showed it was diagnostic, Steel long ago got rid of the chimaeric portion, Eudes-Deslongchamps' stupid attempts to name new type species have no relevance, and you have done the work to finally rectify this taxonomic nightmare.
"Due to these three significant factors (uncertainty of variable characters, poor preservation and
unreasonable name), we have concluded that S. rostromajor, and therefore ‘Steneosaurus’ (MNHN.RJN 134c-d), cannot be confidently assigned to an existing teleosauroid species."
Nope, you just showed it can be assigned to the same species as S. edwardsi.
Actually, I correct myself. THIS is the worst rationale for rejecting Steneosaurus- "In addition, MNHN.RJN 134c-d was initially diagnosed based on significant orbital and temporal characteristics (from the metriorhynchid MNHN.RJN 134a-b), along with generic rostral ones. Because the skull material is now known to be from a metriorhynchid, this ‘hybrid type specimen’ factor adds to the doubtful validity of Steneosaurus. According to Article 23.8 of the ICZN Code, ‘a species-group name established for an animal later found to be a hybrid (Art. 17) must not be used as the valid name for either of the parental species (even if it is older than all other available names for them)’ (this also signifies that the species name rostromajor is itself invalid). As such, MNHN.RJN 134c-d serves as an undiagnostic specimen; we, therefore, consider MNHN.RJN 134c-d to be a nomen dubium and, as such, Steneosaurus is treated as an undiagnostic genus."
If the term "parental species" didn't tip you off, Article 23.8 applies to hybrid individuals (those resulting from different species interbreeding), not type specimens chimaerically combined from multiple species. The Article doesn't even say what Johnson et al. think- it says a name for a hybrid can't be used for either of the species that bred to make it, so that e.g. even if a mule's scientific name was erected prior to that of horse's or ass's, it can't be the name for horse or ass. And indeed even the cited Article 17 says that hybrids and chimaeras can be the basis of valid names- "The availability of a name is not affected even if 17.1. it is found that the original description or name-bearing type specimen(s) relates to more than one taxon, or to parts of animals belonging to more than one taxon; or 17.2. it is applied to a taxon known, or later found, to be of hybrid origin..."
If Johnson et al.'s interpretation were right, there goes Gojirasaurus, Protoavis, Chuandongocoelurus, Chilantaisaurus, Fukuiraptor, Coelurus, Alectrosaurus, Dakotaraptor, etc..
Before the big reveal, we have in the Conclusion what can only be described as a lie- "Through character comparison-and-elimination, the only taxon with which MHNH.RJN 134c-d could hypothetically be referred to is ‘S.’ edwardsi, but the two do not share any clear autapomorphic characters or a unique combination of characters."
What are your seven listed characters if not "a unique combination of characters"? Does any other teleosaurid have them? If not, they are unique. In any case, we get the motivation for dumping Steneosaurus twice at the end of the paper-
"We believe that establishing teleosauroid taxonomy from the beginning with a series of ‘clean’ type species/specimens, with every nomenclatural act correctly formulated, is the best course of action, which we will highlight in a forthcoming paper (Johnson, 2019)."
"We believe that establishing teleosauroid taxonomy from the beginning with a series of ‘clean’ type species/specimens, with every nomenclatural act correctly formulated, is the best course of action. This will necessitate a revised teleosauroid taxonomy, in which species previously referred to the genus Steneosaurus are given new generic names. This work will be published by us in a separate contribution, based on the comprehensive teleosauroid phylogenetic analysis in Johnson’s PhD thesis (2019)."
Basically everything I hate about a current trend in vertebrate paleontology- just throw out old specimens and dishonor their authors who correctly reported what was new at the time to come up with your own names. At least dumping Stegosaurus armatus or designating a neotype for Allosaurus fragilis could be claimed to save time and effort actually analysing the types, if you don't want to do the science to figure out if armatus is actually different from stenops or if fragilis can be distinguished from Saurophaganax. But Johnson et al. already did all the hard work and found Steneosaurus edwardsi was S. rostromajor, they would just rather use Johnson's new genus name for the taxon.
And their reasons are just grasping at straws. 'Sure we identified these seven charactesrs uniquely shared by Steneosaurus rostromajor and S. edwardsi, but uhh.. could be sexually dimorphic? Or anything could be individual variation. Or ontogenetic? Lots of things turn out to be ontogenetic. Plus it's broken. Sooo broken. Sure we could evaluate characters, but who wants a taxon whose holotype isn't pristine? Plus a lot of people had stupid ideas about Steneosaurus over the past two hundred years. What do us scientists do when we have a complicated situation to resolve that was only partially understood historically? Trash their names and give yourselves credit for new genera.' Thus Steneosaurus gets the eternal identity of "all evidence points to it being Johnsonosaurus edwardsi, but ehhh... we just sort of ignore it now as Teleosauridae indet. and it's forgotten."
To conclude, Steneosaurus is really outside my wheelhouse. But if Johnson et al.'s philosophy spreads, we're in danger of losing a lot of historical taxonomy and deserved credit to lazy or selfish authors. Just look at Microraptor for example, whose holotype of M. zhaoianus lacks a decent skull. Some decades down the line, what if cranial differences support various Jiufotang species and someone's like 'the postcranial proportions are unique between the M. zhaoianus type and M. hanqingi, but I want a complete type specimen, so Microraptor is an invalid undiagnostic nomen dubium, and instead I propose Mybetterraptorgenus hanqingi and M. gui.' Just hope they don't pull a Wilson and Upchurch and claim 'Microraptor is invalid and co-ordinate suprageneric Linnean taxa must likewise be abandoned' and replace Microraptorinae with Mybetterraptorgenusinae.
References- Johnson, 2019. The taxonomy, systematics and ecomorphological diversity of Teleosauroidea (Crocodylomorpha, Thalattosuchia), and the evaluation of the genus 'Steneosaurus'. PhD Thesis, University of Edinburgh. 1062 pp.
Johnson, Young and Brusatte, 2020. Emptying the wastebasket: A historical and taxonomic revision of the Jurassic crocodylomorph Steneosaurus. Zoological Journal of the Linnean Society. 189(2), 428-448.
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The Arguable Identity of Paraxenisaurus
Hi everyone. In light of Cau's recent post on the supposed new Mexican deinocheirid "Paraxenisaurus normalensis" (Serrano-Brañas et al., 2020), I figured I'd check the taxon out to see what I thought.
The first thing you might note are the quotation marks surrounding its name, as this is yet another example of authors not including an lsid or reference to ZooBank in their electronic descriptions. ICZN Article 8.5.3. states names published electronically must "be registered in the Official Register of Zoological Nomenclature (ZooBank) (see Article 78.2.4) and contain evidence in the work itself that such registration has occurred", and the pre-print is said to be in preparation for Volume 101 of Journal of South American Earth Sciences, cited as August 2020. Thus it gets to join the ranks of "Thanos" and "Trierarchuncus"as theropods that will eventually be validly named this year. But at least it's not stuck in the purgatory of twelve Scientific Reports Mesozoic theropods, which will never be physically published and thus will remain invalid unless outside action is taken.
One of the big takeaways from Cau's blogpost is that "I am doubtful about the possibility of referring these elements [the paratypes] to the same species of the holotype, since there are very few superimposable elements among the three specimens. Therefore, there is a risk that Paraxenisaurus , - understood as the sum of all three specimens - is a chimera." After reading the paper, Andrea REALLY undersold this critique. Here are the specimen materials lists, with the overlapping elements highlighted in matching colors-
(BENC 2/2-001; proposed holotype) proximal manual phalanx II-2 or III-3, partial astragalocalcaneum, partial metatarsal II, phalanx II-1 (115 mm), proximal phalanx II-2, partial metatarsal III, proximal phalanx III-3, distal metatarsal IV, phalanx IV-1 (104 mm), phalanx IV-3 (67 mm), phalanx IV-4 (45 mm), partial pedal ungual IV
(BENC 1/2-0054) distal metacarpal I, proximal phalanx I-1, partial manual ungual I, distal metacarpal II, distal phalanx II-2
(BENC 1/2-0091) several proximal caudal central fragments (66, 75, 76 mm), proximal metacarpal II, partial metacarpal III, distal femur (155 mm trans), distal metatarsal IV
(BENC 1/2-0092) several distal caudal vertebrae (70, 71 mm)
(BENC 30/2-001) pedal ungual II, pedal ungual III
As you can see, there's only one strict overlap, with BENC 1/2-0091 sharing a distal metatarsal IV with the proposed holotype, found ~14 kilometers away. The paper lists no proposed apomorphies or unique combination of characters for distal metatarsal IV, and indeed the description states they preserve largely non-overlapping portions-
"In the holotype, the distal articular surface is fragmented (Figures 11a1 and 11a2); but in the referred specimen (BENC ½-0091), this surface is nicely preserved and has a non-ginglymoid outline (Figures 11b1 and 11b2). The medial condyle is mostly preserved in the holotype (Figure 11a3), but in the referred specimen it is completely broken (Figure 11b3). Conversely, the lateral condyle is broken in the holotype (Figure 11a4), but is well preserved in the referred specimen (Figure 11b4). Collateral ligament fossae are well developed on both condyles and have approximately the same size and depth (Figures 11a3 and 11b4). In cross-section, the shaft of metatarsal IV near the distal end is thicker dorsoventrally than wide."
Needless to say, metatarsal IV has a shaft which is deeper than wide in all ornithomimosaurs, and the preserved ligament fossae are on opposite sides in each specimen (medial in proposed holotype, lateral in 1/2-0091). Below is a figure comparing the two Mexican specimens with Ornithomimus velox, with 1/2-0091 flipped so that all are comparable as left elements. I don't see anything the "Paraxenisaurus" specimens have in common that could diagnose a taxon.
Left distal metatarsal IV of (left to right) intended "Paraxenisaurus normalensis" holotype BENC 2/2-001, intended "Paraxenisaurus normalensis" paratype BENC 1/2-0091 (right element flipped), and Ornithomomus velox holotype YPM 542 in (top to bottom) dorsal, lateral, ventral and medial views ("Paraxenisaurus" after Serrano-Brañas et al., 2020; Ornithomimus after Claessens and Loewen, 2016).
While no other elements are exactly matched, referred specimen BENC 30/2-001 does include pedal unguals II and III, while the intended holotype has pedal ungual IV. These are again from different localities, although closer this time (~2.8 km), and this time we have characters listed in the diagnosis-
"(9) distinctively broad and ventrally curved pedal unguals that angled downward with respect to the proximal articular surface and depending on the digit, the proximodorsal process becomes slightly enlarge and changes its position from nearly horizontal to mostly vertical, adopting a lipshaped appearance; and (10) pedal unguals with a rounded, large foramen on the medial side* and a deep ventral fossa that surrounds a strongly developed, ridge-like flexor tubercle."
Pedal unguals of (left to right) intended "Paraxenisaurus normalensis" paratype BENC30/2-001 right digit II, right digit III and intended "Paraxenisaurus normalensis" holotype BENC 2/2-001 left digit IV in (top to bottom) right, left, proximal and dorsal views (after Serrano-Brañas et al., 2020). Green lines point to supposedly natural median foramen
Ventral curvature is plesiomorphic, the unguals of BENC 30/2-001 are not broader than other ornithomimosaurs', and ventral angling with the proximal end held vertically is common in theropods and present in e.g. Garudimimus and Beishanlong. The proximodorsal process "changing its position" is using a difference between 30/2-001's mostly horizontal processes and the intended holotype's more vertical process as character, which in itself presupposes they are the same taxon. The ventral fossa surrounding a ridge-like flexor tubercle is also present in Harpymimus, Garudimimus, Beishanlong and large Dinosaur Park unguals (NMC 1349, RTMP 1967.19.145) and is not shown in the intended holotype but is claimed to be "partially broken." This leaves the medial foramen, which might be a valid character in unguals III and IV (II is damaged in that area), but might also be taphonomic, as there are many other small circular areas of damage (e.g. center of proximal surface of ungual IV). While the two unguals in 30/2-001 are similar to each other, that of the intended holotype is more strongly curved, has that smaller more dorsally angled proximodorsal process, is wider in proximal view, and lacks the expanded ventral half characteristic of ornithomimosaurs that is present in the other specimen. But even if these two pedal unguals are correctly referred, they are all that's present in specimen BENC 30/2-001. So they get us nowhere in determining caudal, manual (besides proximal manual phalanx II-2 or III-3) or femoral morphology.
The final issue I noticed was the emphasis on "Paraxenisaurus" having a first pedal digit. This would ironically be unlike Deinocheirus, but plesiomorphically shared with Nedcolbertia, "Grusimimus", Garudimimus, Beishanlong, Archaeornithomimus and Sinornithomimus. The character state is based on metatarsal II, where "a facet on the posterior surface of the distal quarter of this shaft, indicates the presence of an articulation area for metatarsal I." The figure shows a longitudinal groove extending down the posterior center of distal metatarsal II, which as anyone who has scored taxa for Clarke's bird matrix could tell you, is not how non-birds attach their hallux to the metatarsus. Hattori (2016) for instance writes in Allosaurus "there is no attachment scar corresponding to the metatarsal I fossa on either medial or plantar aspect of MT II" and in Citipati "there is no obvious attachment scar of MT I on either medial or plantar aspect of MT II." Serrano-Brañas et al. state "in Garudimimus brevipes ... the attachment site is also placed in the same area as in Paraxenisaurus normalensis", but the feature in Garudimimus is a raised scar with sharp medial demarcation from the shaft. As Middleton (2003) recognized, this scar is for the m. gastrocnemius, specifically the m. gastrocnemius pars medialis (Carrano and Hutchinson, 2002), and I'll note it's present even in Gallimimus which lacks pedal digit I (Osmolska et al., 1972: Plate XLIX Fig. 1b). "Paraxenisaurus"'s groove is then more likely to be the m. flexor digitorum longus II tendon, which "passed through the ventral groove in its respective metatarsal to insert serially on each of the pedal phalanges" in e.g. Tyrannosaurus (Carrano and Hutchinson, 2002).
Left metatarsal II in ventral view of (left to right) intended "Paraxenisaurus normalensis" holotype BENC 2/2-001 (after Serrano-Brañas et al., 2020; yellow arrow points to supposed articulation for metatarsal I), Garudimimus brevipes holotype IGM 100/13 (after Kobayashi and Barsbold, 2005; line points to supposed articulation for metatarsal I), Gallimimus bullatus ZPAL MgD-I/94 (after Osmolska et al., 1972), and Tyrannosaurus rex FMNH PR2081 (after Brochu, 2003).
What exactly is "Paraxenisaurus"? Comparison is hindered by the specimens being figured mixed together, and the figures are not in numerical order in the preprint, being shown in the order of- 1, 10-19, 2, 20-23, 3-9. In addition, the scale bars vary within the same figure (e.g. phalanx IV-1 is proximally ~61 mm wide in figure 14a but ~93 mm wide in figure 14e) and the listed measurements are different yet (e.g. IV-1 is listed as 83 mm wide). Thus any composite reconstruction is necessarily approximate. The supposed manual element is too fragmentary to give much information, but it is of the appropriate size and shape to be a proximal pedal phalanx I-1. This would make more sense preservationally since the other material preserved in the specimen is all from the tarsus and pes. It's a shame the astragalocalcaneum is not described better or figured in more views, as the dorsal (= proximal?) perspective has many broken surfaces and edges, so that e.g. the small calcaneum might be preservational. The fused proximal tarsals are like ceratosaurs, deinocheirids (Deinocheirus plus Hexing), alvarezsaurids and caenagnathids. Having any sense of the ascending process morphology could tell us much. Metatarsal II is not obviously deeper than wide, unlike ornithomimosaurs (except Harpymimus; unreported in deinocheirids), but like carcharodontosaurids, therizinosauroids, some oviraptorids and velociraptorines. The proximal outline of metatarsal III would at first glance appear to be the strangest thing about this material, being reconstructed as strictly dorsoventrally oval unlike all(?) other theropods. Tilting it and adding a posterior tapered tip results in a close match to Majungasaurus however (see figure below). If it is an unreduced proximal metatarsal III, tyrannosauroids, most ornithomimosaurs, alvarezsauroids and pennaraptorans would be excluded. Proximal phalanx II-2 lacks a proximoventral heel, so is not from a deinonychosaur. The pedal phalanges are too elongate to be therizinosauroid, and the pedal ungual is too broad. Phalanges are not as dorsoventrally compressed as Mapusaurus, and as noted above they lack the ventrolateral shelves found in ornithomimosaurs. Abelisaurid phalanges seem similar however. I wonder if we have a case like Camarillasaurus or probably Dandakosaurus involving misidentified elements making the specimen seem stranger than it really was, with so many edges of supposed metatarsal III dotted to indicate incompleteness that it could actually be metatarsal II or IV. Certainly nothing connects this specimen with Deinocheirus. As per the numerous errors illustrated by Hartman et al. (2019) nobody should trust Choiniere et al.'s scorings in any case. The Lori matrix recovers "Paraxenisaurus" as a ceratosaur closest to Aucasaurus as far as taxa with well preserved feet are concerned, but also doesn't include characters particular to ceratosaurs and isn't great with pedal characters in general. So I would place the specimen as Neotheropoda incertae sedis (or even indet.) pending a better description of the tarsus and of the real bone surfaces on supposed proximal metatarsal III.
Pes of "Paraxenisaurus normalensis" holotype (center) in dorsal view compared to Majungasaurus crenatissimus composite (left) and Deinocheirus mirificus referred specimen IGM 100/127 (right). Colored proximal view of "Paraxenisaurus" is after Serrano-Brañas et al., with reoriented metatarsal III as per my interpretation shown above that. Note "Paraxenisaurus" elements were scaled using their scale bars, whereas scaling to listed measurements results in different proportions, so those should be seen as approximate. "Paraxenisaurus" after Serrano-Brañas et al. (2020), Majungasaurus after Carrano (2007) and Deinocheirus after Lee et al. (2014).
References- Osmólska, Roniewicz and Barsbold, 1972. A new dinosaur, Gallimimus bullatus n. gen., n. sp. (Ornithomimidae) from the Upper Cretaceous of Mongolia. Palaeontologica Polonica. 27, 103-143.
Carrano and Hutchinson, 2002. Pelvic and hindlimb musculature of Tyrannosaurus rex (Dinosauria: Theropoda). Journal of Morphology. 253, 207-228.
Brochu, 2003. Osteology of Tyrannosaurus rex: Insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Society of Vertebrate Paleontology Memoir. 7, 138 pp.
Middleton, 2003. Morphology, evolution, and function of the avian hallux. PhD thesis, Brown University. 147 pp.
Carrano, 2007. The appendicular skeleton of Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. In Sampson and Krause (eds.). Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. SVP Memoir 8, 164-179.
Lee, Barsbold, Currie, Kobayashi, Lee, Godefroit, Escuillie and Tsogtbaatar, 2014. Resolving the long-standing enigmas of a giant ornithomimosaur Deinocheirus mirificus. Nature. 515, 257-260.
Kobayashi and Barsbold, 2005. Reexamination of a primitive ornithomimosaur, Garudimimus brevipes Barsbold, 1981 (Dinosauria: Theropoda), from the Late Cretaceous of Mongolia. Canadian Journal of Earth Sciences. 42(9), 1501-1521.
Claessens and Loewen, 2016 (online 2015). A redescription of Ornithomimus velox Marsh, 1890 (Dinosauria, Theropoda). Journal of Vertebrate Paleontology. 36(1), e1034593.
Hattori, 2016. Evolution of the hallux in non-avian theropod dinosaurs. Journal of Vertebrate Paleontology. 36(4), e1116995.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247.
Cau, 2020 online. http://theropoda.blogspot.com/2020/04/paraxenisaurus-un-deinocheiride.html
Serrano-Brañas, Espinosa-Chávez, Maccracken, Gutiérrez-Blando, de León-Dávila and Ventura, 2020. Paraxenisaurus normalensis, a large deinocheirid ornithomimosaur from the Cerro del Pueblo Formation (Upper Cretaceous), Coahuila, Mexico. Journal of South American Earth Sciences. 101, 102610.
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Is Falcatakely a bird?
So this week we got the description of the new Maevarano skull Falcatakely forsterae (O'Connor et al., 2020). It's a pseudo-toucan, with a long, tall snout formed mostly by the maxilla unlike modern birds. O'Connor et al. recover it as an enantiornithine using Brusatte et al.'s TWiG analysis and O'Connor's bird analysis. Our problem is that we only have the anteroventral skull preserved, so no braincase, mandible or postcrania. And being a Maastrichtian deposit in Africa, we don't have the most detailed coelurosaur record. Add in the fact beaks are known to evolve fast on islands (as Madagascar was even back then) and we have a potential problem on our hands.
Follow your nose to the exciting topic of African coelurosaur diversity. Reconstructed holotype skull of Falcatakely forsterae (UA 10015) (after O'Connor et al., 2020).
The Lori analysis places Falcatakely in two potential positions- a therizinosaurid or an omnivoropterygid. The former doesn't make much sense biostratigraphically, but there is a Maevarano synsacrum of the correct size (FMNH PA 741) that was claimed to share characters with Sapeornis by O'Connor and Forster (2010). So there's a possibility. Forcing Falcatakely to be an enantiornithine as in O'Connor et al.'s analyses requires six more steps. Most of the discordant characters relate to the beak, but the wide laterotemporal fenestra would be odd in an enantiornithine. While I was writing this, Andrea Cau published a post on this topic and reported that he recovered Falcatakely as a noasaurid, which would be quite the phylogenetic jump, but it only takes three more steps in the Lori matrix, so is more parsimonious than the enantiornithine option. It falls out as an elaphrosaurine, so could relate to e.g. Afromimus. The non-beak contradictory characters here include a lack of antorbital fossa lacrimal foramen, long posterior lacrimal process and triradiate palatine, which seem more convincing to me. Additional evidence against these latter two positions is the absence of small ceratosaur (Masiaksaurus is twice as big) or large enantiornithine (Maevarano elements are much smaller) postcrania. Andrea reported (translated) "It takes 6 further steps to place it in Coelurosauria, and in that case it is a basal dromaeosaurid: interesting in that regard to note that Rahonavis , known from the same Formation, has also been hypothesized to be a basal dromaeosaurid. Can we rule out that Falcatakely is the (still unknown) skull of Rahonavis? The estimated dimensions of the two animals coincide." Forcing Falcatakely to be Rahonavis only requires one more step, which is pretty impressive. In their (amazing) osteology, Forster et al. (2020) refer an isolated dentary "found near the Rahonavis holotype (its precise location was not recorded during excavation)" which does not match Falcatakely's upper jaw, being upcurved and extensively toothed. But it is similar to other unenlagiines like Buitreraptor and Austroraptor. So much as we have two synsacrum types at this size, unenlagiine-like Rahonavis and Sapeornis-like, we have two cranial types, unenlagiine-like and Falcatakely which is Sapeornis-like in the combination of reduced maxillary dentition, triradiate palatine, modified/reduced antorbital fossa, anteriorly limited naris and strong postorbital-jugal articulation.
Referred dentary of Rahonavis ostromi (FMNH PA 740) as a transparent CT reconstruction (after Forster et al., 2020).
Thus my best guess is that Falcatakely is a basal avialan belonging to the same taxon as FMNH PA 741. But this comes with a huge chunk of salt as it is so far removed temporally and geographically from potential comparable sister taxa. Which is actually a common problem with this part of the tree, as shown by Balaur (= Elopteryx?), Hesperonychus, Imperobator and even Rahonavis itself. We compare these Late Cretaceous taxa to our far more complete Early Cretaceous Jehol record and say Hesperonychus is sorta like Microraptor, Falcatakely is kind of like Sapeornis and Balaur is Jeholornis-grade, but North America, Africa and Europe had their own avialan fauna for 70 million years before them that we're basically unaware of. If the only alvarezsauroid we had was Mononykus' holotype, could we place it correctly as a basal maniraptoran? If the only oviraptorosaur we had was Citipati's skull, would we recover that correctly as the sister taxon of Paraves? I think that's the position we find ourselves in with Falcatakely, and that future discoveries of African small theropods will lead to new interpretations.
References- O'Connor and Forster, 2010. A Late Cretaceous (Maastrichtian) avifauna from the Maevarano Formation, Madagascar. Journal of Vertebrate Paleontology. 30(4), 1178-1201.
Cau, 2020 online. theropoda.blogspot.com/2020/11/falcatakely-eterodossia-e-pluralismo.html
Forster, O'Connor, Chiappe and Turner, 2020. The osteology of the Late Cretaceous paravian Rahonavis ostromi from Madagascar. Palaeontologia Electronica. 23(2):a31.
O'Connor, Turner, Groenke, Felice, Rogers, Krause and Rahantarisoa, 2020. Late Cretaceous bird from Madagascar reveals unique development of beaks. Nature. DOI: 10.1038/s41586-020-2945-x
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Antarctic Ichthyornis solved
So I've been doing some major updates to the Database for what will probably be a New Years upload, including the ornithuromorph section. One rather sad entry as it currently stands is the Antarctic Ichthyornis-
I? sp. (Zinsmeister, 1985)
Late Cretaceous
Seymour Island, Antarctica
Reference- Zinsmeister, 1985. 1985 Seymour Island expedition. Antarctic Journal of U.S. 20, 41-42.
Now with Googling I found the original paper online, which allowed only a bit of improvement-
I? sp. (Zinsmeister, 1985)
Late Maastrictian, Late Cretaceous
Lopez de Bertodano Formation, Seymour Island, Antarctica
Material- several elements
Comments- Zinsmeister (1985) states "several small bones tentatively identified as belonging to the Cretaceous bird Ichthyornis were discovered in the upper Cretaceous Lopez de Bertodano formation."
Reference- Zinsmeister, 1985. 1985 Seymour Island expedition. Antarctic Journal of U.S. 20, 41-42.
So I saw that Zinsmeister worked with Chatterjee in the 80s, who found the Polarornis holotype in the same place two years before that. I emailed Chatterjee about it, who replied-
"It was misidentified in the field. These were some shark teeth."
Mystery solved! But can we do better? Here's an Ichthyornis tooth-
Right eleventh dentary tooth of Ichthyornis dispar (YPM 1450) (after Field et al., 2018).
And here's the array of shark teeth from the Lopez de Bertodano Formation of Seymour Island (from a January 2011 expedition). Can we find any easily confusable matches?
Chondrichthyan teeth from the Lopez de Bertodano Formation (scale 10 mm) (after Otero et al., 2014).
I think the circled 16 and 17 are pretty decent matches for a field identification, though much larger if compared directly. Figures 6-17 are all identified as Odontaspidae indet., which covers any morphology similar to Ichthyornis. Add in the fact that they were by far the most abundant teeth recovered (8 samples versus 1-3 for the other taxa), and I think we have a nice solution on our hands.
I wonder how many other weird records are out there that are based on initial misidentification but stay in the literature because nobody ever publishes a correction?
References- Otero, Gutstein, Vargas, Rubilar-Rogers, Yury-Yañez, Bastías and Ramírez, 2014. New chondrichthyans from the Upper Cretaceous (Campanian-Maastrichtian) of Seymour and James Ross islands, Antarctica. Journal of Paleontology. 88(3), 411-420.
Field, Hanson, Burnham, Wilson, Super, Ehret, Ebersole and Bhullar, 2018. Complete Ichthyornis skull illuminates mosaic assembly of the avian head. Nature. 557, 96-100.
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"Megalosaurus" cloacinus and more - September 2021 Database Update
Hi everyone. I realize it's been ten months since the last post, and that's because I've been prioritizing updating the Database over writing blogs. As a compromise of sorts and to not force people to constantly check the Database updates page, I decided to try out posting when I update including features that could have made it into their own blog post.
One thing I've been doing is working my way through Skawiński et al.'s (2017) paper on Polish Triassic dinosaur reports, which in addition to unnamed fragments, also led to the creation of entries for two supposed Megalosaurus species. silesiacus is a generic carnivorous archosauriform tooth too early to be dinosaurian, while cloacinus has been used for basically every carnivorous archosaur tooth from Rhaetian beds of Germany. The interesting thing about the latter is that workers apparently forgot that it was based on lost teeth described by Quenstedt, not the SMNS tooth figured 47 years later by Huene.
"Zanclodon" silesiacus Jaekel, 1910
= Megalosaurus silesiacus (Jaekel, 1910) Kuhn, 1965
Early Anisian, Middle Triassic
Lower Gogolin Formation, Lower Muschelkalk, Poland
Holotype- (University of Griefswalden/Göttinger coll.; lost?) tooth (24x12x5 mm)
Referred- ?(Geological Museum of the Polish Geological Institute-National Research Institute coll.) tooth (Skawiński, Ziegler, Czepiński, Szermański, Tałanda, Surmik and Niedźwiedzki, 2017)
?(Silesian University of Technology, Faculty of Mining and Geology coll.) tooth (37 mm) (Surmik and Brachaniec, 2013)
Comments- Jaekel (1910) noted (translated) "a dinosaur tooth from the lower shell limestone of Upper Silesia, which would probably be the oldest known dinosaur tooth to date. It comes from the Chorzov strata of the lower shell limestone of Gogolin, Upper Silesia, and came to me through the kindness of engineer Fedder in Opole. The crown shown is 24 mm high, 12 mm wide and 5 mm thick, so it is quite strongly compressed and slightly curved backwards. Its edge is extremely finely serrated (Fig. 16 A). I call the form, which for the time being cannot be specified generically, Zanclodon silesiacus. The only difference between [phytosaur Mesorhinosuchus] and this tooth form lies in the fact that the former is somewhat thicker, somewhat less bent back, and that no notch can be detected on the edge." He referred it to Megalosauridae, and Kuhn (1965) later referred it to the genus Megalosaurus. Carrano et al. (2012) correctly noted "could be considered as Theropoda indet., but we cannot rule out the possibility that it represents a 'rauisuchian' archosaur." Surmik and Brachaniec (2013) describe a tooth from Gogolin Quarry in which "a poor state of preservation makes it impossible to identification of the presence of edge serration, however it still shows a slightly curvature and specific both sides flattening" and identify it as seemingly archosaurian. Skawiński et al. (2017) listed this and another tooth labeled as Megalosaurus silesiacus as other material of Zanclodon silesiacus. The latter tooth is stated to be serrated mesially and distally with a density of 12 per 5 mm. They describe the holotype tooth as "Probably lost" and "lost", and place all three teeth as Archosauromorpha indet.. They are more specifically referred to the Teyujagua plus archosauriform clade here given the recurvature and small serrations, as authors from Kuhn onward have noted plesiomorphic theropod teeth are difficult to distinguish from several clades of archosauriforms (e.g. erythrosuchids, euparkeriids) known from the Anisian. The age is far too early for Megalosaurus or another neotheropod, and the presence of serrations is unlike Zanclodon, so neither genus is appropriate. It should also be noted the three Gogolin teeth differ in shape with the Silesian University specimen less recurved and less tapered than the other two, while the Polish Geological Institute specimen is shorter than the holotype and less concave distally. This could be positional variation, but given the lack of proposed synapomorphies could easily represent multiple taxa.
References- Jaekel, 1910. Ueber einen neuen Belodonten aus dem Buntsandstein von Bernburg. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin. 5, 197-229.
Kuhn, 1965. Fossilium Catalogus 1: Animalia. Pars 109: Saurischia. Ysel Press. 94 pp.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Surmik and Brachaniec, 2013. The large superpredators' teeth from Middle Triassic of Poland. Contemporary Trends in Geoscience. 2, 91-94.
Skawiński, Ziegler, Czepiński, Szermański, Tałanda, Surmik and Niedźwiedzki, 2017 (online 2016). A re-evaluation of the historical 'dinosaur' remains from the Middle-Upper Triassic of Poland. Historical Biology. 29(4), 442-472.
Holotype tooth of "Zanclodon" silesiacus (University of Griefswalden/Göttinger coll.; lost?) in labial (A), basal section (B) and more apical section (C) (after Jaekel, 1910).
"Megalosaurus" cloacinus Quenstedt, 1858
= Plateosaurus cloacinus (Quenstedt, 1858) Huene, 1905
= Gresslyosaurus cloacinus (Quenstedt, 1858) Huene, 1932
= Pachysaurus cloacinus (Quenstedt, 1858) Huene, 1932
Rhaetian, Late Triassic
Exter Formation, Germany
Syntypes- (lost) two teeth
Referred- ?(GPIT and SMNS coll.) many teeth (Huene, 1905)
?(SMNS 52457) tooth (~25x11x? mm) (Huene, 1905)
?(SMNS coll.) teeth (Roemer, 1870)
? seven teeth (Miller Endlich, 1870)
Norian-Rhaetian?, Late Triassic
'Lisów Breccia', Poland
?(University of Wroclaw coll.; lost) two teeth (Roemer, 1870)
Early Hettangian, Early Jurassic
Calcaire de Valognes, Manche, France
?(University of Caen coll.; destroyed) tooth (Rioult, 1978)
Comments- Quenstedt (1858) originally described (translated) "barb-shaped teeth, which are sharp and finely serrated on the concave side, but rounded and smooth on the convex side" with a large mesioapically placed wear facet that makes that edge look straight in side view. He also figures a smaller tooth which has mesial serrations apically that transition to a rounded edge basally. These teeth do not share any obvious synapomorphies and differ in elongation (height/FABL ~300% vs. 138%) and transverse thickness (42% vs. 75% of FABL), so may not belong to the same taxon. Miller Endlich (1870) figured seven teeth from the type locality, stating (translated) they "are mostly flat teeth, slightly curved on one side, with fine serrations on the sharp inner edge. The convex side, the back, does not seem to be serrated, but it is not certain." The figured teeth show a wide range of variation, with figure 13 in particular being stout and unrecurved with large serrations, similar to the Lucianosaurus paratype and similarly referrable to Archosauromorpha incertae sedis. The other teeth have small serrations, with 14 and 18 being straight and 15-17 and 19 being recurved, with 14, 18 and 15 being progressively more transversely compressed. As with the syntypes, these exhibit variation which could be positional or interspecific, and share no obvious characters that connect them to each other or the syntypes. Roemer (1870) wrote (translated) "In the Stuttgart Museum I saw teeth from the bone breccia of Bebenhausen near Tubingen, which show the same fine serration of the side edges as the teeth described by Quenstedt, but are not curved in a sickle shape, but are straight. It is very likely that these latter teeth belong to the same dinosaur as the crooked teeth. With these straight teeth from Bebenhausen, the tooth shown in FIGS. 4 and 5 from the Lisów Breccia from Lubsza near Woźniki completely coincides. The double-edged tooth, which is very delicately and regularly notched at the edges, shows a more strongly curved (outer) and a less curved (inner) surface, both of which are smooth except for a very fine, irregular wrinkle. There is also a much smaller tooth of the same type from the same location." The straight Bebenhausen teeth sound similar to Miller Endlich's figures 14 and 18, although the illustrated straight tooth from Lubsza differs from these in having an increased amount of mesiodistal expansion basally. The Lubsza tooth also has this marked basal expansion labiolingually, and both types of root expansion are atypical of dinosaurs, suggesting this is some other type of vertebrate. Dzik and Sulej (2007) suggested it "may have belonged to a phytosaur" without evidence but Skawiński et al. (2017) stated "phytosaur fossils have not been found in the upper Keuper strata in Silesia" and instead placed it in Archosauromorpha indet.. While this could merely mean phytosaurs were rare in that strata, phytosaur teeth don't seem to have expanded roots either (e.g. Nicrosaurus), and it could even be a fish tooth which often have these types of root expansion. Huene (1905) listed the species as "Plateosaurus" cloacinus within Theropoda, stating it includes Rhaetian dentary "Zanclodon cambrensis". In 1908 he places it in Plateosauridae within Theropoda and states (translated) "The originals can no longer be found. The Tübingen collection still has several teeth from Bebenhausen and Schloßlesmuehle, which can be reconciled well with [Quenstedt's] fig. 12 (l. c.), but are larger. The serrations are coarse and short, the mesial carina does not extend all the way to the base." He illustrated a tooth in figure 274 as "From the Rhaetian Bonebed of Bebenhausen near Tübingen. Tooth in nat. Size. The tip is missing. Original in the natural history cabinet in Stuttgart." Regarding cambrensis, Huene states "The teeth have the greatest resemblance to Plateosaurus cloacinus both in the whole shape and in the serrations. Whether it is really the same or just a very similar species, of course, cannot be decided with certainty given the scanty material", which is not explicit enough to evaluate given published details. Huene later (1932) assigns cloacinus to Teratosauridae within Carnosauria, listed as both Pachysaurus cloacinus (pg. 6) and Gresslyosaurus cloacinus (pg. 72, 114). Steel (1970) calls it Gresslyosaurus cloacinus within Plateosauridae. Buffetaut et al. (1991) mentions "A tooth referred to Megalosaurus cloacinusQuenstedt, from the Lower Hettangian of the Calcaire de Valognes at Valognes (Manche), [which] has been mentioned by Rioult (1978a) as having been destroyed by an air raid on the University of Caen in 1944." Without additional details, it can only be said that the timing suggests a neotheropod. Carrano et al. (2012) incorrectly claimed SMNS 52457, apparently the tooth in Huene's (1908) figure 274, is "the holotype and only specimen" of cloacinus, when Huene stated it was only one of "Many teeth ... in the stone quarries of the Schoenbuch (e.g. Bebenhausen, Schloesslesmuehle), Wuerttemberg; in the university collection in Tubingen and in the natural history cabinet in Stuttgart", and that Quenstedt's originals were lost. SMNS 52457 could be made into a neotype, but this must be done explicitly (ICZN Article 75.3) and so has not been
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paleolibrary - Free ebook download as PDF File (.pdf), Text File (.txt) or read book online for free.
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An ankylosaurian dinosaur from the Cenomanian Dunvegan Formation of northeastern British Columbia, Canada
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Abstract. Fragmentary but associated dinosaur bones collected in 1930 from
the Pine River of northeastern British Columbia are identified here as
originating from an ankylosaur. The specimen represents only the second
occurrence of dinosaur skeletal material from the Cenomanian Dunvegan
Formation and the first from Dunvegan outcrops in the province of British
Columbia. Nodosaurid ankylosaur footprints are common ichnofossils in the
formation, but the skeletal material described here is too fragmentary to
confidently assign to either a nodosaurid or ankylosaurid ankylosaur. The
Cenomanian is a time of major terrestrial faunal transitions in North
America, but many localities of this age are located in the southern United
States; the discovery of skeletal fossils from the Pine River demonstrates
the potential for the Dunvegan Formation to produce terrestrial vertebrate
fossils that may provide important new data on this significant transitional
period during the Cretaceous.
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https://fr.copernicus.org/articles/23/179/2020/
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Terrestrial vertebrate fossils have rarely been reported from the Cenomanian-aged Dunvegan Formation (Burns and Vavrek, 2014; Vavrek et al., 2014), which crops out in northeastern British Columbia and northwestern Alberta. Here we describe dinosaur bones recovered from outcrops of the formation along the Pine River in northeastern British Columbia (Fig. 1a). The specimen (CMN 59667), a block of sandstone with associated ribs and vertebrae, was collected in 1930 from the Pine River. To our knowledge this is the first dinosaur skeletal fossil discovered in the province, predating by more than 40 years the discovery of Ferrisaurus sustutensis (Arbour and Evans, 2019), first reported by Arbour and Graves (2008).
Although bony fish and sharks have been recorded from the Dunvegan Formation in Alberta (Hay et al., 2007; Cook et al., 2008; Vavrek et al., 2014), the terrestrial vertebrate body fossil record of the formation has thus far been limited to a few ankylosaur ossicles from Alberta (Burns and Vavrek, 2014). Dinosaur bones found in the Tumbler Ridge area, first described as originating from the Dunvegan Formation (McCrea, 2003), were later reinterpreted as originating from the Turonian Kaskapau Formation (McCrea and Buckley, 2004; Rylaarsdam et al., 2006). Therefore, well-preserved dinosaur fossils, particularly from an associated skeleton, would contribute significantly to understanding the fauna of the Dunvegan Formation and preservational potential within this unit.
CMNÂ 59667 is noteworthy for being one of very few Cenomanian-aged dinosaur skeletal fossils from Canada (Brown et al., 2015). Terrestrial Cenomanian assemblages are rare in North America but capture an important time which preserves a distinct and transitional assemblage between characteristic faunas of the Early and latest Late Cretaceous. The nearest comparative material of this age is derived from Dunvegan Formation outcrops in Alberta, the Cedar Mountain Formation of Utah (Avrahami et al., 2018; Tucker et al., 2020), the Woodbine (Noto et al., 2012) and Paw Paw formations of Texas, the Frontier Formation of Wyoming (Lull, 1921), the Wayan Formation of Idaho (Krumenacker et al., 2019), and the Dakota Formation of the western US (Eaton, 1960; Eaton, 1993), all of which are either areas that are being actively collected and researched or not well explored or documented. The Dunvegan Formation is the furthest north of the known Cenomanian-aged vertebrate fossil-bearing terrestrial units in North America and is therefore of particular interest for testing proposed latitudinal faunal provinciality in the early Late Cretaceous (e.g. Lehman, 2001; Sampson et al., 2010) and understanding its origin.
CMN 59667 was collected by Merton Yarwood Williams in 1930; this was likely during a survey he headed, the Non-Metallic Mineral Investigation on the Pacific Great Eastern Railway Survey of Resources of the Peace River, West Cariboo and West Lillooet Blocks, B.C., during the summers of 1929 and 1930 (Hives, 1988). Notes associated with CMN 59667 indicate it was collected â1 mile below junction of Murray and Pine Rivers (Peace River district)â. Charles M. Sternberg later that year identified the block as containing an anterior dorsal vertebra and rib fragment of an ornithischian resembling Camptosaurus, according to the label associated with the specimen.
The early to middle Cenomanian Dunvegan Formation crops out in this area. The Dunvegan Formation represents a major deltaic complex deposited over the span of about 2Â million years (Plint, 2000). Characterized by a succession of alluvial and shallow marine sandstones, siltstones, and shales, it has an interfingering and diachronous relationship with the underlying marine Shaftesbury Formation and overlying marine Kaskapau Formation (Bhattacharya, 1994; Plint, 2000).
We undertook a reconnaissance field trip to the Pine RiverâMurray River area in May 2019 to document the lithology and fossils of the original collection area. We were able to access the west bank of the Pine River on foot 1.3âkm downstream from its junction with the Murray River, very close to the locality described in the original notes associated with the specimen. The east bank of the river exposes a large section of Dunvegan Formation approximately 22âm thick (Fig. 1e). At the base of the cliff we observed a distinctive orange-stained palaeosol overlain by two thin dark-grey coal seams or palaeosols. Above these layers, there were seven packages of fine-grained sediments overlain by a more resistant layer of sandstone. The west side of the river lacks cliff-forming outcrop, but small outcrops are present as well as abundant loose pieces of sandstone.
Although we did not encounter any additional vertebrate bone, we observed current ripple marks, siderite nodules, and abundant ichnofossils, including vertically oriented burrows, shallow surface burrows or feeding traces, and the dinosaur ichnotaxon Tetrapodosaurus (RBCM P1473), made by a nodosaurid ankylosaur (Fig. 1b). Previously, Storer (1975) described a slab with tridactyl dinosaur footprints discovered on the north bank of the Pine River âat the site of the present bridge at East Pineâ, very close to the reported collection locality of CMN 59667. These were identified as Colombosauripus ungulatus and interpreted as belonging to a small coelurosaur similar to an ornithomimid. Terrestrial vertebrate footprints and trackways are widespread throughout the Dunvegan formation (both geographically and stratigraphically) and include tracks made by turtles, crocodilians, nodosaurs (Tetrapodosaurus), large ornithopods, small theropods (cf. Irenichnites; Columbosauripus), medium-sized theropods (cf. Magnoavipes), and birds; tracks attributed to large theropods have yet to be reported (McCrea et al., 2014). Plant remains in this area included leaf compressions (including the representative samples we collected, RBCM P1028-P1032), carbonized leaves and cones, and wood impressions (Fig. 1c, d, f). These sedimentological and palaeontological features are consistent with the upper two-thirds of allomember E of the Dunvegan Formation, dominated by lakes, crevasse deltas, and poorly drained palaeosols (Lumsdon-West and Plint, 2005). This region would have been located at about 60ââN during the Cenomanian (van Hinsbergen et al., 2015).
Although the Early to mid-Cretaceous dinosaur fossil record of western Canada is currently less well documented (Brown et al., 2015) in comparison to the southwestern United States, recent discoveries reveal the potential for much more information to be gleaned from this time period. Notably, the Early and mid-Cretaceous dinosaur skeletal record of western Canada is almost entirely represented by ankylosaurs, including osteoderms from the Berriasian-aged Pocaterra Creek Member of the Cadomin Formation in the Rocky Mountains of southwest Alberta (Nagesan et al., 2020), the exceptionally preserved Borealopelta from the Aptian Clearwater Formation (Brown et al., 2017), and isolated ossicles from the Dunvegan Formation of Alberta (Burns and Vavrek, 2014).
Ankylosaur skeletal fossils are widespread across North America during the CampanianâMaastrichtian, but they are never common components of the fossil record for a given formation. Even in the well-sampled Dinosaur Park Formation of southern Alberta, ankylosaurs make up less than 15â% of the skeletal record (Brinkman et al., 1998, 2005). Although similar statistical analyses or censuses have yet to be undertaken for many Early and mid-Cretaceous formations in North America, ankylosaurs appear to make up a greater proportion of the ornithischian fauna compared to the CampanianâMaastrichtian in several localities (e.g. Britt et al., 2009). Most Early and mid-Cretaceous Laramidian ankylosaurs were nodosaurids (Carpenter and Kirkland, 1998; Arbour et al., 2016; Brown et al., 2017), with only a single ankylosaurid, Cedarpelta bilbeyhallo rum, known from the Cenomanian-aged Mussentuchit Member of the Cedar Mountain Formation (Carpenter et al., 2001, 2008).
Ankylosaur footprints are also rare during the CampanianâMaastrichtian of North America, with only a few potential records from the Blackhawk Formation of Utah (McCrea et al., 2001), the Wapiti Formation of northern Alberta (Fanti et al., 2013), and the Chignik Formation of Alaska (Fiorillo et al., 2019). In contrast, the ichnotaxon Tetrapodosaurus, generally considered to have been made by tetradactyl ankylosaurs (McCrea et al., 2001, 2014), is widespread in North America during the Early and mid-Cretaceous, with records from the Nanushuk and Chandler formations of Alaska (McCrea et al., 2001; May and Druckenmiller, 2009); the Gates and Dunvegan formations of Alberta (McCrea et al., 2001); the Gorman Creek, Gething, Dunvegan, Gates, and Kaskapau formations of British Columbia (McCrea et al., 2001, 2014); the Dakota Group of Colorado (Lockley and Gierlinski, 2014); the Naturita and Cedar Mountain formations of Utah (Lockley et al., 1999, 2018); and the Ross River Block in the Yukon (Gangloff et al., 2004). Given the rarity of ankylosaurian footprints in the CampanianâMaastrichtian of North America but their abundance in the Early and mid-Cretaceous (including the Dunvegan Formation), it is notable that the only dinosaur skeletal material thus far described from the Dunvegan Formation, both recorded here and by Burns and Vavrek (2014), is ankylosaurian in nature.
The ichnofauna of the Dunvegan Formation is dominated by Tetrapodosaurus (McCrea et al., 2014), but little is known about its presumed trackmaker because skeletal material from this formation is extremely rare. Both nodosaurid and ankylosaurid ankylosaurs have five functional manual digits, but the number of pedal digits differs between these clades. Four pedal digits are present in the nodosaurids Niobrarasaurus (MU 650âVP; Carpenter et al., 1995), Nodosaurus (YPM 1815; Lull, 1921; Carpenter and Kirkland, 1998), and Sauropelta (AMNH 3032; Ostrom, 1970). In contrast, only three pedal digits are present in the ankylosaurids Anodontosaurus (AMNH 5266; Coombs, 1986), Dyoplosaurus (ROM 784; Arbour et al., 2009), Liaoningosaurus (IVPP V12560; Xu et al., 2001; XHPM-1206; Ji et al., 2016), Pinacosaurus (multiple individuals from Alag Teeg, Currie et al., 2011), âZhejiangosaurusâ (ZMNH M8718; Lü et al., 2007), and the indeterminate articulated Mongolian ankylosaurids MPC 100/1305D (Carpenter et al., 2011) and PIN 614 (Maleev, 1954); Maleev (1956) reports Talarurus PIN 557-3 as having four digits in the pes, but this specimen is composed of multiple individuals, and it is unclear how the pedal elements were associated in situ. No tetradactyl ankylosaurids are known, and as such the tetradactyl prints represented by Tetrapodosaurus are unlikely to have been made by an ankylosaurid ankylosaur.
Based on the prevalence of Tetrapodosaurus in the Dunvegan Formation (McCrea et al., 2014), including at the CMN 59667 locality (e.g. RBCM P1473), and the fact that its trackmaker was probably not an ankylosaurid ankylosaur, we might expect ankylosaur skeletal material from the Dunvegan Formation to derive from a nodosaurid ankylosaur. However, the preserved transverse process of CMN 59667 has a diapophyseal sulcus on its ventral side, a feature that seems to be present in ankylosaurids but absent in nodosaurids. The distribution of the diapophyseal sulcus within ankylosaurs requires additional specimens and investigation, and CMN 59667 is too fragmentary to confidently assess its affinities within Ankylosauria. Nevertheless, the presence of both nodosaurids and ankylosaurids further south in North America during the Cenomanian means that an ankylosaurid identification for this specimen cannot be ruled out, despite the prevalence of nodosaurid footprints in the Dunvegan Formation. Burns and Vavrek (2014) were similarly cautious in their taxonomic appraisal of isolated ankylosaur ossicles from Dunvegan outcrops in Alberta, noting that the morphology and histology of such small elements cannot be as easily mapped to higher-level clades in comparison to larger ankylosaur osteoderms. We note, however, that the ossicles reported by Burns and Vavrek (2014) are extremely small (â¼1âmm diameter), a size that is much more consistent with small interstitial osteoderms from ankylosaurid ankylosaurs such as Jinyunpelta (Zheng et al., 2018), Scolosaurus (NHMUK R5161; Nopcsa, 1928), and Zuul (ROM 75860; Arbour and Evans, 2017). Smaller interstitial osteoderms in nodosaurids are sometimes absent because of closer osteoderm packing (e.g. Borealopelta; Brown, 2017; Brown et al., 2017) or are >8âmm in diameter, such as Sauropelta (AMNH 3032) and Panoplosaurus (CMN 2759).
The Cenomanian is a key stratigraphic interval for interpreting the palaeo-biogeographic and evolutionary history of ankylosaurid ankylosaurs. The earliest records of the characteristic tail club occur in Albian to Cenomanian-aged formations in Asia (Arbour and Currie, 2015; Zheng et al., 2018). Ankylosaurids are also first documented in North America during this time period in the Mussentuchit Member of the Cedar Mountain Formation but then disappear from the North American fossil record until the Santonian (Baszio, 1997; Eaton et al., 1999; Parrish, 1999), possibly indicating a response to high sea levels in Laramidia during the early Late Cretaceous (Arbour et al., 2016), although poor sampling in the TuronianâSantonian cannot be ruled out (e.g. Upchurch et al., 2011). In contrast, nodosaurids have a nearly continuous fossil record throughout the entire Cretaceous of North America (Arbour et al., 2016).
CMN 59667 represents the first associated skeletal material from a dinosaur recovered from the Dunvegan Formation. This new, well-preserved specimen therefore shows that this important taphonomic mode (articulated and associated skeletons, modes A1 and A2 of Eberth and Currie, 2005) can occur in the unit and raises the possibility that continued prospecting efforts could yield substantially more complete dinosaur material from this poorly known time interval in the Cretaceous fossil record of Canada. The presence or absence of ankylosaurids in the Dunvegan Formation can only be determined through the discovery of additional, more diagnostic material, but the morphology of the currently known ankylosaur skeletal remains is tantalizing potential evidence for Cenomanian-aged ankylosaurids in Canada. Similarly, the discovery of unequivocal, diagnostic nodosaurid remains would provide the opportunity to link a trackmaker to the abundant Tetrapodosaurus trackways known from this formation.
Arbour, V. M. and Currie, P. J.: Euoplocephalus tutus and the diversity of ankylosaurid dinosaurs in the Late Cretaceous of Alberta, Canada, and Montana, USA, PLoS ONE, 8, e62421, https://doi.org/10.1371/journal.pone.0062421, 2013.â
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Carpenter, K., Hayashi, S., Kobayashi, Y., MaryaÅska, T., Barsbold, R., Sato, K., and Obata, I.: Saichania chulsanensis (Ornithischia, Ankylosauridae) from the Upper Cretaceous of Mongolia, Palaeontogr. Abt. A, 293, 1â161, 2011.â
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Eberth, D. A. and Currie, P. J.: Vertebrate taphonomy and taphonomic modes, in: Dinosaur Provincial Park: a spectacular ancient ecosystem revealed, edited by: Currie, P. J. and Koppelhus, E. B., Indiana University Press, 453â477, 2005.â
Fanti, F., Bell, P. R., and Sissons, R. L.: A diverse, high-latitude ichnofauna from the Late Cretaceous Wapiti Formation, Alberta, Canada, Cret. Res., 41, 256â269, 2013.â
Fiorillo, A. R., Kobayashi, Y., McCarthy, P. J., Tanaka, T., Tykoski, R. S., Lee, Y.-N., Takasaki, R., and Yoshida, J.: Dinosaur ichnology and sedimentology of the Chignik Formation (Upper Cretaceous), Aniakchak Nationa l Monument, southwestern Alaska; further insights on habitat preferences of high-latitude hadrosaurs, PLOS ONE, 14, e0223471, https://doi.org/10.1371/journal.pone.0223471, 2019.â
Gangloff, R. A., May, K. C., and Storer, J. E.: An early Late Cretaceous dinosaur tracksite in central Yukon Territory, Canada, Ichnos, 11, 299â309, 2004.â
Hay, M. J., Cumbaa, S. L., Murray, A. M., and Plint, A. G.: A new paraclupeid fish (Clupeomorpha, Ellimmichthyiformes) from a muddy marine pro-delta environment: middle Cenomanian Dunvegan Formation, Alberta, Canada, Can. J. Earth Sci., 44, 775â790, 2007.â
Hives, C.: M. Y. Williams fonds, University of British Columbia Archives, 50 pp., 1988.â
Ji, Q., Wu, X., Cheng, Y., Ten, F., Wang, X., and Ji, Y.: Fish hunting ankylosaurs (Dinosauria, Ornithischia) from the Cretaceous of China, J. Geol., 40, 183â190, 2016.â
Kirkland, J. I., Alcalá, L., Loewen, M. A., EspÃlez, E., Mampel, L., and Wiersma, J. P.: The basal nodosaurid ankylosaur Europelta carbonensis n. gen., n. sp. from the Lower Cretaceous (Lower Albian) Escucha Formation of northeastern Spain, PLOS ONE, 8, e80405, https://doi.org/10.1371/journal.pone.0080405, 2013.â
Krumenacker, L. J., Varricchio, D. J., Wilson, J. P., Martin, A., and Ferguson, A.: Taphonomy of and new burrows from Oryctodromeus cubicularis, a burrowing neornithischian dinosaur, from the mid-Cretaceous (AlbianâCenomanian) of Idaho and Montana, USA, Palaeogeogr. Palaeocl., 530, 300â311, 2019.â
Lehman, T. M.: Late Cretaceous dinosaur provinciality, in: Mesozoic Vertebrate Life, edited by: Tanke, D. H. and Carpenter, L., Indiana University Press, 310â328, 2001.â
Lockley, M. G. and Gierlinski, G. D.: Notes on a new ankylosaur track from the Dakota Group (Cretaceous) of northern Colorado, New Mexico Mus. Nat. Hist. Sci. Bull., 62, 301â306, 2014.â
Lockley, M., Burton, R., and Grondel, L.: A large assemblage of tetrapod tracks from the Cretaceous Naturita Formation, Cedar Canyon region, southwestern Utah, Cret. Res., 92, 108â121, 2018.â
Lockley, M. G., Kirkland, J. I., DeCourten, F. L., Britt, B. B., and Hasiotis, S. T.: Dinosaur tracks from the Cedar Mountain Formation of eastern Utah: a preliminary report, in: Vertebrate Paleontology in Utah, edited by: Gilette, D. D., Geol. Soc. Am., Miscellaneous Publication 99-1, 253â258, 1999.â
Lü, J., Jin, X., Sheng, Y., Li, Y., Wang, G., and Azuma, Y.: New nodosaurid dinosaur from the Late Cretaceous of Lishui, Zhejiang Province, China, Acta Geol. Sin., 81, 344â350, 2007.â
Lull, R.: The Cretaceous armored dinosaur, Nodosaurus textilis Marsh, Am. J. Sci. Ser., 5, 97â126, 1921.â
Lumsdon-West, M. P. and Plint, A. G.: Changing alluvial style in response to changing accommodation rate in a proximal foreland basin setting: Upper Cretaceous Dunvegan Formation, north-east British Columbia, Canada, in: Fluvial Sedimentology VII, edited by: Blum, M. D., Marriott, S. B., and Leclair, S. F., Int. As. Sed., IAS Special Publication 35, 493â515, 2005.â
Maleev, E. A.: The armored dinosaurs of the Cretaceous period in Mongolia (Family Syrmosauridae), Dokl. Akad. Nauk SSSR, 48, 142â170, 1954 (in Russian; translation by Robert Welch).â
Maleev, E. A.: Armored dinosaurs of the Upper Cretaceous of Mongolia, Family Ankylosauridae, Trudy Paleontol. Inst. Akad. Nauk SSSR, 62, 51â91, 1956 (in Russian; translation by R. Welch).â
May, K. C. and Druckenmiller, P. S.: A dinosaurian ichnofossil assemblage from the Nanushuk Formation, National Petroleum Reserve â Alaska, The Alaska Geological Society 2009 Technical Conference Abstracts Volume, 32 pp., 2009.â
McCrea, R. T. and Buckley, L. G.: Excavating British Columbia's first dinosaurs and other palaeontological projects in the Tumbler Ridge area, Alberta Palaeontological Society Eighth Annual Symposium, Calgary, Alberta, Canada, March 2004, 24â33, 2004.â
McCrea R. T., Lockley, M. G., and Meyer, C. A.: Global distribution of purported ankylosaur track occurrences, in: The Armored Dinosaurs, edited by: Carpenter, K., Indiana University Press, 413â454, 2001.â
McCrea, R. T., Buckley, L. G., Plint, A. G., Currie, P. J., Haggart, J. W., Helm, C. W., and Pemberton, S. G.: A review of vertebrate track-bearing formations from the Mesozoic and earliest Cenozoic of western Canada with a description of a new theropod ichnospecies and reassignment of an avian ichnogenus, New Mexico Mus., Nat. Hist. Sci. Bull., 62, 5â93, 2014.â
Nagesan, R. S., Campbell, J. A., Pardo, J. D., Lennie, K. I., Vavrek, M. J., and Anderson, J. S.: An Early Cretaceous (Berriasian) fossil-bearing locality from the Rocky Mountains of Alberta, yielding the oldest dinosaur skeletal remains from western Canada, Can. J. Earth Sci., 57, 542â552, 2020.â
Nopcsa, F.: Die dinosaurier der Siebenbürgischen Landesteile Ungarns, Mit. A. d. Jahrb. D. kgl. Ungar. Geolog. Reichsanst., 23, 1â26, 1915.â
Nopcsa, B. F.: Palaeontological notes on reptiles, V. On the skull of the Upper Cretaceous dinosaur Euoplocephalus, Geol. Hungarica, Ser. Palaeontol., 1, 1â84, 1928.â
Noto, C. R., Main, D. J., and Drumheller, S. K.: Feeding traces and paleobiology of a Cretaceous (Cenomanian) crocodyliform: example from the Woodbine Formation of Texas, Palaios, 27, 105â115, 2012.â
Osborn, H. F.: Two Lower Cretaceous dinosaurs from Mongolia, Am. Mus. Novit., 95, 1â10, 1923.â
Åsi, A.: Hungarosaurus tormai, a new ankylosaur (Dinosauria) from the Upper Cretaceous of Hungary, J. Vertebr. Paleontol., 25, 370â383, 2005.â
Ostrom, J. H.: Stratigraphy and paleontology of the Cloverly Formation (Lower Cretaceous) of the Bighorn Basin Area, Wyoming and Montana, B. Peabody Mus. Nat. Hist., 35, 12â34, 1970.â
Owen, R.: Report on British fossil reptiles, Reports of the British Association for the Advancement of Science, 11, 60â204, 1842.â
Parrish, J. M.: Dinosaur teeth from the Upper Cretaceous (Turonian-Judithian) of southern Utah, in: Gillette DD (ed) Vertebrate paleontology in Utah, Geol. Soc. Am., 319â321, https://doi.org/10.1371/journal.pone.0012292, 1999.â
Plint, A. G.: Sequence stratigraphy and paleogeography of a Cenomanian deltaic complex: the Dunvegan and lower Kaskapau formations in subsurface and outcrop, Alberta and British Columbia, Canada, B. Can. Petrol. Geol., 48, 43â79, 2000.â
Rylaarsdam, J. R., Varban, B. L., Plint, A. G., Buckley, L. G., and McCrea, R. T.: Middle Turonian dinosaur paleoenvironments in the Upper Cretaceous Kaskapau Formation, northeast British Columbia, Can. J. Earth Sci., 43, 631â652, 2006.â
Sampson, S. D., Loewen, M. A., Farke, A. A., Roberts, E. M., Forster, C. A., Smith, J. A., and Titus, A. L.: New horned dinosaurs from Utah provide evidence fo r intracontinental dinosaur endemism, PLOS ONE, 5, e12292, https://doi.org/10.1016/j.cretres.2020.104384, 2010.â
Seeley, H. G.: On the classification of the fossil animals commonly named Dinosauria, Proc. R. Soc. Lond., 43, 165â171, 1887.â
Storer, J.: Dinosaur tracks, Columbosauripus ungulatus (Saurischia: Coelurosauria), from the Dunvegan Formation (Cenomanian) of northeastern British Columbia, Can. J. Earth Sci., 12, 1805â1807, 1975.â
Tucker, R. T., Zanno, L. E., Huang, H.-Q., and Makovicky, P. J.: A refined temporal framework for newly discovered fossil assemblages of the upper Cedar Mountain Formation (Mussentuchit Member), Mussentuchit Wash, Central Utah. Cret. Res., 110, 104384, https://doi.org/10.1371/journal.pone.012694, 2020.â
Upchurch, P., Mannion, P. D., Benson, R. B., Butler, R. J., and Carrano, M. T.: Geological and anthropogenic controls on the sampling of the terrestrial fossil record: a case study from the Dinosauria, Geol. Soc. SP., 358, 209â240, 2011. â
van Hinsbergen, D. J. J., de Groot, L. V., van Schaik, S. J., Spakman, W., Bijl, P. K., Sluijis, A., Langereis, C. G., and Brinkhuis, H.: A paleolatitude calculator for paleoclimate studies (model version 2.1), PLOS ONE, 10, e0126946, https://doi.org/10.1371/journal.pone.0126946, 2015.â
Vavrek, M. J., Murray, A. M., and Bell, P. R.: An early Late Cretaceous (Cenomanian) sturgeon (Acipenseriformes) from the Dunvegan Formation, northwestern Alberta, Canada, Can. J. Earth Sci., 51, 677â681, 2014.â
Wiersma, J. P. and Irmis, R. B.: A new southern Laramidian ankylosaurid, Akainacephalus johnsoni gen. et sp. nov., from the upper Campanian Kaiparowits Formation of southern Utah, USA, Peer J., 6, e5016, https://doi.org/10.7717/peerj.5016, 2018.â
Xu, X., Wang, X.-L., and You, H.-L.: A juvenile ankylosaur from China, Naturwissenschaften, 88, 297â300, 2001.â
Zheng, W., Jin, X., Azuma, Y., Wang, Q., Miyata, K., and Xu, X.: The most basal ankylosaurine dinosaur from the AlbianâCenomanian of China, with implications for the evolution of the tail club, Sci. Rep. UK., 8, 3711, https://doi.org/10.1016/j.cretres.2020.104384, 2018.â
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Zhejiangosaurus (meaning "Zhejiang lizard") is an extinct genus of nodosaurid dinosaur from the Upper Cretaceous (Cenomanian stage) of Zhejiang, eastern China. It was first named by a group of Chinese and Japanese authors Junchang Lü, Xingsheng Jin, Yiming Sheng and Yihong Li in 2007 and the...
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https://static.wikia.nocookie.net/dinosaurs/images/4/4a/Site-favicon.ico/revision/latest?cb=20240402071814
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Dinopedia
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https://dinopedia.fandom.com/wiki/Zhejiangosaurus
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Zhejiangosaurus (meaning "Zhejiang lizard") is an extinct genus of nodosaurid dinosaur from the Upper Cretaceous (Cenomanian stage) of Zhejiang, eastern China. It was first named by a group of Chinese and Japanese authors Junchang Lü, Xingsheng Jin, Yiming Sheng and Yihong Li in 2007 and the type species is Zhejiangosaurus lishuiensis ("from Lishui", Chinese administrative unit on which the fossil was found).[1]
Description[]
Zhejiangosaurus could grow up to 4.5 m (17 ft) in length and was 1.4 metric tons in weigh.[3]
Material[]
Material for Zhejiangosaurus consists of the holotype, ZNHM M8718, a partial skeleton which has preserved a sacrum with eight vertebrae, a complete right ilium and partial left ilium, a complete right pubis, the proximal end of the right ischium, two complete hindlimbs, fourteen caudal vertebrae, and some unidentified bones. These remains come from Liancheng, in the Chinese administrative unit of Lishui on the province of Zhejiang and they were collected from the Cenomanian-age Chaochuan Formation.[1]
Systematics[]
On the species description, Lü et al. (2007) found Zhejiangosaurus to belong to the ankylosaurian family Nodosauridae.[1] It is the only known nodosaurid from As
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Z is for Zuul – THE PAGE TURNS
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2024-05-02T16:29:47+00:00
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Yes, Zuul from Ghostbusters. Zuul who inhabits Sigourney Weaver’s body in order to search for the Keymaster, schlubby Rick Moranis, so that their coupling will release the demon Gozer into the world. A nerdy fantasy written by nerds for nerds. I was never a fan of the movie, but yesterday, when I was running down … Continue reading "Z is for Zuul"
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THE PAGE TURNS
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https://kajmeister.com/z-is-for-zuul
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Yes, Zuul from Ghostbusters. Zuul who inhabits Sigourney Weaver’s body in order to search for the Keymaster, schlubby Rick Moranis, so that their coupling will release the demon Gozer into the world. A nerdy fantasy written by nerds for nerds.
I was never a fan of the movie, but yesterday, when I was running down the list of which “Z” dinosaur would get the honor to front my very last post, and I said Zuul, my spouse immediately said Oh! The Gatekeeper of Gozer. Paleontologists, I suppose, are as nerdy as romance writers, medieval historians, and Hollywood directors, so, yes….
They did indeed name a dinosaur for Zuul.
Instead, it could have been one of the following… (list thoughtfully provided by thoughtco.com although it’s incomplete, since they forgot about Zuul).
Zalmoxes – A strange-looking ornithopod from Romania.
Zanabazar – Named after a Buddhist spiritual leader.
Zapalasaurus – This “diplodocoid” sauropod lived in early Cretaceous South America.
Zby – This dinosaur’s name was inversely proportional to its size.
Zephyrosaurus – Otherwise known as the Western Wind Lizard.
Zhanghenglong – A transitional hadrosaur of late Cretaceous Asia.
Zhejiangosaurus – The first identified nodosaur from Asia.
Zhenyuanlong – Also known as the “fluffy feathered poodle from hell.”
Zhongyuansaurus – The only known ankylosaur to lack a tail club.
Zhuchengceratops – It probably figured on the lunch menu of Zhuchengtyrannus.
Zhuchengosaurus – This hadrosaur was even bigger than Shantungosaurus.
Zhuchengtyrannus – This Asian tyrannosaur was the size of T. Rex.
Zuniceratops – This horned dinosaur was discovered by an eight-year-old boy.
Zuolong – It was named after General Tso, of Chinese restaurant fame.
Zupaysaurus – This “devil lizard” was one of the earliest theropods
But, nope, I picked Zuul.
For this last post on dinosaurs (*sniff* now I’m getting a bit verklempt), I will share some information on dinosaur naming conventions, as well as a few final thoughts on why we find dinosaurs so fascinating.
What’s in a Name?
Dinosaur names, like any other kind of creature extant or extinct, have two parts. There’s a genus part and a species part. Even though the classification system now has lots more tree branches than the old Kingdom-Phylum system, the branches always ends with genus and species. Remember those clades of letter “C,” which turn out to run everything? Good thing it was so high in my alphabet! Genus-species is at the tip of the clade.
The first part, the genus defines a group of animals that have similar characters. Homo is a genus because there were versions of humans before us sapiens—Homo neanderthalis, Homo erectus. The first part of the name, the genus, is always capitalized. The second part, the species, is never capitalized. Note: You are supposed to put the Latin names in italics, but I reserve the right for this post to be inconsistent. If I forget the italics, just ignore it.
Oops! I’ve been doing it wrong. It’s Tyrannosaurus rex, or T. rex. Never T. Rex… Now I have to back to previous posts and fix that!
By the way, I asked last week, why it wasn’t T. regina? Apparently, there is a species with that name! Just in 2022, paleontologists Person, Van Raalte, and Paul noted that everyone was constantly chucking anything that looked like a T. rex skull into the same pot, but they were actually different species. They argue for a T. imperator, too, a T. king, queen, and emperor. This analysis is causing quite a stir, as scientists disagree that the distinguishing features, such as number of teeth and size of the femurs, are not different enough. If the scientists have their way, it might reclassify famous statues, Sue in particular. The scientific debate will rage on!
Construction of Names and Clades
These two-pronged dinosaur names follow the general International Code for Zoological Nomenclature–back to the polysyllabic scientific world, whoopee! Also called the ICZN code, this shared agreement on naming conventions dates back to the late 19th century. Scientists were making up their own naming systems (Merton’s rules, Strickland’s codes) which got very confusing, so they got together and agreed they had to pick one. The ICZN is kept updated, but there’s some suggestion of tension in the wikipedia description of it that mentions…
Such new editions of the ICZN Code are not democratically approved by those taxonomists who are forced to follow the code’s provisions, neither do taxonomists have the right to vote for the members of the commission or the editorial committee.
Wikipedia explanation of ICZN codes
Sounds to me like some of those taxonomists weren’t too happy. But what’re they going to do, just make up a name? They have to get everyone to agree. Taxonomist, remember, is the person classifying the “thing.” If you want to give a dinosaur a name, you have to figure out what kind of dinosaur it is.
Most of the dinosaur naming relates to finding new dinosaurs. You can even find a list of all the scientific publications about dinosaurs, per year, which describes all the new and pending research arguments about new names. Just as of today, there are 18 pending new names for dinosaurs alone, including things like Titanomachya gimenezi, a small “big” titanosaur named after (1) the battle of Titans and (2) famous Argentine paleontologist Olga Gimenez. That’s a classic strategy, to use a combination of Greek and a person.
More often, the Greek genus name is descriptive of body parts. In 2015, a skeleton that was mis-classified was renamed Sefapanosaurus zastronensis. Sefapanosaurus referred to the cross-shaped ankle bone, while the species name mentions the South African town of Zastron, where the bones originated. These were actually found 80 years ago, and misclassified with Aardonyx. But scientists re-examined the structure and changed the name.
That’s the second part of the classification dance. While much of naming relates to new dinosaur parts found and constructed, there is a lot of re-classification occurring as well. What happens over time is that more bones are found, more evidence, which allows the tree to shift and split and reorganize. Start with what you have, but when you know more, then fix your mistakes.
When Zuul was first discovered, they thought it was another type of Euplocephalus (“well-armored head”). But they found a lot of pieces of Zuul, enough to be sure that it was on a distinct branch of the tree and that it should be put with the other ankylosaurs. They did call it Zuul, after the movie character who had that horned head. But then they added the species of crurivastator, which means destroyer of shins, because they thought the club tail would basically take out the bottom parts of attacking predators.
Life Imitates Art Imitates Life
Lots of dinosaurs have been named for fictional creatures–like Zuul– which is funny since the fictional creatures are often given names that sound like real creatures. There are new species discovered all the time. Actually, the majority of new names seem to be going to spiders, wasps, and beetles, so if you want to see all the creatures named after H.P. Lovecraft characters, look in the insect section of the ICZN.
Meanwhile every single species, dinosaur or otherwise, has a torturous path to its name. For example, the pterosaur (not a dinosaur!) named Targaryendraco wiedenrothi was originally named Ornithocheirus wiedenrothi, and grouped with other ornithocheirids. (Wiedenroth was an amateur fossil hunter who found it). After five more studies, though, it was put into its own tree branch and renamed, indeed for the House of Dragons from Game of Thrones.
More fun fictional-character-inspired dinosaurs (mostly):
Pantydraco caducus, after Pant-y-ffynnon in Wales, a thecodont
Hagryphus giganteus, Egyptian god Ra + griffin + big, an oviraptor
Gojirasaurus quayi, already noted after Godzilla, a theropod
Bambiraptor feinbergi, a theropod
Medusaceratops lokii, because of the Loki-like horns, a ceratopsian
Thanos simonattoi, a mean-looking theropod
Ozraptor subotaii, a thief, egg-stealer, from Subotai of the Conan series
Bradycneme draculae, found in Transylvania, another theropod
Irritator challengeri, from Conan Doyle’s Lost World, yet another theropod
Borogovia gracilicrus, like Carroll’s “borogoves,” still another theropod
Lohuecotitan pandafilandi, after Pandafilando, a giant in Don Quixote, a titanosaur
Dracorex hogwartsia
Now, the story about this last one is a bit sad. They found pieces of this dinosaur in South Dakota. After the skeleton was assembled and acquired by an Indiana children’s science museum, the museum had a naming contest. It resulted in the name related to Harry Potter, Dragon King of Hogwarts in Greek.
However, they apparently named it too quickly because it’s now thought to be a juvenile version of Pacycephalosaurus, a well-known and widespread group that includes the bony head vegeterians. It will probably get renamed and reclassified before ere long. Maybe Pacycephalosaurus hogwartsia???
Could Have Had a Spielberg Action Figure!
I talked about the Jurassic legacy back under letter “J,” that Spielberg had actually hired dinosaur experts for his 1993 film which made his dinosaurs more accurate than had ever been seen before and, in turn, influenced and inspired future dinosaur enthusiasts. The dinosaur enthusiasts wanted to return the favor.
In 1993, the group that had discovered the impressive Utahraptor wanted to give it the genus name spielbergi in anticipatory honor of the director. (Letter “U”–I didn’t even come across this story last week when I was writing about Utahraptors.) At the time, the group said they were looking for a funding source from either Spielberg or Universal Studios, and, when it wasn’t forthcoming, they changed the name.
But more recently, a different story has emerged that makes more sense because scientists usually aren’t blackmailers. According to a 2021 story in Inverse, Universal screwed it up for Steven, or themselves, depending on your point of view. There was a tiny Pennsylvania museum that planned to exhibit the Utahraptor, and they used the word Jurassic in their promotional materials. It’s kind of like using the word Gondwanaland or Oligocene–if you know what that is scientifically, you do, but other people don’t. At the time, in late 1992, it wasn’t a household word. But Universal was thinking about the trademark side of things, so they threatened this poor little Erie Zoological Society with legal action. The Putnam Museum had printed up T-shirts with Utahraptor spielbergi already on it, but they had to take the word “Jurassic” off all their signage, even though they had been using it for months before the movie was even advertised.
As a result, the Utahraptor scientist decided to choose the species name ostrommyi since the Spielberg Studios were making such a fuss. This was after none other than John Ostrom, the guy who made Deinonychus famous and helped push forward the dinosaur renaissance, which had influenced Spielberg’s choices.
Dreaming of Dinosaurs
It’s curious that humans have such a strong affinity for these creatures, which lived so long ago and have left mostly only bones. We don’t generally feel that way about trilobites, bacteria, or bark beetles even though they, in various ways, also ruled. We don’t even feel that way about our nearest cousins, the chimpanzees and gorillas. Instead, we write dystopian stories where we go to war with the apes on our clade.
But dinosaurs fill stories–I’ve even got one I’m playing around with that has to do with the stegosaurus shaped like a statistical chi-squared graph. It’s a children’s story, I’m working on it, still a little rough. KK reminded me the other day of the wonderful story by Guy Endore, “Day of the Dragon,” where a scientist fixes a flaw in a crocodile’s heart and it becomes a dragon. Are dragons dinosaurs? They seem like close cousins. Perhaps there was an undiscovered clade-line in remote parts of Siberia or China where pterosaur/dinosaurs really were dragons. Still, dragons are imaginary, but dinosaurs were real.
We’d like to think that we could tame them, or that they could teach us something, even though it’s hard to imagine actually doing that. And as the Jurassic movies showed, it’s dangerous to think about trying.
As others have pointed out, they feel half real and half imagined. Even though we can draw flesh on their bones, they seem like something we’ve only dreamed about. Because they ruled the earth and succumbed to climate change, we want to understand what happened to them, as hard as that may be (see “Y”). We want to know what these bones are trying to tell us, 200 million years later. Given that we’ve only been studying them in earnest ourselves for a couple of centuries, we’ve probably barely scratched the surface.
There’s a lot more, I’m sure, that they could tell us.
Thanks for reading!
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Alvarezsaurs in the Lori matrix
This time our topology is-
I provide a new definition for Alvarezsauroidea that adds Therizinosaurus as an external specifier since I find it most parsimonious for Therizinosauria to be its sister group, and uses Alvarezsaurus as the internal specifier unlike Sereno's that uses Shuvuuia- (Alvarezsaurus calvoi< - Ornithomimus velox, Therizinosaurus cheloniformis, Passer domesticus) . Alvarezsauroids have had a controversial phylogenetic placement, with the Lori matrix recovering them as basal maniraptorans sister to therizinosaurs. Yet they can be outside therizinosaurs plus pennaraptorans in 3 steps, become avemetatarsalians in 4 steps (can bring therizinosaurs or not), non-maniraptoriforms in 6 steps (they bring therizinosaurs), closer to pennaraptorans than therizinosaurs in 6 steps, paravians in 11 steps (therizinosaurs move with), closer to Compsognathus than to birds in 15 steps, closer to birds than deinonychosaurs in 27 steps, and closer to Archaeopteryx and other birds than to dromaeosaurids and troodontids in 30 steps.
Fukuivenator is an odd taxon, recovered here as the basalmost alvarezsauroid. But it can be a therizinosaurian in only two steps, and outside Maniraptoriformes in 4 steps (it emerges in Coeluridae). One thing I don't think it is is a dromaeosaurid, as that takes 27 more steps, and getting it into Paraves or Pennaraptora requires 11 and 7 steps respectively. Still, I wouldn't be surprised to see this taxon work its way around the base of Maniraptoriformes once an osteology comes out.
Shuvuuia deserti IGM 100/975 axial elements in ventral view and pelvis in dorsal view (courtesy AMNH).
Nqwebasaurus was recently redescribed by Sereno (2017), which I incorporated into its scorings. Choiniere et al. (2012) recovered it in Ornithomimosauria, but note most of the characters they list to support that are also said to be present in alvarezsauroids. Even they could place it in Alvarezsauroidea with only 4 steps. The Lori matrix needs 6 steps to place it in Ornithomimosauria, which I think is higher partially due to it finding Pelecanimimus to be an alvarezsauroid too. So similarities between the two like their teeth being in a common groove and maxillary teeth being confined to the anterior third of the bone are no longer ornithomimosaur-like. As recently noted by Cerroni et al. (2019), this makes more sense biogeographically as well. Oh, and note that the Lori matrix found Afromimus to be a ceratosaur as in that paper. In any case, Nqwebasaurus takes 10 steps to move to Compsognathidae, and 7 steps to move sister to Pennaraptora.
As for Pelecanimimus itself, it seems plausibly alvarezsauroid if you think about it. The skull is famously similar to Shuvuuia, the posterior tympanic recess is in the otic recess, ossified sterna are otherwise unknown for ornithomimosaurs, the long manual digit I was always out of place compared to Harpymimus, and Europe makes more sense for otherwise Gondwanan clades in the Cretaceous. Now if only someone would release Perez-Moreno's thesis describing it in detail...
Shuvuuia deserti IGM 100/975 pectoral and forelimb elements. Note the tiny phalanx from digit II or III at the bottom (courtesy AMNH).
Patagonykus and Bonapartenykus are usually closer to parvicursorines than Alvarezsaurus and Achillesaurus, but the Lori matrix found them just outside Alvarezsauridae instead. Interestingly, Xu et al. (2018) recovered the same results. It takes 3 steps to move Patagonykus closer to parvicursorines, and 4 steps to join Alvarezsaurus and Patagonykus to the exclusion of parvicursorines as in Alifanov and Barsbold (2009). Xu et al. recover these in 5 and 7 steps respectively, and the most recent version of Longrich and Currie's alvarezsaurid matrix (Lu et al., 2018) recovers a basal Patagonykus and a basal Parvicursorinae in 3 steps each.
One odd result is that the newly described Xiyunykus and Bannykus fall in Patagonykinae too. Yet only 2 steps move them outside the Patagonykus plus Parvicursorinae clade, where they form a clade. Another step breaks that up to place Xiyunykus more basal as in Xu et al.. Them being basal certainly fits better stratigraphically, and Xu et al. use several characters designed for alvarezsauroids that the Lori matrix didn't include yet. Hopefully full osteologies will be published as well.
Mononykus olecranus cast YPM 56693 (of holotype) pes in plantar view (courtesy of Senter).
A patagonykine Achillesaurus as suggested by Agnolin et al. (2012) takes 7 additional steps in the Lori matrix where it instead emerges just closer to parvicursorines than Alvarezsaurus. On the other hand, only a single step joins it with Alvarezsaurus as in Longrich and Currie (2009) and only 2 steps makes it just further from parvicursorines than Alvarezsaurus as in Xu et al. (2018).
Alnashetri is known from type hindlimb material, but now also from MPCA 377, a nearly complete specimen with interesting characters like flat and unfused sternal plates. Makovicky et al. (2016) used this data to recover it as the sister group to Alvarezsauridae, and while the few published details left it more derived in the Lori tree, it can go to a more basal position with only two steps. It should be interesting to compare to e.g. Bannykus once it is published.
Mononykus olecranus cast YPM 56693 (of holotype) (courtesy of Senter).
The arctometatarsal clade has a unique topology, but no other analysis has included nearly as many characters or all of these taxa, with Lu et al. omitting Albinykus and Ceratonykus among non-fragmentary specimens, and Xu et al. omitting the more recently described Qiupanykus. Enforcing the Lori topology in Lu et al.'s matrix is only 5 steps longer, and doing so in Xu et al.'s matrix is only 6 steps longer. On the other hand, Xu et al.'s topology is so unresolved at this level, the only difference in mine is placing the Albinykus plus Xixianykus clade basally near Albertonykus, which takes 5 steps to do in the Lori matrix.
It should be noted that Lu et al.'s illustrated topology (their Figure 3) is not their matrix's real result, as they did not fully analyze tree space. Instead of 20 trees, there are 214 trees. These differ in that Albertonykus, YPM 1049 and undescribed 41HIII-0104 can fall out anywhere more derived than Patagonykus, and that Parvicursor, the Tugriken Shireh taxon, Shuvuuia and Mononykus form an unresolved polytomy. This leaves Linhenykus, Qiupanykus and Xixianykus unresolved between that polytomy and Patagonykus, which is perfectly compatible with the Lori topology. This may also show that the small alvarezsauroid-specific matrix of Longrich and Currie is insufficient given all the new taxa described since 2009. YPM 1049 was far too fragmentary to include (distal metatarsal III) but I tried testing undescribed Quipa specimen 41HIII-0104. Didn't make it into the publication, but here's its scorings-
'41HIII0104' ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ???????-?? ?????????1 ?10??????? ?????????? ?????????? ???0?????? ????1????? ???1?????{01} ?????????? ?????????? ?????????? ????????0? ????????3? ?????????? ?????????? ?????????? ?????????? ?????????1 ?????????1 1????????? 1????????? ?????????? ?????????? ?????????? ???1?????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ????????0? ?????????? ?????????? ?????????? ?????????? ?????????? ???1?????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????{123}???? ?????????? ?????????? ?????????? ?????????? ?????????? ??1???0???
Tugriken Shireh parvicursorine (IGM 100/99) vertebrae and ilia in ventral view, forelimb and fibula in lower right (courtesy AMNH).
Interestingly, Agnolin et al., Xu et al. and the Lori analysis all recovered Albinykus sister to Xixianykus outside Parvicursorinae. Wonder if that's a real signal? Unfortunately, the only attempt to name this clade was Agnolin et al. who also recovered Ceratonykus in there and called it Ceratonykini. Xu et al. place Ceratonykus closer to parvicursorines, while I found it more basal than either, sister to Qiupanykus which neither of the other studies used. Forcing Ceratonykus sister to Albonykus plus Xixianykus takes 3 more steps in the Lori matrix. Forcing Ceratonykus sister to Mononykus as in its original description (with or without Qiupanykus) takes 5 more steps. As stated in the paper, we were the first analysis to include Hateg tibiotarsi Bradycneme and Heptasteornis. While the former can fall into many positions in Maniraptora, the latter was resolved as an alvarezsaurid as proposed by Naish and Dyke (2004). Note this used only the tibiotarsus and not alvarezsaurid-like distal femur FGGUB R.1957. A single step moves Heptasteornis to Troodontidae.
We also provide an updated definition for Parvicursorinae (Mononykus olecranus + Parvicursor remotus), like Choiniere et al.'s (2010) but using species. One accident of our definitional and discovery history is that all these newer arctometatarsal alvarezsaurids (Xixianykus, Albertonykus, Albinykus, Linhenykus, Qiupanykus, Ceratonykus, etc.) emerge outside the originally discovered and defined Parvicursorinae. We could really use some clade defining taxa closer to Mononykus than Patagonykus, Alvarezsaurus or Achillesaurus. In any case, I got a lot of experience with parvicursorine specimens, examining Shuvuuia and the Tugriken Shireh specimen IGM 100/99 in person, and having photos of high quality casts of Mononykus thanks to Senter. I found the Tugriken Shireh taxon closer to Shuvuuia, but moving it closer to Parvicursor as in Longrich and Currie is just 1 step longer.
Tugriken Shireh parvicursorine (IGM 100/99) vertebrae and ilia in dorsal view, forelimb and fibula in lower right (courtesy AMNH).
Next time, therizinosaurs...
References- Naish and Dyke, 2004. Heptasteornis was no ornithomimid, troodontid, dromaeosaurid or owl: The first alvarezsaurid (Dinosauria: Theropoda) from Europe. Neus Jahrbuch für Geologie und Paläontologie. 7, 385-401.
Alifanov and Barsbold, 2009. Ceratonykus oculatus gen. et sp. nov., a new dinosaur (?Theropoda, Alvarezsauria) from the Late Cretaceous of Mongolia. Paleontological Journal. 43(1), 94-106.
Longrich and Currie, 2009. Albertonykus borealis, a new alvarezsaur (Dinosauria: Theropoda) from the Early Maastrichtian of Alberta, Canada: Implications for the systematics and ecology of the Alvarezsauridae. Cretaceous Research. 30(1), 239-252.
Choiniere, Xu, Clark, Forster, Guo and Han, 2010. A basal alvarezsauroid theropod from the early Late Jurassic of Xinjiang, China. Science. 327, 571-574.
Agnolin, Powell, Novas and Kundrat, 2012. New alvarezsaurid (Dinosauria, Theropoda) from uppermost Cretaceous of north-western Patagonia with associated eggs. Cretaceous Research. 35, 33-56.
Makovicky, Apesteguia and Gianechini, 2016. A new, almost complete specimen of Alnashetri cerropoliciensis(Dinosauria: Theropoda) impacts our understanding of alvarezsauroid evolution. XXX Jornadas Argentinas de Paleontologia de Vertebrados. Libro de resumenes, 74.
Sereno, 2017. Early Cretaceous ornithomimosaurs (Dinosauria: Coelurosauria) from Africa. Ameghiniana. 54, 576-616.
Lu, Xu, Chang, Jia, Zhang, Gao, Zhang, Zhang and Ding, 2018. A new alvarezsaurid dinosaur from the Late Cretaceous Qiupa Formation of Luanchuan, Henan Province, central China. China Geology. 1, 28-35.
Xu, Choiniere, Tan, Benson, Clark, Sullivan, Zhao, Han, Ma, He, Wang, Xing and Tan, 2018. Two Early Cretaceous fossils document transitional stages in alvarezsaurian dinosaur evolution. Current Biology. 28, 1-8. DOI: 10.1016/j.cub.2018.07.057
Cerroni, Agnolin, Egli and Novas, 2019. The phylogenetic position of Afromimus tenerensis Sereno, 2017 and its paleobiogeographical implications. Journal of African Earth Sciences. DOI: 10.1016/j.jafrearsci.2019.103572
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
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Therizinosaurs in the Lori matrix
Next up are therizinosaurs. These are one of the best analyzed clades because I incorporated all of Zanno's (2010) characters, which is by far the largest and most recent analysis of the group until the Lori paper was published. The topology is-
Falcarius is the most basal taxon shown of course, but Martharaptor was pruned a posteriori and can fall out anywhere in Therizinosauria outside the Alxasaurus plus Segnosaurus clade. I tried including Thecocoelurus, but the Lori matrix is pretty terrible when it comes to scoring single vertebrae-
Thecocoelurus ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ???????(01)0? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ???1?????? ?????????? ?????2???? ?????????? ?????????? ?????????? ?????????? ???????0?? ?????????? ?????????? ?????????? ?????1???? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ??0??????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ??????????
Jianchangosaurus fell out in the same place as its original description, with Cau's (2018) placement in Alvarezsauroidea taking 18 more steps so is very unlikely. Mine is the only published matrix besides Senter (2011) and its derivatives to use information from the second Beipaiosaurus specimen, and incorporated photos from Zanno and the new paper on the holotype skull elements too.
Beipiaosaurus inexpectus holotype (IVPP V11559) cervical vertebra in dorsal view (courtesy of Zanno).
Zanno also provided photos of Alxasaurus and Enigmosaurus, and its depressing how much of the former is lost. Enigmosaurus rather famously was shown by Zanno to not resemble Barsbold's original illustration that was the only reference picture known for over two decades. Placing Enigmosaurus closer to Segnosaurus than Neimongosaurus or Erliansaurus as in Zanno's tree takes 4 more steps. Forcing Enigmosaurus and Erlikosaurus to be sister taxa to simulate the synonymy mentioned by Barsbold (1983) takes 4 steps, so seems unlikely. The duo moves between Nanshiungosaurus and the Segnosaurus plus Nothronychus clade. We were the first analysis to include "Chilantaisaurus" zheziangensis, which emerged in a polytomy with Alxasaurus, Enigmosaurus and therizinosaurids.
Alxasaurus elesitaiensis holotype (IVPP V88402a) chevrons in right lateral view (natural order reversed) (courtesy of Zanno).
As was the case with Archaeornithomimus? bissektensis, we didn't include the possible chimaera of Bissekty Therizinosauria as an OTU, unlike Sues and Averianov (2015). But if you do want to experiment with it, here's the scorings. It emerges in a polytomy in the Suzhousaurus plus Therizinosaurus clade of therizinosaurids. Btw, Archaeornithomimus? bissektensis does fall out most parsimoniously sister to A. asiaticus when all Bissekty material is used.
'Bissekty-Therizinosauroidea' ????1??0?? ????1????? ?????1???? ?1???????? ?????{01}1000 ?01??????? 00???????? 2??21?0??? ???????00{123} {12}00(01)2011?? ??0???0(01)00 ??11????{12}? ?{01}0?100??? ?????????? ?1{01}?0????? ??0?0???{012}0 000(01)?010?? ?????????? ?????????? ?????0{12}?00 1?0???0??0 ??0??{01}0??? 0?1??????? ????0?1?0? {01}0?1???{01}?? ??0??????? ?????????? ?????????0 ???001??00 ??00?01?1? 10???????? ?????????1 ???0?????? ???1?????? ??????0??1 0???1(12)???? ????0???1? ????0?011? ???0????0? ???0{12}0???? ????????0? 0?0????1?? ?????????? ??010101?1 ??0?0(01)???? ?????{01}010? ???100???? ????????11 1010?????? ??0??0???? ?????????? ?????????? ??????0??? ??00?????? ?00??????? ?????????? ???????0-- -??00??010 ?????????? ??????-??? ???-010??1 0100????00 10?1?00??? ???0?00??? ?????????? ????0????? ???000???? ?0???-???? ?????????? ?????001??
Enigmosaurus mongoliensis holotype (IGM 100/84) synsacrum and ilium in ventral view (courtesy of Zanno).
Next is Therizinosauridae itself, which we refined Zhang et al.'s (2001) definition of to include type species. Therizinosaurids first split into a clade of Erliansaurus, Neimongosaurus, Suzhousaurus and Therizinosaurus. Forcing the former two to be outside a clade of Suzhousaurus, Therizinosaurus and the taxa below, as in Zanno's tree, takes 5 more steps. Notably, we did not include the hindlimb IGM 100/45 in the Therizinosaurus OTU since there's no overlap and its not even particularly large. But here's the Therizinosaurus OTU including the hindlimb. Using this version of Therizinosaurus leaves the tree basically the same but destabilizes it somewhat in that Therizinosaurus and Erlikosaurus can now go in multiple positions within Therizinosauridae, and the Nanchao embryos are in a trichotomy with the Suzhousaurus clade and the Nothronychus clade. Is this an indication the hindlimb produces homoplasy and so might not belong to Therizinosaurus?
Therizinosaurus ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ????????01 0110021000 01?0001100 00001110?? ?????????? ?????????? ?????????? ???0???010 1?0000000? 0?1??1???? ?????????? ?????????? 000??0???? ??????3??1 ?????????0 ???00????? 0?1?0?1??? ?????0???? ?????????1 ?????????? ?????0???? ?????????? ?????????? 1011010101 100100??1? ????{12}?00?? 00?12110?? 101??????? 0????0?111 00?010???? ????????1? ?????????? ????100111 1101110??? ?????????? ????????1? ?1???????? ?????????1 ???0?????? ?000?????? ?????0???1 ?????????? ?????????? ?????????? ?????????? ?????0???? ??????-00? ?00-000001 0?0100??0? 10000???0{01} ?{01}?0000??0 {01}0???????? ????????00 ?00?????0- -??11-???? ?????????? ???1?0???0
Segnosaurus galbinensis paratype (IGM 100/83) cervical neural arch in right lateral view (courtesy of Zanno).
Now comes the Nanchao therizinosaur embryos, those described by Kundrat et al. inside dendroolithid eggs. While including such young specimens might be seen as risky, my ontogenetically conservative scoring method with state N seems to have worked fine here. They fall out where you'd expect a Santonian-Campanian therizinosaur to do so. Following that is Nanshiungosaurus brevispinus, which Senter et al. (2012) recovered as the next most derived therizinosaur after Alxasaurus. Forcing it into this basal position takes 4 steps. Nanshiungosaurus? bohlini was included but pruned a posteriori since it can go anywhere in the Segnosaurus plus Nothronychus clade. Forcing Nanshiungosaurus monophyly is just a single step longer though, while forcing bohlini to be sister to the contemporaneous Suzhousaurus takes 2 steps.
Segnosaurus itself (which Zanno also provided photos of) pairs with ex-Alectrosaurus forelimb AMNH 6368, which has only previously been analyzed by Zanno (2006) where it pairs with Erliansaurus. Forcing that here compared to other taxa she included results in trees 3 steps longer. Erlikosaurus groups with the Nothronychus species in a trichotomy where it can be sister to either species. Forcing Nothronychus monophyly takes only a single step, but note that no proposed Nothronychus characters involve elements that can be compared to Erlikosaurus (humerus and pes). Forcing Erlikosaurus to group with Therizinosaurus as in Senter et al. requires only a single step, with Erlikosaurus moving to the Therizinosaurus clade.
Next time, oviraptorosaurs...
References- Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Transactions of the Joint Soviet-Mongolian Palaeontological Expedition. 19, 117 pp.
Zhang, Xu, Sereno, Kwang and Tan, 2001. A long-necked therizinosauroid dinosaur from the Upper Cretaceous Iren Dabasu Formation of Nei Mongol, People’s Republic of China. Vertebrata PalAsiatica. 39(4), 282-290.
Zanno, 2006. The pectoral girle and forelimb of the primitive therizinosauroid Falcarius utahensis (Theropoda, Maniraptora): Analyzing evolutionary trends within Therizinosauroidea. Journal of Vertebrate Paleontology. 26(3), 636-650.
Zanno, 2010. A taxonomic and phylogenetic re-evaluation of Therizinosauria (Dinosauria: Maniraptora). Journal of Systematic Palaeontology. 8(4), 503-543.
Senter, 2011. Using creation science to demonstrate evolution 2: Morphological continuity within Dinosauria. Journal of Evolutionary Biology. 24(10), 2197-2216.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid tail. PLoS ONE. 7(5), e36790.
Sues and Averianov, 2015. Therizinosauroidea (Dinosauria: Theropoda) from the Upper Cretaceous of Uzbekistan. Cretaceous Research. 59, 155-178.
Cau, 2018. The assembly of the avian body plan: A 160-million-year long process. Bollettino della Società Paleontologica Italiana. 57(1), 1-25.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
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Happy New Year 2020
Hi all. A Theropod Database update is online, with the main additions being troodontid information and info from the Hayashibara Museum of Natural Sciences Research Bulletins 1-3. I love these publications and wish more like them existed for other collections. They detail the expeditions into Mongolia with exact discovery dates and field numbers for taxa like Nomingia, Elsornis and Aepyornithomimus, and tons of still undescribed specimens. It's amazing just how many ornithomimosaurs are known from the Bayanshiree Formation for instance, when only the Garudimimus holotype has been described. There are over twenty more including the sort-of-described "Gallimimus""mongoliensis" specimen IGM 100/14. So often for new taxa, especially those from the Jehol biota, no information is provided in the description as to when the specimen was discovered. I get that many are found by non-professionals and given to museums, but at least say "the specimen was given to the museum on x-x-xx by someone who said it was excavated around year y." Next up, halszkaraptorine and dromaeosaurid updates...
undescribed ?Gobivenator skull (HMNS coll.; field number 940801 TS-I WTB) (after Tsogtbaatar and Chinzorig, 2010).
Reference- Tsogtbaatar and Chinzorig, 2010. Fossil specimens prepared in Mongolian Paleontological Center: 2002–2008. Hayashibara Museum of Natural Sciences Research Bulletin. 3, 155-166.
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Details on Teinurosaurus and random musings
Hi all. When updating The Theropod Database I noticed my entry for Teinurosaurus is pathetically bad- wrong authors, wrong age, wrong size, and generally missing the complicated history of this innocuous vertebra. How embarrassing! So here's the revised version that will be uploaded-
Teinurosaurus Nopcsa, 1928
= Saurornithoides Nopsca, 1928 (preoccupied Osborn, 1924)
= Caudocoelus Huene, 1932
T. sauvagei (Huene, 1932) Olshevsky, 1978
= Caudocoelus sauvagei Huene, 1932
Tithonian, Late Jurassic
Mont-Lambert Formation, Hauts-de-France, France
Holotype- (BHN2R 240; = Boulogne Museum 500) incomplete distal caudal vertebra (75 mm)
Diagnosis- Provisionally indeterminate relative to Kaijiangosaurus, Tanycolagreus and Ornitholestes.
Other diagnoses- (after Huene, 1932; compared to Elaphrosaurus) centrum wider; narrower ventral surface; ventral median groove wider; transversely narrower prezygapophyses.
While Huene attmpted to distinguish Teinurosaurus from Elaphrosaurus, only the wider median ventral groove is apparent in existing photos of the former. This is compared to the one distal caudal of the latter figured in ventral view, but as Kobayashi reports grooves become distally narrower in Harpymimus while Ostrom reports they become distally wider in Deinonychus, groove width is not considered taxonomically distinctive at our current level of understanding. Indeed, this lack of data is most relevent to both diagnosing and identifying Teinurosaurus. Very few taxa have detailed descriptions of distal caudal vertebrae or more than lateral views figured, let alone indications of variation within the distal caudal series. So the facts that Fukuiraptor and Deinonychus share ventrally concave central articulations with Teinurosaurus in their single anteriorly/posteriorly figured distal caudal vertebra, or that Afromimus, "Grusimimus" and Falcariusalso have have wide ventral grooves in their few ventrally figured distal caudals, are not considered taxonomically important.
Comments- Sauvage (1897-1898; in a section written in January 1898) first mentioned a distal caudal vertebra he referred to the ornithischian Iguanodon prestwichii (now recognized as the basal styracosternan Cumnoria prestwichii) - "We are disposed to regard as belonging to the same species the caudal vertebra of a remote region, the part which we figure under n ° 7, 8" [translated]. Note Galton (1982) was incorrect in claiming Sauvage reported on this specimen in his 1897 paper (written December 6), which includes a section on prestwichiinearly identical to the 1897-1898 one but which lacks the paragraph describing this vertebra. This could provide a specific date of December 1897 to January 1898 for the discovery and/or recognition of the specimen. Huene (1932) correctly noted Sauvage mislabeled plate VII figure 8 as dorsal view, when it is in ventral view as understood by the text. Compared to Cumnoria, the caudal is more elongate (length 3.93 times posterior height compared to 2.54 times at most), has a ventral median groove instead of a keel, and the prezygapophyseal base in 71% of the anterior central height compared to ~30-40%, all typical of avepods. Nopcsa (1928) recognized its theropod nature and in his list of reptile genera meant to use a footnote to propose Teinurosaurusas a "new name for the piece described and figured by Sauvage (Direct. Traveaux Geol. Portugal Lisbonne 1897-1898, plate VII, Fig. 7-10) as late caudal of Iguanodon Prestwichi." Teinurosaurusis listed as an aublysodontine megalosaurid (not as an ornithomimine, contra Galton), roughly equivalent to modern Eutyrannosauria. However due to a typographical error, the footnote's superscript 1 was placed after Saurornithoides instead of Teinurosaurus. Sauvage (1929) corrected this in an addendum- "footnote 1 does not refer to Saurornithoides (line 19 from below) but to Teinurosaurus(last line of text)." Unfortunately, Huene missed the addendum, and thus wrote "Nopcsa recognized in 1927 (43, p. 183) that this was a coelurosaur and intended to give it a name, but used one already used by Osborn, namely "Saurornithoides" (91, 1924, p. 3- 7). For this reason, a new name had to be given here" [translated]. Huene's proposed new name was Caudocoelus sauvagei, placed in Coeluridae and "somewhat reminiscent of Elaphrosaurus." Huene is also perhaps the first of several authors to place the specimen in the Kimmeridgian, when it is actually from the Tithonian (Buffetaut and Martin, 1993; as Portlandian). Galton wrote "Lapparent and Lavocat (1955: 801) gave a line drawing of the vertebra after Sauavage (1898) and included it in the section on Elaphrosaurus" and that the specimen "was referred to Elaphrosaurusby Lapparent and Lavocat (1955)." This was perhaps done because Huene explicitly compared the two, ironically making it the only taxon distinguished from Teinurosaurus at the time. Most of Huene's characters cannot be checked in the few published photos of Teinurosaurus, but the ventral median sulcus is indeed much wider than Elaphrosaurus. Ostrom (1969) was the first author to detail Nopcsa's (1929) addendum, stating "Nopcsa's name Teinurosaurus has clear piority over Huene's Caudocoelus, but since Nopcsa failed to provbide a specific name, Teinurosaurus is not valid." Olshevsky (1978) solved this by writing "Teinurosaurus has clear priority over Caudocoelus, as noted in Ostrom 1969, and it is certainly a valid generic name. The species Caudocoelus sauvagei is proposed here as the type species of the genus Teinurosaurus, resulting in the new combination Teinurosaurus sauvagei(von Huene 1932) as the proper name of the type specimen." He also claimed "the specimen itself, unfortunately, was destroyed during World War II and thus must remain a nomen dubium." This was repeated by Galton, but as Buffetaut et al. (1991) wrote- "Contrary to a widespread opinion (expressed, for instance, by Lapparent, 1967), the vertebra in question has survived two world wars and years of neglect, like a large part of the other fossil reptile remains in the collections of the Boulogne Natural History Museum (see Vadet and Rose, 1986)." Olshevsky noted Steel misunderstood Nopsca in a different way, believing Teinurosaurus instead of Aublysodon was a "name, proposed by Cope in 1869 ... used instead of Deinodon", as stated under superscript 2. Galton did have the first modern opinion on Teinurosaurus' affinities, stating "In addition to Elaphrosaurus, elongate prezygapophyses occur in the allosaurid Allosaurus and the dromaeosaurid Deinonychus, so this caudal vertebra can only be identified as theropod, family incertae sedis." Buffetaut and Martin (1993) agreed, saying "no really distinctive characters that would allow a familial assignment can be observed." Ford (2005 online) gave the type repository as "Dortigen Museum", but this is a misunderstanding based on Huene's "Boulogne-sur-mer (Nr. 500 im dortigen Museum)", which roughly translated is "Boulogne-sur-mer region (No. 500 in the museum there)", referring to the Boulogne Museum where it has always been held. It was originally number 500, but was recatalogued at some point.
Sauvage lists the vertebra's length as 75 mm and his plate at natural size would have it be 79 mm, Huene lists it as 11 cm (110 mm) and his figure at 1:2 size would have it be 152 mm. Galton's drawing with supposed 5 cm scale would have it be 235 mm, while Buffetaut and Martin's plate with scale would leave it at 74 mm. As Huene's and Galton's figures are taken from Sauvage's original plate and the newest and unique photo matches Sauvage's reported length almost exactly, 75 mm is taken as the correct length.
Relationships- While prior authors haven't specified Teinurosaurus' relationships past Theropoda (besides Lapparent and Lavocat's apparent synonymy with Elaphrosaurus), there are several ways to narrow down its identity. Only neotheropods are known from the Late Jurassic onward, so coelophysoid-grade taxa are excluded. Some theropod clades were too small to have a 75 mm caudal, including most non-tyrannosauroid coelurosaurs besides ornithomimosaurs, therizinosaurs and eudromaeosaurs. The former two are unknown from the Jurassic, and additionally paravians like eudromaeosaurs lack any neural spine by the time the centrum gets as elongate as Teinurosaurus (e.g. by caudal 12 in Deinonychus at elongation index of 2.4). Teinurosaurus has an elongation index (centrum length/height) of 3.9, which also excludes Ceratosauridae, Beipiaosaurus+ therizinosauroids and oviraptorosaurs. Prezygapophyses basal depth is significantly less in ceratosaurids, megalosaurids, carnosaurs except Neovenator, compsognathids, Fukuivenator and Falcarius. Remaining taxa are elaphrosaur-grade ceratosaurs, piatnitzkysaurids, Neovenator and basal tyrannosauroids.
References- Sauvage, 1897. Notes sur les Reptiles Fossiles (1). Bulletin de la Société géologique de France. 3(25), 864-875.
Sauvage, 1897-1898. Vertebres Fossiles du Portugual, Contributions a l'etude des poissions et des reptiles du Jurassique et du Cretaceous. Direction des Travaux Geologiques Portugal. 1-46.
Osborn, 1924. Three new Theropoda, Protoceratops zone, central Mongolia. American Museum Novitates. 144, 1-12.
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Nopcsa, 1929. Addendum "The genera of reptiles". Palaeobiologica. 2, 201.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1), 361 pp.
Lapparent and Lavocat, 1955. Dinosauriens. In Piveteau (ed.). Traite de Paleontologie. Masson et Cie. 5, 785-962.
Lapparent, 1967. Les dinosaures de France. Sciences. 51, 4-19.
Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Peabody Museum of Natural History Bulletin. 30, 1-165.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia of Paleoherpetology. Gustav Fischer Verlag. 87 pp.
Olshevsky, 1978. The archosaurian taxa (excluding the Crocodylia). Mesozoic Meanderings. 1, 50 pp.
Galton, 1982. Elaphrosaurus, an ornithomimid dinosaur from the Upper Jurassic of North America and Africa. Paläontologische Zeitschrift. 56, 265-275.
Vadet and Rose, 1986. Catalogue commente des types et figures de dinosauriens, ichthyosauriens, sauropterygiens, pterosauriens et cheloninens du Musée d'Histoire Naturelle de Boulogne-sur-Mer. In E. Buffetaut, Rose and Vadet (eds.). Vértébrés Fossiles du Boulonnais. Mémoires de la Société Académique du Boulonnais. 1(2), 85-97.
Rose, 1987. Redecouverte d'une vertebre caudale reptilienne (Archosauriens) de status controverse et provenant des terrains jurassiques superieurs du Boulonnais. Bulletin de la Société académique du Boulonnais. 1(5), 150-153.
Buffetaut, Cuny and le Loeuff, 1991. French Dinosaurs: The best record in Europe? Modern Geology. 16(1-2), 17-42.
Buffetaut and Martin, 1993. Late Jurassic dinosaurs from the Boulonnais (northern France): A review. Revue de Paléobiologie. 7(vol. spéc.), 17-28.
Ford, 2005 online. http://www.paleofile.com/Dinosaurs/Theropods/Teinurosaurus.asp
And before we go, here are a couple more tidbits I've noticed in the upcoming update...
- That theropod tail preserved in Burmese amber (DIP-V-15103) described by Xing et al. (2016) was only placed as specifically as a non-pygostylian maniraptoriform. But as the deposits are Gondwanan (e.g. Poinar, 2018), the range of potential Cenomanian theropods is better understood. And only one group has caudal centra over three times longer than tall- unenlagiines. I bet DIP-V-15103 is our first sample of preserved plumage in an unenlagiine, which makes you wonder if the weird alternating barb placement was a feature that evolved on Gondwana, and if so did Rahonavis' remiges exhibit it too?
- Does anyone realize both "Tralkasaurus" (Cerroni et al., 2019) and "Thanos" (Delcourt and Iori, 2018) are nomina nuda? Neither are in an official volume yet, though "Tralkasaurus" is scheduled for March and "Thanos" will probably make it this year if the average papers per volume of Historical Biology holds up. "Tralkasaurus" has an empty space in its "Zoobank registration:" section, while the "Thanos" paper doesn't mention ZooBank at all, and neither show up in ZooBank searches. Also, one of "Thanos"' supposed autapomorphies is a deep prezygapophyseal spinodiapophyseal fossa, which does not exist in abelisaurs as it would require a spinodiapophyseal lamina. The labeled structure seems internal, probably the centroprezygapophyseal fossa or prezygapophyseal centrodiapophyseal fossa based on CT-scanned noasaurid cervical DGM929-R. That leaves axial pleurocoel size and distance from each other, and ventral keel strength as suggested characters. Which can only be compared to Carnotaurus among brachyrostrans. Hmmm...
References- Xing, McKellar, Xu, Li, Bai, Persons IV, Miyashita, Benton, Zhang, Wolfe, Yi, Tseng, Ran and Currie, 2016. A feathered dinosaur tail with primitive plumage trapped in Mid-Cretaceous amber. Current Biology. 26(24), 3352-3360.
Delcourt and Iori, 2018. A new Abelisauridae (Dinosauria: Theropoda) from São José do Rio Preto Formation, Upper Cretaceous of Brazil and comments on the Bauru Group fauna. Historical Biology. DOI: 10.1080/08912963.2018.1546700
Poinar, 2018. Burmese amber: Evidence of Gondwanan origin and Cretaceous dispersion. Historical Biology. DOI: 10.1080/08912963.2018.1446531
Cerroni, Motta, Agnolín, Aranciaga Rolando, Brissón Egli and Novas, 2019. A new abelisaurid from the Huincul Formation (Cenomanian-Turonian; Upper Cretaceous) of Río Negro province, Argentina. Journal of South American Earth Sciences. 98, 102445.
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Oculudentavis is not a theropod
Hi all. This week we got the announcement of a tiny theropod skull in Myanmar amber, which was bound to happen eventually as amazing finds from that deposit keep being published. Alas, whatever Oculudentavis is, it's not a theropod.
Oculudentavis skull (after Xing et al., 2020).
Just look at it. No antorbital fenestra, incomplete ventral bar to the laterotemporal fenestra, huge posttemporal fenestrae, teeth that extend posteriorly far under the orbit...
All of which might be coincidental, but then look at the mandible.
Oculudentavis mandible (after Xing et al., 2020).
That spike-like coronoid process is classic lepidosaur, plus the dentary is way too long compared to the post-dentary elements, then the description says "The tooth geometry appears to be acrodont to pleurodont; no grooves or sockets are discernable." And of course "the scleral ring is very large and is formed by elongated spoon-shaped ossicles; a morphology similar to this is otherwise known only in lizards (for example, Lacerta viridis)."
Add to this the size of this partially fused specimen being smaller than any extant bird (14 mm), and no feather remains, and why is this a theropod again? The endocast is big, but why not a clade of brainier lizards or late surviving megalancosaurs by the Cenomanian?
The authors add it to Jingmai's bird analysis where it ends in a huge polytomy closer to Aves than Archaeopteryx, but outside fake Ornithuromorpha. That's often what happens when a taxon is wrongly placed in a clade. Note the figured placement between Archaeopteryx and Jeholornis is only found using implied weights. At least add it to e.g. Nesbitt's or Ezcurra's archosauromorph analyses, or Cau's theropod analyses before assuming it's a bird.
Thanks to Ruben Molina Perez for suggesting this issue in the first place.
Reference- Xing, O'Connor, Schmitz, Chiappe, McKellar, Yi and Li, 2020. Hummingbird-sized dinosaur from the Cretaceous period of Myanmar. Nature. 579, 245-249.
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What is Oculudentavis if it's not a theropod?
In my last post, I argued the recently described Oculudentavis (Xing et al., 2020) is not a theropod. So what is it? To answer that question, I entered it into Simoes et al.'s (2018) sauropsid analysis which emphasizes basal lepidosauromorphs and comes out with basal gekkos and nested iguanians even using just morphological characters. To test Jingmai's avialan hypothesis, I also added Archaeopteryx to the matrix. The result is 384 MPTs of 2337 steps each.
Strict consensus of 384 MPTs of Simoes et al.'s (2018) analysis after adding Oculudentavis and Archaeopteryx. Compare to Extended Data Figure 3 of Simoes et al..
As you can see, Oculudentavis resolves as a stem-squamate in a trichotomy with Huehuecuetzpalli and squamates, while Archaeopteryx is an archosauromorph sister to Erythrosuchus. And this matrix didn't score for scleral ossicle shape, posttemporal fenestra size or maxillary tooth row length. After scoring Oculudentavis, its teeth are clearly not acrodont, it seems to have a ventral parietal fossa and lacks an ossified laterosphenoid. The authors could have made it easier to evaluate by separating the cranial elements in the 3D pdf file. As it is, a lot of palatal and braincase info is uncertain. But Huehuecuetzpalli is Albian compared to Oculudentavis' Cenomanian, and has a skull length of 32 mm (19 mm in the juvenile) versus 14 mm in Oculudentavis.
Huehuecuetzpalli skull (top; after Reynoso, 1998), Oculudentavis skull and separate mandible (middle; after Xing et al., 2020), and Archaeopteryx skull (after Rauhut, 2014).
References- Reynoso, 1998. Huehuecuetzpalli mixtecus gen. et sp. nov: A basal squamate (Reptilia) from the Early Cretaceous of Tepexi de Rodriguez, central Mexico. Philosophical Transactions of the Royal Society B: Biological Sciences. 353, 477-500.
Rauhut, 2014. New observations on the skull of Archaeopteryx. Paläontologische Zeitschrift. 88(2), 211-221.
Simōes, Caldwell, Talanda, Bernardi, Palci, Vernygora, Bernardini, Mancini and Nydam, 2018. The origin of squamates revealed by a Middle Triassic lizard from the Italian Alps. Nature. 557(7707), 706-709.
Xing, O'Connor, Schmitz, Chiappe, McKellar, Yi and Li, 2020. Hummingbird-sized dinosaur from the Cretaceous period of Myanmar. Nature. 579, 245-249.
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It's finally January 1, 200n and Phylonyms is published!
Ah the PhyloCode, the so-called future of biological nomenclature whose release has always kept on slipping ever more distantly into the future. After 20 years of waiting, we now have Phylonyms: A Companion to the PhyloCode, by de Queiroz et al. (2020), "a turning point in the history of phylogenetic nomenclature" according to its introduction. As the book states "Phylonyms serves as the starting point for phylogenetic nomenclature governed by the PhyloCode. According to the preamble, “This code will take effect on the publication of Phylonyms: A Companion to the PhyloCode, and it is not retroactive.” Thus, names and definitions published here have precedence over any competing names and definitions published either before (or after) the publication of Phylonyms." So for anyone invested in standardized phylogenetic nomenclature, this is it. Nothing better is coming down the pipeline in our lifetimes, so let's see what we're stuck with.
First of all, it's expensive. You can get an ebook for $222 on Amazon or a hardcover sometime after June 9th for $234. I found an electronic version for $169 plus tax on VitalSource, but you have to use their reader. It's 1323 pages though, so isn't a bad deal. That's less than five $37.95 Cretaceous Research pdfs, and I figure this is one of those historical volumes it's good to have, like Sibley and Ahlquist's bird phylogeny book.
The format is an encyclopedia-style list of clades in phylogenetic order with Registration Number, Definition, Etymology, Reference Phylogeny, Composition, Apomorphies, Synonyms, Comments and Literature Cited. Rather like my Theropod Database, so no complaints there. Well, one complaint is really more to do with the PhyloCode itself where they decided to abbreviate definitions with the non-standard del/nabla triangle symbol ∇. If you want people to start using your format, you might want to choose symbols that exist on a standard keyboard. Alt+2207 is supposed to generate it in Windows, but results in ƒ here in Blogger. Anyone know the correct Unicode numbers?
On to the substance, where Phylonyms covers all life. Dinosaurs are the last section of the book, and non-avian dinosaurs get all of four definitions-
Dinosauria R. Owen 1842 [M. C. Langer, F. E. Novas, J. S. Bittencourt, M. D. Ezcurra, and J. A. Gauthier], converted clade name
Registration Number: 194
Definition: The smallest clade containing Iguanodon bernissartensis Boulenger in Beneden 1881 (Ornithischia/Euornithopoda) Megalosaurus bucklandii Mantell 1827 (Theropoda/Megalosauroidea) and Cetiosaurus oxoniensis Phillips 1871 (Sauropodomorpha).
I'm glad we've standardized which theropod, ornithischian and sauropodomorph are used (or so I thought, see below), but otherwise there's not much to say. The caveats around which apomorphies are also found in Nyasasaurus and at least some silesaurs illustrate why apomorphy-based definitions are bad. The reference phylogeny for this and Saurischia is Lloyd et al.'s (2008) supertree, which is quite outdated and has a lot of artifacts from being a supertree.
Saurischia H. G. Seeley 1888 [J. A. Gauthier, M. C. Langer, F. E. Novas, J. Bittencourt, and M. D. Ezcurra], converted clade name
Registration Number: 195
Definition: The largest clade containing Allosaurus fragilis Marsh 1877 (Theropoda/Carnosauria) and Camarasaurus supremus Cope 1877 (Sauropodomorpha), but not Stegosaurus stenops Marsh 1887 (Ornithischia/Stegosauridae).
It's rather odd the same authors didn't choose the same specifiers for each dinosaurian clade as they did in the previous definition, leaving us without a neat node-stem triplet. Instead they went with the Kischlatian approach of using taxa"mentioned and figured as examples of their respective groups by Seeley (1888)." This is funny because I don't think this rationale is ever suggested in the PhyloCode, whereas Dinosauria and Saurischia are actually the official examples used for Recommendation 11F encouraging node-stem triplets ("If it is important to establish two names as applying to sister clades regardless of the phylogeny, reciprocal maximum-clade definitions should be used in which the single internal specifier of one is the single external specifier of the other, and vice versa"). Specifically- "If one wishes to define the names Saurischia and Ornithischia such that they will always refer to sister clades, Saurischia might be defined as the largest clade containing Megalosaurus bucklandiiMantell 1827 but not Iguanodon bernissartensisBoulenger in Beneden 1881, and Ornithischia would be defined as the largest clade containing Iguanodon bernissartensis but not Megalosaurus bucklandii. To stabilize the name Dinosauria as referring to the clade comprising Saurischia and Ornithischia, Dinosauria should be defined as the smallest clade containing Megalosaurus bucklandii and Iguanodon bernissartensis."
Ornithoscelida and its consequences are mentioned, but I'm glad more time is not taken up with it as I expect the hypothesis to fall away as Baron's phylogenetic mistakes are not followed by future authors.
Sauropodomorpha F. R. von Huene 1932 [M. Fabbri, E. Tschopp, B. McPhee, S. Nesbitt, D. Pol, and M. Langer], converted clade name
Registration Number: 295
Definition: The largest clade containing Saltasaurus loricatus Bonaparte and Powell 1980 (Sauropodomorpha) but not Allosaurus fragilis Marsh 1877 (Theropoda) and Iguanodon bernissartensis Boulenger in Beneden 1881 (Ornithischia).
I dislike the use of Saltasaurus as the internal specifier, which is a holdover of Sereno's weird use of deeply nested OTUs when others would be more historically relevant and/or eponymous. Fabbri et al. defend the choice because "Fossil specimens referred to Saltasaurus loricatus are abundant, the species is well known, and its phylogenetic position is consistent among phylogenetic analyses", but this would be even more true for e.g. Camarasaurus supremus used in Saurischia's definition. The other specifiers are a mix of those in Dinosauria's and Saurischia's definition, so there's absolutely no consistency. The reference phylogeny is Otero et al.'s (2015) Sefapanosaurus description using Yates' matrix, so is fine.
There's a rare error in the comments for this entry. Fabbri et al. state "Segnosaurus galbinensis from the Cretaceous was briefly thought to be a relatively early diverging sauropodomorph (Paul, 1984; Gauthier, 1986; Olshevsky, 1991). More material referable to that species and the discovery of closely related taxa later showed that Segnosaurus galbinensis is part of the Therizinosauria", but material of S. galbinensis besides that initially recovered in the 1970s is not known.
Theropoda O. C. Marsh 1881 [D. Naish, A. Cau, T. R. Holtz, Jr., M. Fabbri, and J. A. Gauthier], converted clade name
Registration Number: 216
Definition: The largest clade containing Allosaurus fragilis Marsh 1877 (Theropoda) but neither Plateosaurus engelhardti Meyer 1837 (Sauropodomorpha) nor Heterodontosaurus tucki Crompton and Charig 1962 (Ornithischia).
Here we've chosen two completely different specifiers for Sauropodomorpha and Ornithischia, so again we have no consistency. The reference phylogeny is Cau (2018), which is ideal.
What about the rest? It's a HUGE volume, and obviously most of Pan-Biota is outside my area of expertise. One obvious issue is the wildly varying coverage of different clades. Apparently nobody could be bothered with the vast majority of vertebrates (euteleosts) or animals (insects, except one definition for Trichoptera), and Molluska doesn't even get a definition. But we do get several entries for edrioasterid taxa down to subfamily-level, generally obscure Paleozoic echinoderms. Closer to dinosaurs, there's nothing at all for pan-crocs, but we get an entry for Pterosauromorpha for which only Scleromochlus is given as a plausible non-pterosaurian example (perhaps wrongly- Bennett, 2020).
Then there are the apomorphy-based definitions which will cause headaches in the future. Look at Apo-Chiroptera- "Definition: The clade for which the unique modifications of the hand, forearm, humerus, scapula, hip, and ankle (see Diagnostic Apomorphies) associated with flapping flight, as inherited by Vespertilio murinus Linnaeus 1758, are apomorphies." Then you go down to the nine listed sets of Diagnostic Apomorphies like "Modification of the scapula: Scapular spine originates at the posterior edge of the glenoid fossa. Long axis of scapular spine offset 20–30 degrees from axis of rotation of the humeral head. Scapular spine reduced in height—acromion process appears more strongly arched and less well supported than in other mammals. Presence of at least two facets in infraspinous fossa." These are all going spread out as more stem bats are discovered, and indeed the authors already note "Simmons and Geisler (1998) included the absence of claws on wing digits III-V with this suite of modifications; however, the presence of claws on all the wing digits of Onychonycteris suggests that claws were present primitively in Apo-Chiroptera."
Ungulata is defined by Archibald as "The least inclusive crown clade containing Bos primigenius Bojanus 1827 (= Bos taurus Linnaeus 1758) (Artiodactyla) and Equus ferus Boddaert 1785 (= Equus caballus Linnaeus 1758) (Perissodactyla), provided that this clade does not include Felis silvestris Schreber 1777 (= Felis catus Linnaeus 1758) (Carnivora), Manis pentadactyla Linnaeus 1758 (Pholidota), Vespertilio murinus Linnaeus 1758 (Chiroptera), or Erinaceus europaeus Linnaeus 1758 (Lipotyphla)." But this doesn't exist in molecular studies, including those of ultraconserved elements, which consistently place carnivorans, pangolins and bats closer to perrisodactyls. So this is likely to be a historical footnote, as well established molecular relationships end up trumping morphological relationships in every example I know of.
Finally, we get Pan-Lepidosauria for the total group of lepidosaurs, which has been Lepidosauromorpha for over thirty years. Yet Archosauromorpha is retained as "The least inclusive clade containing Gallus (originally Phasianus) gallus (Aves) (Linnaeus 1758), Alligator (originally Crocdilus) mississippiensis (Daudin 1802) (Crocodylia), Mesosuchus browni Watson 1912 (Rhynchosauria), Trilophosaurus buettneri Case 1928 (Trilophosauridae), Prolacerta broomi Parrington 1935 (Prolacertiformes), and Protorosaurus speneri von Meyer 1830 (Protorosauria)" even though Pan-Archosauria is also used for the total group of archosaurs, traditionally the definition of Archosauromorpha. I agree our new Archosauromorpha deserved a name for being a generally recognized group, whereas whether e.g. choristoderes or sauropterygians fell out closer to lizards or birds is highly unstable. But I would have rather kept the tradition of -omorpha for the stem clades and gave this a new name.
Overall, I'm not very impressed for something 20 years in the making that intends to be so important. How do you contradict your own example for choosing specifiers in four papers, where two share the same author list, the other two share another author (Fabbri), and each of those shares an author with both of the first two (Langer and Gauthier)? And one of those is an editor for the volume. Nothing could be negotiated in over two decades? But it's what we have to work with now, and in the name of consistancy I'll adopt the definitions proposed. Now to see what happens when RegNum goes online.
References- Lloyd, Davis, Pisani, Tarver, Ruta, Sakamoto, Hone, Jennings and Benton, 2008. Dinosaurs and the Cretaceous terrestrial revolution. Proceedings of the Royal Society B. 275, 2483-2490.
Otero, Krupandan, Pol, Chinsamy and Choiniere, 2015. A new basal sauropodiform from South Africa and the phylogenetic relationships of basal sauropodomorphs. Zoological Journal of the Linnean Society. 174, 589-634.
Cau, 2018. The assembly of the avian body plan: A 160-million-year long process. Bollettino della Società Paleontologica Italiana. 57(1), 1-25.
Bennett, 2020. Reassessment of the Triassic archosauriform Scleromochlus taylori: Neither runner nor biped, but hopper. PeerJ. 8:e8418.
de Queiroz, Cantiono and Gauthier, 2020. Phylonyms: A Companion to the PhyloCode, 1st Edition. Taylor & Francis Group. 1323 pp.
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The Unecessary Death of Steneosaurus
Not a dinosaur, but a new paper on the classic crocodylomorph Steneosaurus exemplifies a troubling trend in recent vertebrate taxonomy. Johnson et al. (2020) reexamine the original material of Steneosaurus, an aquatic croc from the Jurassic of France. It hadn't been seriously looked at since the 1860s, so this is one of my favorite kinds of paleontology papers- restudying a fragmentary old specimen in a modern light. What do they find?
We first get a detailed recount of its history, with two decades as Cuvier's "tête à museau plus allongé" (= head with a more elongated snout; I have to praise the authors for translating all the French to English, even in our spoiled era of Google Translate it saves time), before it was named Steneosaurus rostro-major by Geoffroy Saint-Hilaire in 1825. Eudes Deslongchamps and son tackled it in the 1860s, where they viewed the specimen as too poorly preserved and so "stated that the taxon to represent the genus Steneosaurus should be either ‘Steneosaurus’ megistorhynchus Eudes-Deslongchamps, 1866, or ‘Steneosaurus’ edwardsi Eudes-Deslongchamps, 1868c." Ha! You don't get to just take somebody's genus and affix your new species as its type. They were the last to examine the specimen in detail however, making that a pretty bad note to end on.
Johnson et al. then reexamine the type snout of Steneosaurus, correcting the species name by eliminating the hyphen, officially making it the lectotype, noting Steel had determined the posterior skull to be Metriorhynchus, and illustrating and redescribing the specimen. Excellent work and very well done. After eliminating Mycterosuchus nasutus, 'Steneosaurus' leedsi, 'S.' heberti and Lemmysuchus and other machimosaurins based on numerous dissimilar characters, the authors come to the contemporaneous 'Steneosaurus' edwardsi.
"As mentioned before, this was a second species that Eudes-Deslongchamps (1867–69) considered identical to S. rostromajor. These two taxa share a combination of features including:
1. A subcircular, moderately interdigitating premaxilla-maxilla suture.
2. Maxillae ornamented with irregular grooves.
3. A shallower mediolateral compression of the posterior maxillae, as opposed to ‘S.’ heberti (MNHN.F 1890-13).
4. Horizontally flat posterior premaxilla in lateral view.
5. Deep anterior and mid-maxillary reception pits that gradually become shallower towards the posterior maxilla.
6. Subcircular to circular alveoli that remain relatively the same size throughout the maxilla.
7. Teeth with well-pronounced enamel ridges at the base."
Well how cool is that? They put in the hard work, found the matching more complete specimens, and now we have Steneosaurus edwardsi as a junior synonym of S. rostromajor, giving us a good look at what Steneosaurus really was after two hundred years.
Lectotype of Steneosaurus rostromajor (MNHN.RJN 134c-d) in dorsal (A, B) and ventral (C, D) views. (after Johnson et al., 2020).
But no.
Johnson et al. immediately say "it is important to note that many of these characters may, in fact, be related to sexual dimorphism, ontogeny and intraspecific variation." True, but that could be said for basically every character supposed to diagnose Mesozoic croc genera, or theropod genera, pterosaur genera, etc.. Unless you have some specific example like 'enamel ridges have been shown to develop with age and both S. rostromajor and S. edwardsi are larger than S? leedsi or S? heberti with weak ridges', then it's just hand-waving. And no, Johnson et al. never develop such an argument for one of those characters, let alone all seven.
Next, we get "In addition to the sexual dimorphism/ontogeny problem, one of the critical issues about MNHN.RJN 134c-d is that it is poorly preserved." Sure, but you were still able to perform many comparisons. Again, the authors never say any of their seven characters are taphonomic, so it's another objection without substance.
Yet the worst rationale for rejecting Steneosaurus is "in reality, the name Steneosaurus is extremely impractical. It was used for many metriorhynchid specimens (e.g. ‘Steneosaurus’ gracilis, ‘Steneosaurus’ palpebrosus and ‘Steneosaurus’ manselii) during much of the 19th century, largely in part due to Cuvier’s metriorhynchid skull region (MNHN. RJN 134a-b) being attributed to the teleosauroid rostral section (MNHN.RJN 134c-d). Indeed, the concise, classical definition of ‘Steneosaurus’ as we interpret it today was not given until the work of both Eudes-Deslongchampses (1868c, 1867–69)"
Substitute Megalosaurus in there to see how ridiculous it is. That has had over 45 species assigned to it, and was named in the 1820s but didn't have a modern concept associated with it until the 1980s. When Johnson et al. lament that for Steneosaurus "rather than comparing characters outright, comparison is by process of elimination (or the question of ‘what features does this specimen lack?’)", that perfectly describes the Megalosaurus paralectotype dentary.
"After the Eudes-Deslongchampses’ treatment, what was left was an undiagnostic, chimeric type specimen for S. rostromajor (MNHN.RJN 134) and the genus Steneosaurus was redefined using a new type species that was not accepted by some researchers. In addition, since the Eudes-Deslongchampses, there has been no attempt to rectify this taxonomic nightmare;"
You just showed it was diagnostic, Steel long ago got rid of the chimaeric portion, Eudes-Deslongchamps' stupid attempts to name new type species have no relevance, and you have done the work to finally rectify this taxonomic nightmare.
"Due to these three significant factors (uncertainty of variable characters, poor preservation and
unreasonable name), we have concluded that S. rostromajor, and therefore ‘Steneosaurus’ (MNHN.RJN 134c-d), cannot be confidently assigned to an existing teleosauroid species."
Nope, you just showed it can be assigned to the same species as S. edwardsi.
Actually, I correct myself. THIS is the worst rationale for rejecting Steneosaurus- "In addition, MNHN.RJN 134c-d was initially diagnosed based on significant orbital and temporal characteristics (from the metriorhynchid MNHN.RJN 134a-b), along with generic rostral ones. Because the skull material is now known to be from a metriorhynchid, this ‘hybrid type specimen’ factor adds to the doubtful validity of Steneosaurus. According to Article 23.8 of the ICZN Code, ‘a species-group name established for an animal later found to be a hybrid (Art. 17) must not be used as the valid name for either of the parental species (even if it is older than all other available names for them)’ (this also signifies that the species name rostromajor is itself invalid). As such, MNHN.RJN 134c-d serves as an undiagnostic specimen; we, therefore, consider MNHN.RJN 134c-d to be a nomen dubium and, as such, Steneosaurus is treated as an undiagnostic genus."
If the term "parental species" didn't tip you off, Article 23.8 applies to hybrid individuals (those resulting from different species interbreeding), not type specimens chimaerically combined from multiple species. The Article doesn't even say what Johnson et al. think- it says a name for a hybrid can't be used for either of the species that bred to make it, so that e.g. even if a mule's scientific name was erected prior to that of horse's or ass's, it can't be the name for horse or ass. And indeed even the cited Article 17 says that hybrids and chimaeras can be the basis of valid names- "The availability of a name is not affected even if 17.1. it is found that the original description or name-bearing type specimen(s) relates to more than one taxon, or to parts of animals belonging to more than one taxon; or 17.2. it is applied to a taxon known, or later found, to be of hybrid origin..."
If Johnson et al.'s interpretation were right, there goes Gojirasaurus, Protoavis, Chuandongocoelurus, Chilantaisaurus, Fukuiraptor, Coelurus, Alectrosaurus, Dakotaraptor, etc..
Before the big reveal, we have in the Conclusion what can only be described as a lie- "Through character comparison-and-elimination, the only taxon with which MHNH.RJN 134c-d could hypothetically be referred to is ‘S.’ edwardsi, but the two do not share any clear autapomorphic characters or a unique combination of characters."
What are your seven listed characters if not "a unique combination of characters"? Does any other teleosaurid have them? If not, they are unique. In any case, we get the motivation for dumping Steneosaurus twice at the end of the paper-
"We believe that establishing teleosauroid taxonomy from the beginning with a series of ‘clean’ type species/specimens, with every nomenclatural act correctly formulated, is the best course of action, which we will highlight in a forthcoming paper (Johnson, 2019)."
"We believe that establishing teleosauroid taxonomy from the beginning with a series of ‘clean’ type species/specimens, with every nomenclatural act correctly formulated, is the best course of action. This will necessitate a revised teleosauroid taxonomy, in which species previously referred to the genus Steneosaurus are given new generic names. This work will be published by us in a separate contribution, based on the comprehensive teleosauroid phylogenetic analysis in Johnson’s PhD thesis (2019)."
Basically everything I hate about a current trend in vertebrate paleontology- just throw out old specimens and dishonor their authors who correctly reported what was new at the time to come up with your own names. At least dumping Stegosaurus armatus or designating a neotype for Allosaurus fragilis could be claimed to save time and effort actually analysing the types, if you don't want to do the science to figure out if armatus is actually different from stenops or if fragilis can be distinguished from Saurophaganax. But Johnson et al. already did all the hard work and found Steneosaurus edwardsi was S. rostromajor, they would just rather use Johnson's new genus name for the taxon.
And their reasons are just grasping at straws. 'Sure we identified these seven charactesrs uniquely shared by Steneosaurus rostromajor and S. edwardsi, but uhh.. could be sexually dimorphic? Or anything could be individual variation. Or ontogenetic? Lots of things turn out to be ontogenetic. Plus it's broken. Sooo broken. Sure we could evaluate characters, but who wants a taxon whose holotype isn't pristine? Plus a lot of people had stupid ideas about Steneosaurus over the past two hundred years. What do us scientists do when we have a complicated situation to resolve that was only partially understood historically? Trash their names and give yourselves credit for new genera.' Thus Steneosaurus gets the eternal identity of "all evidence points to it being Johnsonosaurus edwardsi, but ehhh... we just sort of ignore it now as Teleosauridae indet. and it's forgotten."
To conclude, Steneosaurus is really outside my wheelhouse. But if Johnson et al.'s philosophy spreads, we're in danger of losing a lot of historical taxonomy and deserved credit to lazy or selfish authors. Just look at Microraptor for example, whose holotype of M. zhaoianus lacks a decent skull. Some decades down the line, what if cranial differences support various Jiufotang species and someone's like 'the postcranial proportions are unique between the M. zhaoianus type and M. hanqingi, but I want a complete type specimen, so Microraptor is an invalid undiagnostic nomen dubium, and instead I propose Mybetterraptorgenus hanqingi and M. gui.' Just hope they don't pull a Wilson and Upchurch and claim 'Microraptor is invalid and co-ordinate suprageneric Linnean taxa must likewise be abandoned' and replace Microraptorinae with Mybetterraptorgenusinae.
References- Johnson, 2019. The taxonomy, systematics and ecomorphological diversity of Teleosauroidea (Crocodylomorpha, Thalattosuchia), and the evaluation of the genus 'Steneosaurus'. PhD Thesis, University of Edinburgh. 1062 pp.
Johnson, Young and Brusatte, 2020. Emptying the wastebasket: A historical and taxonomic revision of the Jurassic crocodylomorph Steneosaurus. Zoological Journal of the Linnean Society. 189(2), 428-448.
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The Arguable Identity of Paraxenisaurus
Hi everyone. In light of Cau's recent post on the supposed new Mexican deinocheirid "Paraxenisaurus normalensis" (Serrano-Brañas et al., 2020), I figured I'd check the taxon out to see what I thought.
The first thing you might note are the quotation marks surrounding its name, as this is yet another example of authors not including an lsid or reference to ZooBank in their electronic descriptions. ICZN Article 8.5.3. states names published electronically must "be registered in the Official Register of Zoological Nomenclature (ZooBank) (see Article 78.2.4) and contain evidence in the work itself that such registration has occurred", and the pre-print is said to be in preparation for Volume 101 of Journal of South American Earth Sciences, cited as August 2020. Thus it gets to join the ranks of "Thanos" and "Trierarchuncus"as theropods that will eventually be validly named this year. But at least it's not stuck in the purgatory of twelve Scientific Reports Mesozoic theropods, which will never be physically published and thus will remain invalid unless outside action is taken.
One of the big takeaways from Cau's blogpost is that "I am doubtful about the possibility of referring these elements [the paratypes] to the same species of the holotype, since there are very few superimposable elements among the three specimens. Therefore, there is a risk that Paraxenisaurus , - understood as the sum of all three specimens - is a chimera." After reading the paper, Andrea REALLY undersold this critique. Here are the specimen materials lists, with the overlapping elements highlighted in matching colors-
(BENC 2/2-001; proposed holotype) proximal manual phalanx II-2 or III-3, partial astragalocalcaneum, partial metatarsal II, phalanx II-1 (115 mm), proximal phalanx II-2, partial metatarsal III, proximal phalanx III-3, distal metatarsal IV, phalanx IV-1 (104 mm), phalanx IV-3 (67 mm), phalanx IV-4 (45 mm), partial pedal ungual IV
(BENC 1/2-0054) distal metacarpal I, proximal phalanx I-1, partial manual ungual I, distal metacarpal II, distal phalanx II-2
(BENC 1/2-0091) several proximal caudal central fragments (66, 75, 76 mm), proximal metacarpal II, partial metacarpal III, distal femur (155 mm trans), distal metatarsal IV
(BENC 1/2-0092) several distal caudal vertebrae (70, 71 mm)
(BENC 30/2-001) pedal ungual II, pedal ungual III
As you can see, there's only one strict overlap, with BENC 1/2-0091 sharing a distal metatarsal IV with the proposed holotype, found ~14 kilometers away. The paper lists no proposed apomorphies or unique combination of characters for distal metatarsal IV, and indeed the description states they preserve largely non-overlapping portions-
"In the holotype, the distal articular surface is fragmented (Figures 11a1 and 11a2); but in the referred specimen (BENC ½-0091), this surface is nicely preserved and has a non-ginglymoid outline (Figures 11b1 and 11b2). The medial condyle is mostly preserved in the holotype (Figure 11a3), but in the referred specimen it is completely broken (Figure 11b3). Conversely, the lateral condyle is broken in the holotype (Figure 11a4), but is well preserved in the referred specimen (Figure 11b4). Collateral ligament fossae are well developed on both condyles and have approximately the same size and depth (Figures 11a3 and 11b4). In cross-section, the shaft of metatarsal IV near the distal end is thicker dorsoventrally than wide."
Needless to say, metatarsal IV has a shaft which is deeper than wide in all ornithomimosaurs, and the preserved ligament fossae are on opposite sides in each specimen (medial in proposed holotype, lateral in 1/2-0091). Below is a figure comparing the two Mexican specimens with Ornithomimus velox, with 1/2-0091 flipped so that all are comparable as left elements. I don't see anything the "Paraxenisaurus" specimens have in common that could diagnose a taxon.
Left distal metatarsal IV of (left to right) intended "Paraxenisaurus normalensis" holotype BENC 2/2-001, intended "Paraxenisaurus normalensis" paratype BENC 1/2-0091 (right element flipped), and Ornithomomus velox holotype YPM 542 in (top to bottom) dorsal, lateral, ventral and medial views ("Paraxenisaurus" after Serrano-Brañas et al., 2020; Ornithomimus after Claessens and Loewen, 2016).
While no other elements are exactly matched, referred specimen BENC 30/2-001 does include pedal unguals II and III, while the intended holotype has pedal ungual IV. These are again from different localities, although closer this time (~2.8 km), and this time we have characters listed in the diagnosis-
"(9) distinctively broad and ventrally curved pedal unguals that angled downward with respect to the proximal articular surface and depending on the digit, the proximodorsal process becomes slightly enlarge and changes its position from nearly horizontal to mostly vertical, adopting a lipshaped appearance; and (10) pedal unguals with a rounded, large foramen on the medial side* and a deep ventral fossa that surrounds a strongly developed, ridge-like flexor tubercle."
Pedal unguals of (left to right) intended "Paraxenisaurus normalensis" paratype BENC30/2-001 right digit II, right digit III and intended "Paraxenisaurus normalensis" holotype BENC 2/2-001 left digit IV in (top to bottom) right, left, proximal and dorsal views (after Serrano-Brañas et al., 2020). Green lines point to supposedly natural median foramen
Ventral curvature is plesiomorphic, the unguals of BENC 30/2-001 are not broader than other ornithomimosaurs', and ventral angling with the proximal end held vertically is common in theropods and present in e.g. Garudimimus and Beishanlong. The proximodorsal process "changing its position" is using a difference between 30/2-001's mostly horizontal processes and the intended holotype's more vertical process as character, which in itself presupposes they are the same taxon. The ventral fossa surrounding a ridge-like flexor tubercle is also present in Harpymimus, Garudimimus, Beishanlong and large Dinosaur Park unguals (NMC 1349, RTMP 1967.19.145) and is not shown in the intended holotype but is claimed to be "partially broken." This leaves the medial foramen, which might be a valid character in unguals III and IV (II is damaged in that area), but might also be taphonomic, as there are many other small circular areas of damage (e.g. center of proximal surface of ungual IV). While the two unguals in 30/2-001 are similar to each other, that of the intended holotype is more strongly curved, has that smaller more dorsally angled proximodorsal process, is wider in proximal view, and lacks the expanded ventral half characteristic of ornithomimosaurs that is present in the other specimen. But even if these two pedal unguals are correctly referred, they are all that's present in specimen BENC 30/2-001. So they get us nowhere in determining caudal, manual (besides proximal manual phalanx II-2 or III-3) or femoral morphology.
The final issue I noticed was the emphasis on "Paraxenisaurus" having a first pedal digit. This would ironically be unlike Deinocheirus, but plesiomorphically shared with Nedcolbertia, "Grusimimus", Garudimimus, Beishanlong, Archaeornithomimus and Sinornithomimus. The character state is based on metatarsal II, where "a facet on the posterior surface of the distal quarter of this shaft, indicates the presence of an articulation area for metatarsal I." The figure shows a longitudinal groove extending down the posterior center of distal metatarsal II, which as anyone who has scored taxa for Clarke's bird matrix could tell you, is not how non-birds attach their hallux to the metatarsus. Hattori (2016) for instance writes in Allosaurus "there is no attachment scar corresponding to the metatarsal I fossa on either medial or plantar aspect of MT II" and in Citipati "there is no obvious attachment scar of MT I on either medial or plantar aspect of MT II." Serrano-Brañas et al. state "in Garudimimus brevipes ... the attachment site is also placed in the same area as in Paraxenisaurus normalensis", but the feature in Garudimimus is a raised scar with sharp medial demarcation from the shaft. As Middleton (2003) recognized, this scar is for the m. gastrocnemius, specifically the m. gastrocnemius pars medialis (Carrano and Hutchinson, 2002), and I'll note it's present even in Gallimimus which lacks pedal digit I (Osmolska et al., 1972: Plate XLIX Fig. 1b). "Paraxenisaurus"'s groove is then more likely to be the m. flexor digitorum longus II tendon, which "passed through the ventral groove in its respective metatarsal to insert serially on each of the pedal phalanges" in e.g. Tyrannosaurus (Carrano and Hutchinson, 2002).
Left metatarsal II in ventral view of (left to right) intended "Paraxenisaurus normalensis" holotype BENC 2/2-001 (after Serrano-Brañas et al., 2020; yellow arrow points to supposed articulation for metatarsal I), Garudimimus brevipes holotype IGM 100/13 (after Kobayashi and Barsbold, 2005; line points to supposed articulation for metatarsal I), Gallimimus bullatus ZPAL MgD-I/94 (after Osmolska et al., 1972), and Tyrannosaurus rex FMNH PR2081 (after Brochu, 2003).
What exactly is "Paraxenisaurus"? Comparison is hindered by the specimens being figured mixed together, and the figures are not in numerical order in the preprint, being shown in the order of- 1, 10-19, 2, 20-23, 3-9. In addition, the scale bars vary within the same figure (e.g. phalanx IV-1 is proximally ~61 mm wide in figure 14a but ~93 mm wide in figure 14e) and the listed measurements are different yet (e.g. IV-1 is listed as 83 mm wide). Thus any composite reconstruction is necessarily approximate. The supposed manual element is too fragmentary to give much information, but it is of the appropriate size and shape to be a proximal pedal phalanx I-1. This would make more sense preservationally since the other material preserved in the specimen is all from the tarsus and pes. It's a shame the astragalocalcaneum is not described better or figured in more views, as the dorsal (= proximal?) perspective has many broken surfaces and edges, so that e.g. the small calcaneum might be preservational. The fused proximal tarsals are like ceratosaurs, deinocheirids (Deinocheirus plus Hexing), alvarezsaurids and caenagnathids. Having any sense of the ascending process morphology could tell us much. Metatarsal II is not obviously deeper than wide, unlike ornithomimosaurs (except Harpymimus; unreported in deinocheirids), but like carcharodontosaurids, therizinosauroids, some oviraptorids and velociraptorines. The proximal outline of metatarsal III would at first glance appear to be the strangest thing about this material, being reconstructed as strictly dorsoventrally oval unlike all(?) other theropods. Tilting it and adding a posterior tapered tip results in a close match to Majungasaurus however (see figure below). If it is an unreduced proximal metatarsal III, tyrannosauroids, most ornithomimosaurs, alvarezsauroids and pennaraptorans would be excluded. Proximal phalanx II-2 lacks a proximoventral heel, so is not from a deinonychosaur. The pedal phalanges are too elongate to be therizinosauroid, and the pedal ungual is too broad. Phalanges are not as dorsoventrally compressed as Mapusaurus, and as noted above they lack the ventrolateral shelves found in ornithomimosaurs. Abelisaurid phalanges seem similar however. I wonder if we have a case like Camarillasaurus or probably Dandakosaurus involving misidentified elements making the specimen seem stranger than it really was, with so many edges of supposed metatarsal III dotted to indicate incompleteness that it could actually be metatarsal II or IV. Certainly nothing connects this specimen with Deinocheirus. As per the numerous errors illustrated by Hartman et al. (2019) nobody should trust Choiniere et al.'s scorings in any case. The Lori matrix recovers "Paraxenisaurus" as a ceratosaur closest to Aucasaurus as far as taxa with well preserved feet are concerned, but also doesn't include characters particular to ceratosaurs and isn't great with pedal characters in general. So I would place the specimen as Neotheropoda incertae sedis (or even indet.) pending a better description of the tarsus and of the real bone surfaces on supposed proximal metatarsal III.
Pes of "Paraxenisaurus normalensis" holotype (center) in dorsal view compared to Majungasaurus crenatissimus composite (left) and Deinocheirus mirificus referred specimen IGM 100/127 (right). Colored proximal view of "Paraxenisaurus" is after Serrano-Brañas et al., with reoriented metatarsal III as per my interpretation shown above that. Note "Paraxenisaurus" elements were scaled using their scale bars, whereas scaling to listed measurements results in different proportions, so those should be seen as approximate. "Paraxenisaurus" after Serrano-Brañas et al. (2020), Majungasaurus after Carrano (2007) and Deinocheirus after Lee et al. (2014).
References- Osmólska, Roniewicz and Barsbold, 1972. A new dinosaur, Gallimimus bullatus n. gen., n. sp. (Ornithomimidae) from the Upper Cretaceous of Mongolia. Palaeontologica Polonica. 27, 103-143.
Carrano and Hutchinson, 2002. Pelvic and hindlimb musculature of Tyrannosaurus rex (Dinosauria: Theropoda). Journal of Morphology. 253, 207-228.
Brochu, 2003. Osteology of Tyrannosaurus rex: Insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Society of Vertebrate Paleontology Memoir. 7, 138 pp.
Middleton, 2003. Morphology, evolution, and function of the avian hallux. PhD thesis, Brown University. 147 pp.
Carrano, 2007. The appendicular skeleton of Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. In Sampson and Krause (eds.). Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. SVP Memoir 8, 164-179.
Lee, Barsbold, Currie, Kobayashi, Lee, Godefroit, Escuillie and Tsogtbaatar, 2014. Resolving the long-standing enigmas of a giant ornithomimosaur Deinocheirus mirificus. Nature. 515, 257-260.
Kobayashi and Barsbold, 2005. Reexamination of a primitive ornithomimosaur, Garudimimus brevipes Barsbold, 1981 (Dinosauria: Theropoda), from the Late Cretaceous of Mongolia. Canadian Journal of Earth Sciences. 42(9), 1501-1521.
Claessens and Loewen, 2016 (online 2015). A redescription of Ornithomimus velox Marsh, 1890 (Dinosauria, Theropoda). Journal of Vertebrate Paleontology. 36(1), e1034593.
Hattori, 2016. Evolution of the hallux in non-avian theropod dinosaurs. Journal of Vertebrate Paleontology. 36(4), e1116995.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247.
Cau, 2020 online. http://theropoda.blogspot.com/2020/04/paraxenisaurus-un-deinocheiride.html
Serrano-Brañas, Espinosa-Chávez, Maccracken, Gutiérrez-Blando, de León-Dávila and Ventura, 2020. Paraxenisaurus normalensis, a large deinocheirid ornithomimosaur from the Cerro del Pueblo Formation (Upper Cretaceous), Coahuila, Mexico. Journal of South American Earth Sciences. 101, 102610.
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Is Falcatakely a bird?
So this week we got the description of the new Maevarano skull Falcatakely forsterae (O'Connor et al., 2020). It's a pseudo-toucan, with a long, tall snout formed mostly by the maxilla unlike modern birds. O'Connor et al. recover it as an enantiornithine using Brusatte et al.'s TWiG analysis and O'Connor's bird analysis. Our problem is that we only have the anteroventral skull preserved, so no braincase, mandible or postcrania. And being a Maastrichtian deposit in Africa, we don't have the most detailed coelurosaur record. Add in the fact beaks are known to evolve fast on islands (as Madagascar was even back then) and we have a potential problem on our hands.
Follow your nose to the exciting topic of African coelurosaur diversity. Reconstructed holotype skull of Falcatakely forsterae (UA 10015) (after O'Connor et al., 2020).
The Lori analysis places Falcatakely in two potential positions- a therizinosaurid or an omnivoropterygid. The former doesn't make much sense biostratigraphically, but there is a Maevarano synsacrum of the correct size (FMNH PA 741) that was claimed to share characters with Sapeornis by O'Connor and Forster (2010). So there's a possibility. Forcing Falcatakely to be an enantiornithine as in O'Connor et al.'s analyses requires six more steps. Most of the discordant characters relate to the beak, but the wide laterotemporal fenestra would be odd in an enantiornithine. While I was writing this, Andrea Cau published a post on this topic and reported that he recovered Falcatakely as a noasaurid, which would be quite the phylogenetic jump, but it only takes three more steps in the Lori matrix, so is more parsimonious than the enantiornithine option. It falls out as an elaphrosaurine, so could relate to e.g. Afromimus. The non-beak contradictory characters here include a lack of antorbital fossa lacrimal foramen, long posterior lacrimal process and triradiate palatine, which seem more convincing to me. Additional evidence against these latter two positions is the absence of small ceratosaur (Masiaksaurus is twice as big) or large enantiornithine (Maevarano elements are much smaller) postcrania. Andrea reported (translated) "It takes 6 further steps to place it in Coelurosauria, and in that case it is a basal dromaeosaurid: interesting in that regard to note that Rahonavis , known from the same Formation, has also been hypothesized to be a basal dromaeosaurid. Can we rule out that Falcatakely is the (still unknown) skull of Rahonavis? The estimated dimensions of the two animals coincide." Forcing Falcatakely to be Rahonavis only requires one more step, which is pretty impressive. In their (amazing) osteology, Forster et al. (2020) refer an isolated dentary "found near the Rahonavis holotype (its precise location was not recorded during excavation)" which does not match Falcatakely's upper jaw, being upcurved and extensively toothed. But it is similar to other unenlagiines like Buitreraptor and Austroraptor. So much as we have two synsacrum types at this size, unenlagiine-like Rahonavis and Sapeornis-like, we have two cranial types, unenlagiine-like and Falcatakely which is Sapeornis-like in the combination of reduced maxillary dentition, triradiate palatine, modified/reduced antorbital fossa, anteriorly limited naris and strong postorbital-jugal articulation.
Referred dentary of Rahonavis ostromi (FMNH PA 740) as a transparent CT reconstruction (after Forster et al., 2020).
Thus my best guess is that Falcatakely is a basal avialan belonging to the same taxon as FMNH PA 741. But this comes with a huge chunk of salt as it is so far removed temporally and geographically from potential comparable sister taxa. Which is actually a common problem with this part of the tree, as shown by Balaur (= Elopteryx?), Hesperonychus, Imperobator and even Rahonavis itself. We compare these Late Cretaceous taxa to our far more complete Early Cretaceous Jehol record and say Hesperonychus is sorta like Microraptor, Falcatakely is kind of like Sapeornis and Balaur is Jeholornis-grade, but North America, Africa and Europe had their own avialan fauna for 70 million years before them that we're basically unaware of. If the only alvarezsauroid we had was Mononykus' holotype, could we place it correctly as a basal maniraptoran? If the only oviraptorosaur we had was Citipati's skull, would we recover that correctly as the sister taxon of Paraves? I think that's the position we find ourselves in with Falcatakely, and that future discoveries of African small theropods will lead to new interpretations.
References- O'Connor and Forster, 2010. A Late Cretaceous (Maastrichtian) avifauna from the Maevarano Formation, Madagascar. Journal of Vertebrate Paleontology. 30(4), 1178-1201.
Cau, 2020 online. theropoda.blogspot.com/2020/11/falcatakely-eterodossia-e-pluralismo.html
Forster, O'Connor, Chiappe and Turner, 2020. The osteology of the Late Cretaceous paravian Rahonavis ostromi from Madagascar. Palaeontologia Electronica. 23(2):a31.
O'Connor, Turner, Groenke, Felice, Rogers, Krause and Rahantarisoa, 2020. Late Cretaceous bird from Madagascar reveals unique development of beaks. Nature. DOI: 10.1038/s41586-020-2945-x
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Antarctic Ichthyornis solved
So I've been doing some major updates to the Database for what will probably be a New Years upload, including the ornithuromorph section. One rather sad entry as it currently stands is the Antarctic Ichthyornis-
I? sp. (Zinsmeister, 1985)
Late Cretaceous
Seymour Island, Antarctica
Reference- Zinsmeister, 1985. 1985 Seymour Island expedition. Antarctic Journal of U.S. 20, 41-42.
Now with Googling I found the original paper online, which allowed only a bit of improvement-
I? sp. (Zinsmeister, 1985)
Late Maastrictian, Late Cretaceous
Lopez de Bertodano Formation, Seymour Island, Antarctica
Material- several elements
Comments- Zinsmeister (1985) states "several small bones tentatively identified as belonging to the Cretaceous bird Ichthyornis were discovered in the upper Cretaceous Lopez de Bertodano formation."
Reference- Zinsmeister, 1985. 1985 Seymour Island expedition. Antarctic Journal of U.S. 20, 41-42.
So I saw that Zinsmeister worked with Chatterjee in the 80s, who found the Polarornis holotype in the same place two years before that. I emailed Chatterjee about it, who replied-
"It was misidentified in the field. These were some shark teeth."
Mystery solved! But can we do better? Here's an Ichthyornis tooth-
Right eleventh dentary tooth of Ichthyornis dispar (YPM 1450) (after Field et al., 2018).
And here's the array of shark teeth from the Lopez de Bertodano Formation of Seymour Island (from a January 2011 expedition). Can we find any easily confusable matches?
Chondrichthyan teeth from the Lopez de Bertodano Formation (scale 10 mm) (after Otero et al., 2014).
I think the circled 16 and 17 are pretty decent matches for a field identification, though much larger if compared directly. Figures 6-17 are all identified as Odontaspidae indet., which covers any morphology similar to Ichthyornis. Add in the fact that they were by far the most abundant teeth recovered (8 samples versus 1-3 for the other taxa), and I think we have a nice solution on our hands.
I wonder how many other weird records are out there that are based on initial misidentification but stay in the literature because nobody ever publishes a correction?
References- Otero, Gutstein, Vargas, Rubilar-Rogers, Yury-Yañez, Bastías and Ramírez, 2014. New chondrichthyans from the Upper Cretaceous (Campanian-Maastrichtian) of Seymour and James Ross islands, Antarctica. Journal of Paleontology. 88(3), 411-420.
Field, Hanson, Burnham, Wilson, Super, Ehret, Ebersole and Bhullar, 2018. Complete Ichthyornis skull illuminates mosaic assembly of the avian head. Nature. 557, 96-100.
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"Megalosaurus" cloacinus and more - September 2021 Database Update
Hi everyone. I realize it's been ten months since the last post, and that's because I've been prioritizing updating the Database over writing blogs. As a compromise of sorts and to not force people to constantly check the Database updates page, I decided to try out posting when I update including features that could have made it into their own blog post.
One thing I've been doing is working my way through Skawiński et al.'s (2017) paper on Polish Triassic dinosaur reports, which in addition to unnamed fragments, also led to the creation of entries for two supposed Megalosaurus species. silesiacus is a generic carnivorous archosauriform tooth too early to be dinosaurian, while cloacinus has been used for basically every carnivorous archosaur tooth from Rhaetian beds of Germany. The interesting thing about the latter is that workers apparently forgot that it was based on lost teeth described by Quenstedt, not the SMNS tooth figured 47 years later by Huene.
"Zanclodon" silesiacus Jaekel, 1910
= Megalosaurus silesiacus (Jaekel, 1910) Kuhn, 1965
Early Anisian, Middle Triassic
Lower Gogolin Formation, Lower Muschelkalk, Poland
Holotype- (University of Griefswalden/Göttinger coll.; lost?) tooth (24x12x5 mm)
Referred- ?(Geological Museum of the Polish Geological Institute-National Research Institute coll.) tooth (Skawiński, Ziegler, Czepiński, Szermański, Tałanda, Surmik and Niedźwiedzki, 2017)
?(Silesian University of Technology, Faculty of Mining and Geology coll.) tooth (37 mm) (Surmik and Brachaniec, 2013)
Comments- Jaekel (1910) noted (translated) "a dinosaur tooth from the lower shell limestone of Upper Silesia, which would probably be the oldest known dinosaur tooth to date. It comes from the Chorzov strata of the lower shell limestone of Gogolin, Upper Silesia, and came to me through the kindness of engineer Fedder in Opole. The crown shown is 24 mm high, 12 mm wide and 5 mm thick, so it is quite strongly compressed and slightly curved backwards. Its edge is extremely finely serrated (Fig. 16 A). I call the form, which for the time being cannot be specified generically, Zanclodon silesiacus. The only difference between [phytosaur Mesorhinosuchus] and this tooth form lies in the fact that the former is somewhat thicker, somewhat less bent back, and that no notch can be detected on the edge." He referred it to Megalosauridae, and Kuhn (1965) later referred it to the genus Megalosaurus. Carrano et al. (2012) correctly noted "could be considered as Theropoda indet., but we cannot rule out the possibility that it represents a 'rauisuchian' archosaur." Surmik and Brachaniec (2013) describe a tooth from Gogolin Quarry in which "a poor state of preservation makes it impossible to identification of the presence of edge serration, however it still shows a slightly curvature and specific both sides flattening" and identify it as seemingly archosaurian. Skawiński et al. (2017) listed this and another tooth labeled as Megalosaurus silesiacus as other material of Zanclodon silesiacus. The latter tooth is stated to be serrated mesially and distally with a density of 12 per 5 mm. They describe the holotype tooth as "Probably lost" and "lost", and place all three teeth as Archosauromorpha indet.. They are more specifically referred to the Teyujagua plus archosauriform clade here given the recurvature and small serrations, as authors from Kuhn onward have noted plesiomorphic theropod teeth are difficult to distinguish from several clades of archosauriforms (e.g. erythrosuchids, euparkeriids) known from the Anisian. The age is far too early for Megalosaurus or another neotheropod, and the presence of serrations is unlike Zanclodon, so neither genus is appropriate. It should also be noted the three Gogolin teeth differ in shape with the Silesian University specimen less recurved and less tapered than the other two, while the Polish Geological Institute specimen is shorter than the holotype and less concave distally. This could be positional variation, but given the lack of proposed synapomorphies could easily represent multiple taxa.
References- Jaekel, 1910. Ueber einen neuen Belodonten aus dem Buntsandstein von Bernburg. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin. 5, 197-229.
Kuhn, 1965. Fossilium Catalogus 1: Animalia. Pars 109: Saurischia. Ysel Press. 94 pp.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Surmik and Brachaniec, 2013. The large superpredators' teeth from Middle Triassic of Poland. Contemporary Trends in Geoscience. 2, 91-94.
Skawiński, Ziegler, Czepiński, Szermański, Tałanda, Surmik and Niedźwiedzki, 2017 (online 2016). A re-evaluation of the historical 'dinosaur' remains from the Middle-Upper Triassic of Poland. Historical Biology. 29(4), 442-472.
Holotype tooth of "Zanclodon" silesiacus (University of Griefswalden/Göttinger coll.; lost?) in labial (A), basal section (B) and more apical section (C) (after Jaekel, 1910).
"Megalosaurus" cloacinus Quenstedt, 1858
= Plateosaurus cloacinus (Quenstedt, 1858) Huene, 1905
= Gresslyosaurus cloacinus (Quenstedt, 1858) Huene, 1932
= Pachysaurus cloacinus (Quenstedt, 1858) Huene, 1932
Rhaetian, Late Triassic
Exter Formation, Germany
Syntypes- (lost) two teeth
Referred- ?(GPIT and SMNS coll.) many teeth (Huene, 1905)
?(SMNS 52457) tooth (~25x11x? mm) (Huene, 1905)
?(SMNS coll.) teeth (Roemer, 1870)
? seven teeth (Miller Endlich, 1870)
Norian-Rhaetian?, Late Triassic
'Lisów Breccia', Poland
?(University of Wroclaw coll.; lost) two teeth (Roemer, 1870)
Early Hettangian, Early Jurassic
Calcaire de Valognes, Manche, France
?(University of Caen coll.; destroyed) tooth (Rioult, 1978)
Comments- Quenstedt (1858) originally described (translated) "barb-shaped teeth, which are sharp and finely serrated on the concave side, but rounded and smooth on the convex side" with a large mesioapically placed wear facet that makes that edge look straight in side view. He also figures a smaller tooth which has mesial serrations apically that transition to a rounded edge basally. These teeth do not share any obvious synapomorphies and differ in elongation (height/FABL ~300% vs. 138%) and transverse thickness (42% vs. 75% of FABL), so may not belong to the same taxon. Miller Endlich (1870) figured seven teeth from the type locality, stating (translated) they "are mostly flat teeth, slightly curved on one side, with fine serrations on the sharp inner edge. The convex side, the back, does not seem to be serrated, but it is not certain." The figured teeth show a wide range of variation, with figure 13 in particular being stout and unrecurved with large serrations, similar to the Lucianosaurus paratype and similarly referrable to Archosauromorpha incertae sedis. The other teeth have small serrations, with 14 and 18 being straight and 15-17 and 19 being recurved, with 14, 18 and 15 being progressively more transversely compressed. As with the syntypes, these exhibit variation which could be positional or interspecific, and share no obvious characters that connect them to each other or the syntypes. Roemer (1870) wrote (translated) "In the Stuttgart Museum I saw teeth from the bone breccia of Bebenhausen near Tubingen, which show the same fine serration of the side edges as the teeth described by Quenstedt, but are not curved in a sickle shape, but are straight. It is very likely that these latter teeth belong to the same dinosaur as the crooked teeth. With these straight teeth from Bebenhausen, the tooth shown in FIGS. 4 and 5 from the Lisów Breccia from Lubsza near Woźniki completely coincides. The double-edged tooth, which is very delicately and regularly notched at the edges, shows a more strongly curved (outer) and a less curved (inner) surface, both of which are smooth except for a very fine, irregular wrinkle. There is also a much smaller tooth of the same type from the same location." The straight Bebenhausen teeth sound similar to Miller Endlich's figures 14 and 18, although the illustrated straight tooth from Lubsza differs from these in having an increased amount of mesiodistal expansion basally. The Lubsza tooth also has this marked basal expansion labiolingually, and both types of root expansion are atypical of dinosaurs, suggesting this is some other type of vertebrate. Dzik and Sulej (2007) suggested it "may have belonged to a phytosaur" without evidence but Skawiński et al. (2017) stated "phytosaur fossils have not been found in the upper Keuper strata in Silesia" and instead placed it in Archosauromorpha indet.. While this could merely mean phytosaurs were rare in that strata, phytosaur teeth don't seem to have expanded roots either (e.g. Nicrosaurus), and it could even be a fish tooth which often have these types of root expansion. Huene (1905) listed the species as "Plateosaurus" cloacinus within Theropoda, stating it includes Rhaetian dentary "Zanclodon cambrensis". In 1908 he places it in Plateosauridae within Theropoda and states (translated) "The originals can no longer be found. The Tübingen collection still has several teeth from Bebenhausen and Schloßlesmuehle, which can be reconciled well with [Quenstedt's] fig. 12 (l. c.), but are larger. The serrations are coarse and short, the mesial carina does not extend all the way to the base." He illustrated a tooth in figure 274 as "From the Rhaetian Bonebed of Bebenhausen near Tübingen. Tooth in nat. Size. The tip is missing. Original in the natural history cabinet in Stuttgart." Regarding cambrensis, Huene states "The teeth have the greatest resemblance to Plateosaurus cloacinus both in the whole shape and in the serrations. Whether it is really the same or just a very similar species, of course, cannot be decided with certainty given the scanty material", which is not explicit enough to evaluate given published details. Huene later (1932) assigns cloacinus to Teratosauridae within Carnosauria, listed as both Pachysaurus cloacinus (pg. 6) and Gresslyosaurus cloacinus (pg. 72, 114). Steel (1970) calls it Gresslyosaurus cloacinus within Plateosauridae. Buffetaut et al. (1991) mentions "A tooth referred to Megalosaurus cloacinusQuenstedt, from the Lower Hettangian of the Calcaire de Valognes at Valognes (Manche), [which] has been mentioned by Rioult (1978a) as having been destroyed by an air raid on the University of Caen in 1944." Without additional details, it can only be said that the timing suggests a neotheropod. Carrano et al. (2012) incorrectly claimed SMNS 52457, apparently the tooth in Huene's (1908) figure 274, is "the holotype and only specimen" of cloacinus, when Huene stated it was only one of "Many teeth ... in the stone quarries of the Schoenbuch (e.g. Bebenhausen, Schloesslesmuehle), Wuerttemberg; in the university collection in Tubingen and in the natural history cabinet in Stuttgart", and that Quenstedt's originals were lost. SMNS 52457 could be made into a neotype, but this must be done explicitly (ICZN Article 75.3) and so has not been acc
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https://dinotoyblog.com/stegouros-deluxe-by-collecta/
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Stegouros (Deluxe by CollectA) – Dinosaur Toy Blog
|
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2024-06-15T22:22:34+00:00
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https://dinotoyblog.com/stegouros-deluxe-by-collecta/
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Most ankylosaurs are classified as either Ankylosauridae or Nodosauridae. Ankylosaurids are easily distinguishable by their wide, blocky heads and tails terminating in solid bone clubs, and include the likes of Ankylosaurus itself, Euoplocephalus, Jinyunpelta, Pinacosaurus, and Zuul. By contrast, nodosaurids like Animantarx, Borealopelta, Edmontonia, Gastonia, and Sauropelta are distinguished by their narrower, more triangular heads and clubless tails. But there’s also a third family, Parankylosauria. They lived in Gondwana during the Cretaceous period and they were much smaller and more basal than the ankylosaurids and nodosaurids. Examples include Antarctopelta from Antarctica, Kunbarrasaurus from Australia, and the subject of this review, Stegouros from Chile.
Discovered in 2018 and formally named and described in 2021, Stegouros elengassen was only around two metres in length at most and roughly the size of a domestic dog, making it one of the smallest ankylosaurs known. CollectA’s 2023 Deluxe Stegouros, however, is quite the gargantuan toy, measuring slightly over 24 cm long, 9 cm tall at the hips, and 7 cm wide. This makes it their biggest ankylosaur toy to date, even bigger than the Deluxe Ankylosaurus as shown below. Some collectors may be put off by its great size, but personally, scale has never been that much of an issue for me. Also, I think it’s kind of funny how it dwarfs every other ankylosaur in my collection when it was quite the opposite in reality.
The main colour on this toy is medium brown with dark wash, dull sandy yellow for the underbelly, dark brown for the osteoderms and spikes, dark grey for the claws and beak, a dull pink tongue, and pinkish grey streaks over the glossy black eyes. Overall, it’s pretty standard coloration for an ankylosaur toy, nothing out of the ordinary, but as always, it looks perfectly plausible.
The Stegouros is posed with its front limbs bent and its feet firmly planted. Both the head and the tail are raised and turning to the right, and the mouth is wide open. Presumably it is either attempting to ward off a predator or engaging in combat with another of its kind. Again, quite a few other ankylosaur toys are posed similar to like this, but it always does work for them. Between its pose and its size, this armoured beast looks quite formidable indeed.
In addition to being one of the smallest known ankylosaurs, Stegouros is notable for a number of anatomical features which are well reflected on this toy. The head is relatively elongated, V-shaped when viewed from above, and very big, proportionally bigger than those of most other ankylosaurs. There are no scutes projecting from the back of the head, but there are large, flattened scales that probably would have afforded it some decent protection.
The entire body is covered in fine pebbled scales and as usual, there are heavy wrinkles around the limb joints and hanging down from the neck and flanks. The limbs are fairly long, indicating that Stegouros could have tried to flee from large predators as an alternative to as standing and fighting. The hind feet have three large toes and one tiny inner toe, all with claws, while the front feet have five toes with claws on all but the outermost. Multiple rows of keeled osteoderms run down the neck, back, and tail, and on the limbs as well. The armour overall more closely resembles an ankylosaurid’s than a nodosaurid’s. All in keeping with the known fossil material, which, happily, is nearly complete.
And that brings us at last to the Stegouros‘ tail. Instead of a rounded club or no weapon at all, the tip has seven pairs of large, flattened osteoderms that are merged together to form a most unique structure. The shape strongly resembles either the profile of a pinecone or an ancient Aztec macuahuitl, which is a wooden club equipped with sharpened shards of obsidian along the outer edge. Antarctopelta may have possessed a similar tail as well. Indeterminate theropod remains have been recovered from the Dorotea Formation where Stegouros was found, so it probably had need for a good defensive weapon. Its primary purpose, though, was likely intraspecific conflict. In any case, a blow from that tail would undoubtedly have been painful. And can you imagine if this animal was the same size as Ankylosaurus? Yikes!
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https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4131922/
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A New Basal Ankylosaurid (Dinosauria: Ornithischia) from the Lower Cretaceous Jiufotang Formation of Liaoning Province, China
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[
"Fenglu Han",
"Wenjie Zheng",
"Dongyu Hu",
"Xing Xu",
"Paul M. Barrett"
] |
2014-08-25T00:00:00
|
A new ankylosaurid, Chuanqilong chaoyangensis gen. et sp. nov., is described here based on a nearly complete skeleton from the Lower Cretaceous Jiufotang Formation of Baishizui Village, Lingyuan City, Liaoning Province, China. Chuanqilong chaoyangensis ...
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https://www.ncbi.nlm.nih.gov/coreutils/nwds/img/favicons/favicon.ico
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PubMed Central (PMC)
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https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4131922/
|
PLoS One. 2014; 9(8): e104551.
PMCID: PMC4131922
PMID: 25118986
A New Basal Ankylosaurid (Dinosauria: Ornithischia) from the Lower Cretaceous Jiufotang Formation of Liaoning Province, China
, 1 , 2 , * , 2 , 3 , 4 , 2 and 5
Fenglu Han
1 Faculty of Earth Sciences, China University of Geosciences (Wuhan), Wuhan, China
2 Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China
Find articles by Fenglu Han
Wenjie Zheng
2 Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China
3 University of Chinese Academy of Sciences, Beijing, China
Find articles by Wenjie Zheng
Dongyu Hu
4 Paleontological Institute, Shenyang Normal University, Shenyang, China
Find articles by Dongyu Hu
Xing Xu
2 Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China
Find articles by Xing Xu
Paul M. Barrett
5 Department of Earth Sciences, Natural History Museum, London, United Kingdom
Find articles by Paul M. Barrett
Peter Dodson, Editor
1 Faculty of Earth Sciences, China University of Geosciences (Wuhan), Wuhan, China
2 Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China
3 University of Chinese Academy of Sciences, Beijing, China
4 Paleontological Institute, Shenyang Normal University, Shenyang, China
5 Department of Earth Sciences, Natural History Museum, London, United Kingdom
University of Pennsylvania, United States of America
Competing Interests: The authors have declared that no competing interests exist.
Conceived and designed the experiments: XX. Performed the experiments: FLH. Analyzed the data: FLH. Contributed reagents/materials/analysis tools: XX DYH WJZ. Contributed to the writing of the manuscript: FLH PB XX.
Copyright © 2014 Han et al
This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are properly credited.
Associated Data
Supplementary Materials
GUID: 12FE3422-7959-49A2-997E-3369508387AE
Data Availability Statement
The authors confirm that all data underlying the findings are fully available without restriction. All relevant data are within the paper and its Supporting Information files.
Abstract
A new ankylosaurid, Chuanqilong chaoyangensis gen. et sp. nov., is described here based on a nearly complete skeleton from the Lower Cretaceous Jiufotang Formation of Baishizui Village, Lingyuan City, Liaoning Province, China. Chuanqilong chaoyangensis can be diagnosed on the basis of two autapomorphies (glenoid fossa for quadrate at same level as the dentary tooth row; distally tapering ischium with constricted midshaft) and also a unique combination of character states (slender, wedge-like lacrimal; long retroarticular process; humerus with strongly expanded proximal end; ratio of humerus to femur length = 0.88). Although a phylogenetic analysis places Chuanqilong chaoyangensis as the sister taxon of the sympatric Liaoningosaurus near the base of the Ankylosauridae, the two taxa can be distinguished on the basis of many features, such as tooth morphology and ischial shape, which are not ontogeny-related. Chuanqilong chaoyangensis represents the fourth ankylosaurid species reported from the Cretaceous of Liaoning, China, suggesting a relatively high diversity in Cretaceous Liaoning.
Introduction
Ankylosauria is a group of quadrupedal herbivorous dinosaurs characterised by parasagittal rows of osteoderms on the dorsolateral surface of the body and a heavily armored skull [1]. The earliest records of the group have been reported from various Early or Middle Jurassic localities, and include Bienosaurus lufengensis, Tianchisaurus nedegoapefererima, and Sarcolestes leedsi [2]–[4], although all of these records have been considered either nomina dubia or dubiously referable to Ankylosauria [1], [5]. Definitive ankylosaur taxa are known to occur from the Late Jurassic (e.g., Gargoyleosaurus from western North America: [6]) to the end of the Cretaceous and their remains have been reported from all continents except Africa [1]. In Liaoning Province, China, three ankylosaurian species have been reported: Liaoningosaurus paradoxus from the Lower Cretaceous Yixian Formation [7] and Crichtonsaurus bohlini [8] and C. benxiensis [9] from the Upper Cretaceous Sunjiawan Formation. Liaoningosaurus was originally considered to be a possible nodosaurid [7], but a recent study suggests that it is a basal ankylosaurid [10]. C. bohlini and C. benxiensis are also referable to the Ankylosauridae (and are probably basal ankylosaurines) [9], [10].
Here, we describe a fourth ankylosaur species from Liaoning based on a specimen collected from the Lower Cretaceous Jiufotang Formation. This specimen preserves a nearly complete skeleton, and it provides new information on the morphology and taxonomy of the Ankylosauria.
Materials and Methods
The permits for this research were obtained from the Chaoyang Jizantang Paleontological Museum of Liaoning, China. All of the materials described herein were collected from a single quarry by local farmers. Locality information was provided by staff at the Chaoyang Jizantang Paleontological Museum. The fossils are two-dimensionally preserved and visible in ventral view only. The material includes a skull and articulated postcranial material referable to a single individual. The skull and mandible are nearly complete, but have been strongly compressed dorsoventrally. The vertebral column includes the cervicals, dorsals, sacrals, and most of the caudals, but most of them are disarticulated. Both of the fore- and hind limbs are well preserved and articulated. Armor is preserved around the entire body but is only visible in ventral view.
In order to compare the ratios of humerus/tibia length to femur length with those of other ankylosaurs, the data were analysed in the software package SPSS 16.0 using the linear fit function. The best fit lines, regression equation and R2 values are presented in the Results.
The phylogenetic position of Chuanqilong chaoyangensis was inferred using parsimony analysis. The new taxon was incorporated into a previously published data matrix built to examine ankylosaurian interrelationships [10]. Liaoningosaurus paradoxus was also re-scored in the matrix based on our firsthand observations of the holotype specimen (Text S1). The modified data matrix consists of 170 characters and 52 taxa. The matrix was analyzed using TNT [11], and all of the characters were treated as equally weighted and unordered. The analysis was conducted using a heuristic search with 1000 replicates. TBR branch swapping was employed and 100 parsimonious trees were saved per replicate. A reduced consensus analysis was performed to identify wildcard taxa within TNT to provide maximum phylogenetic resolution for the new taxa [11]. Standard bootstrap values (absolute frequencies) were calculated using a traditional heuristic search with 1000 replications. Bremer supports were calculated by running the script “Bremer. run” automatically.
Nomenclatural Acts
The electronic edition of this article conforms to the requirements of the amended International Code of Zoological Nomenclature (ICZN), and hence the new names contained herein are available under that Code from the electronic edition of this article. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information viewed through any standard web browser by appending the LSID to the prefix “http://zoobank.org/”. The LSID for this publication is: urn:lsid:zoobank.org:pub:9D60475B-FA91-464B-8EBF-D335582AE23E. The electronic edition of this work was published in a journal with an ISSN (1932–6203), and has been archived and is available from the following digital repositories: PubMed Central (http://www.ncbi.nlm.nih.gov/pmc), LOCKSS (http://www.lockss.org).
Institutional Abbreviations
AMNH, American Museum of Natural History, New York, USA; BXGM, Benxi Geological Museum, Liaoning Province, China; CEUM, Prehistoric Museum, College of Eastern Utah, Price, Utah, USA; CJPM, Chaoyang Jizantang Paleontological Museum; DMNH, Denver Museum of Natural History, Denver, Colorado, USA; IVPP, Institute of Vertebrate Paleontology and Paleoanthropology, Beijing, China; LPM, Liaoning Paleontological Museum, Shenyang, Liaoning Province, China; MPC, Mongolian Paleontological Center, Ulaanbaatar, Mongolia; MTM, Hungarian Natural History Museum, Budapest, Hungary; MU, University of Missouri, Columbia, Missouri, USA; NHMUK, Natural History Museum, London, UK; PIN, Paleontological Institute, Russian Academy of Sciences, Moscow, Russia; QM, Queensland Museum, Brisbane, Australia; ROM, Royal Ontario Museum, Toronto, Canada; SMU, Southern Methodist University, Dallas, Texas, USA; TMP, Royal Tyrrell Museum of Palaeontology, Drumheller, Alberta, Canada; UALVP, University of Alberta Laboratory for Vertebrate Paleontology, Edmonton, Alberta, Canada; YPM, Yale Peabody Museum, New Haven, Connecticut, USA.
Results
Systematic Paleontology
Dinosauria Owen, 1842 [12]
Ornithischia Seeley, 1887 [13]
Thyreophora Nopcsa, 1915 [14]
Ankylosauria Osborn, 1923 [15]
Ankylosauridae Brown, 1908 [16]
Chuanqilong gen. nov. urn:lsid:zoobank.org:act:76DD6D3F-F23C-4AC0-B480-20AC73B50279
Type Species
Chuanqilong chaoyangensis gen. et sp. nov. urn:lsid:zoobank.org:act:EE6564A0-33AE-4CC7-B4EA-8CD682E1EB43
Etymology
The generic name is derived from Chinese Chuanqi (legendary, referring to western Liaoning providing a spectacular assemblage of Mesozoic terrestrial fossils) + long (dragon). The specific name is derived from the broader geographical area including the type locality.
Holotype
CJPM V001, a nearly complete skeleton missing only the distal portion of the caudal series. The specimen is housed in the Chaoyang Jizantang Paleontological Museum. A cast of the holotype specimen is housed in the Institute of Vertebrate Paleontology and Paleoanthropology as IVPP FV 1978.
Locality and Horizon
Baishizui Village, Goumenzi County, Lingyuan City, Liaoning Province, China ( ); the quarry is in the Jiufotang Formation (Lower Cretaceous, Aptian) [17] ( ). A detailed stratigraphical investigation of this quarry is required to establish its relationships to other exposures of the Jiufotang Formation in the area.
Diagnosis
An ankylosaur distinguished from other ankylosaurs by two autapomorphies: the glenoid fossa for the quadrate is at the same level as the dentary tooth row; and the distally tapering ischium is constricted at midshaft length. Chuanqilong also differs from all other ankylosaurians in having the following unique combination of character states: presence of a long retroarticular process (differs from all other ankylosaurians except Gargoyleosaurus); presence of a slender, wedge-like lacrimal (differs from all other known ankylosaurians except Minmi); ratio of humerus to femur length of 0.88 (notably higher than in most known ankylosaurians except Hungarosaurus and Liaoningosaurus); the width of the proximal end of the humerus is half of the length of the humeral shaft (substantially different from that of Liaoningosaurus, in which this ratio is 0.38); presence of subtriangular unguals (absent in all other ankylosaurs except Liaoningosaurus and Dyoplosaurus).
Description and Comparisons
The holotype skeleton is exposed mainly in ventral view ( ) and as result numerous anatomical details are not visible. In addition, the presence of some elements obscures large portions of the other bones present limiting the amount of information available. Nevertheless, preservation is generally good. Although the specimen represents a large animal (approximately 4.5 m in total body length; see for measurements of the holotype skeleton), it is likely to be a juvenile individual based on several features. For example, the vertebral centra are not fused to their neural arches in all visible vertebrae including cervicals, dorsals, and caudals. In addition, the sacral ribs are not fused to the sacral centra. Consequently, an adult individual is likely to have been greater than 4.5 m in length. Jurassic ankylosaurians are mostly relatively small in size with body lengths no greater than 4 meters. For example, Mymoorapelta and Gargoyleosaurus each have lengths of approximately 3 m [18], [19]. By comparison, most Cretaceous ankylosaurians have body lengths greater than 5 m. For example, the primitive ankylosaurid Cedarpelta has an estimated body length of 7.5–8.5 m [20].
Table 1
BoneL/RLengthWpWdWm1* 2* ScapulaL——19.59——R40—199——HumerusL3518145.5——R35———20—UlnaL18135———R17136———RadiusL236.56.54——R23884——FemurL4018179——R4020189——TibiaL—15197——R3612176——FibulaL32653.5——R35653.5——IliumL76———2015R————2015IschiumL421875——R411975.5——Metacarpal IL7.54.54.5———Metacarpal IIL844———Metacarpal IIIL845———Metacarpal IVL724———Metatarsal IIL137.553.5——Metatarsal IIIL15.57.563.5——Metatarsal IIR136.553.5——Metatarsal IIIR15663——Metatarsal IVR13561.5——
Skull
The skull and mandibles are strongly compressed dorsoventrally. The skull is triangular in ventral view, with a transverse width that was probably greater than its length, as in ankylosaurids ( , ) [21]. The maxilla is partially exposed in lateral view and exhibits a shallow, flattened buccal emargination. A large, triangular antorbital fossa or fenestra seems to be present, located in the caudodorsal region of the maxilla ( ) in lateral view. An antorbital fenestra is also present in juvenile ankylosaurids such as Liaoningosaurus and Pinacosaurus, though it is replaced by a small concavity in adult Pinacosaurus [7], [22]. A small antorbital fenestra is also present in the probably adult Minotaurasaurus [22],[23], whereas it is either unknown or absent in all other ankylosaurians [1]. It seems likely that the presence of an open antorbital fenestra in Chuanqilong chaoyangensis is due to its juvenile status. However, the orbit is circular in outline, and relatively small in comparison to skull length, contrary to what would be expected in a juvenile individual. The lacrimal is slender and wedge-shaped ( ), forming the rostral margin of the orbit, as in the basal ankylosaurid Minmi, whereas it is sub-rectangular in other known ankylosaurians, such as Pinacosaurus and Cedarpelta [1]. A long and dorsoventrally compressed supraorbital is visible in lateral view, contacting the lacrimal rostroventrally. Caudally, the postorbital is damaged and only partially exposed. The ‘squamosal’ may be composed of both the squamosal and a portion of the postorbital. It is subrectangular in outline and ornamented with subparallel grooves, as in Minmi [24] ( ). The left quadrate is exposed in rostral view. It is long and straight with a rectangular head, and there is no indication that the quadrate head was fused with the squamosal, thereby differing from the condition seen in nodosaurids [25]. Below the quadrate head, the shaft is transversely expanded to form a wide, shallow, and rostrally-opening depression. The quadrate constricts ventral to this point and is narrowest at midshaft length. A crescentic depression is present on the cranioventral surface of the quadrate for reception of the quadratojugal. The pterygoid process is thin, short, and sub-triangular in outline. The transversely expanded ventral end is composed of two well-defined mandibular condyles, which are separated by a shallow groove ventrally. The medial condyle is transversely wider and extends further ventrally than the lateral condyle, as in most other ankylosaurians. No other features of the skull are visible.
Mandible
The paired mandibular rami are preserved separately and both are visible in medial view only ( , ). The predentary is missing. The mandible is long and shallow, as in other basal ankylosaurids, but it differs from other taxa in the apparent absence of an osteoderm from its ventral margin. The lack of an osteoderm in this area may be either due to incomplete preservation, suggesting that they were not fused to the mandibular bones and thereby supporting the suggestion that the holotype Chuanqilong was not fully grown. In adult individuals of other ankylosaurians, such as Pinacosaurus and Saichania, osteoderms are fused to the lateral surface of the mandible [22], [26]. Alternatively, an osteoderm, if present, might be restricted to the lateral-most corner of the mandible and hence obscured from view (this condition is present in juvenile individuals of Pinacosaurus grangeri e.g., IVPP V16853; V. Arbour, pers. comm.), which would also represent a juvenile feature. The dentary tooth row is straight or slightly arched dorsally, but it is not as strongly sinusoidal as those of derived ankylosaurians, such as Euoplocephalus [27] or Pinacosaurus [26]. In dorsal view, the rostral end of the dentary tooth row is curved medially. At least 20 alveoli are present. The ventral margin of the mandible is relatively straight in its rostral and middle regions, but curves caudodorsally in its caudal part. The right dentary symphysis is preserved and is slightly downturned, short, robust, and sub-triangular in cross-section. The Meckelian canal is open, long, and deep. The coronoid eminence is prominent and projects above the level of the dentary tooth row, as in nodosaurids, whereas the coronoid eminence is situated at approximately the same level as the dentary tooth row in ankylosaurids, including Pinacosaurus [26], Euoplocephalus [27], and Ankylosaurus [28]. The splenial and prearticular are missing, exposing the adductor fossa, which is large and located below the coronoid eminence. The articular is small and oval in outline in lateral view. The retroarticular process is long and slender, as in Gargoyleosaurus, but differs from those of all other ankylosaurians, which possess relatively short and deep retroarticular processes [19]. Unusually, the glenoid fossa is situated in a relatively dorsal position, lying at approximately the same level as the dentary tooth row, unlike the condition present in all other ankylosaurians known from appropriate material, in which the glenoid fossa is situated at a level ventral to the dentary tooth row (e.g., Pinacosaurus: [26]). Unfortunately, this feature cannot be assessed in Liaoningosaurus [7].
Dentition
Premaxillary teeth are unknown due to breakage of the snout. Both maxillary and dentary teeth are preserved, with the former exposed labially and the latter exposed lingually.
There are at least 20 alveoli in the left maxilla. The maxillary tooth counts of most ankylosaurians are around 20: Ankylosaurus has the largest number of maxillary teeth (34–35) [28], whereas Liaoningosaurus has the smallest number (about 10: [7]). Tooth count probably increases during growth [7] and adult tooth count also varies between species [28]. The teeth and their marginal denticles are small in comparison to the size of skull, as occurs typically in ankylosaurids [21].
The four preserved rostral maxillary teeth are smaller than the caudal teeth. The crowns are as tall as their width with sub-triangular outlines, as in most ankylosaurians ( ). The base of the crown is strongly swollen with a weak cingulum, as in ankylosaurids [21]. There is no shallow notch at the base of tooth crowns, unlike the notched condition present in the ankylosaurid Crichtonsaurus [8] and the nodosaurid Edmontonia [29]. Additionally, the teeth of C. bohlini bear much larger denticles and a more distinct cingulum than those of Chuanqilong. There is no distinct primary ridge, and secondary vertical ridges and grooves are present on the labial surfaces of the tooth crowns. These ridges usually terminate apical to the cingulum, but some ridges extend across the cingulum to the basal margin of the crown. The crescentic cingula seen in some ankylosaurs, such as Texasetes, are absent. Small denticles and cusps are present on one rostral maxillary tooth crown. The denticles are small and tapering with a round cross-section at their base. However, most of the teeth do not bear these structures, though it is not clear if this absence is due to poor preservation or tooth wear. Most of the dentary teeth are missing. Those that are preserved are only partially exposed in the left dentary. Dentary teeth seem to have been similar in size and shape to the maxillary teeth.
Axial Skeleton
Several cervical and dorsal vertebrae are scattered on the slab. The cervical centra are spool-like and shorter than they are wide: their articular surfaces are all obscured. The dorsal centra are also spool-like in ventral view, slightly amphicoelous, and possess concave lateral surfaces. Dorsal centra are longer than tall. A ventral keel is absent from all of the preserved dorsal centra. One sacral vertebra is exposed on the slab in ventral view. Its centrum is wider than it is long, and its exposed articular surface is rugose, suggesting that it had not yet fused with the other sacrals. Several sacral ribs are preserved separately. They are robust and dumbbell-shaped in outline. Approximately 20 caudal vertebrae are preserved, but most of them are disarticulated. The centra of the proximal and middle caudal vertebrae are shorter than they are wide. Deep longitudinal grooves are present on the ventral surfaces of the proximal caudal vertebrae. Chevron facets are well developed, with the caudal facets more prominent than the cranial ones.
One middle caudal vertebra is well preserved. Its centrum is relatively longer and transversely narrower than those of the cranial caudals and in lateral view it has a square outline. The transverse process is reduced to a small nodular process and is located on the dorsal part of the lateral surface of the centrum. The neural spines are elongate, oriented caudodorsally, and possess arc-shaped outlines. The prezygapophyseal facets are oval in outline and face craniomedially, whereas the postzygapophyses are positioned near the tip of the neural spine and face caudolaterally.
Caudually, three additional distal caudal vertebrae are tightly articulated. Their centra are more elongate and transversely narrower than those of the proximal and middle caudals. In these vertebrae, the neural spine has merged with the postzygapophyses to form a single midline caudal process that extends caudally. The caudal process terminates cranial to the midpoint of following vertebra, as in nodosaurids, but unlike the condition in derived ankylosaurids, in which the process is longer [21]. The prezygapophyses are short and reduced in size, corresponding to the size reduction of the postzygapophyses. No chevrons are preserved.
Several ossified hypaxial tendons are present near the distal region of the tail ( ). During preservation, they have moved from their original positions and arrangement so that they are now aligned in different orientations. On the basis of the morphology of the preserved caudals, which does not conform with that of the tail club handle morphology seen in ankylosaurine taxa, a tail club knob was probably absent, as in all nodosaurids and some basal ankylosaurids (e.g., Minmi: [30]). Presence of a tail club was formerly an important diagnostic feature of ankylosaurids [21], but recent work has indicated that tail clubs may have been present in ankylosaurines only, a clade that includes Ankylosaurus [28], Euoplocephalus [31], Pinacosaurus [32], Talarurus [33], Saichania [34], Tarchia [35], Tianzhenosaurus [36], Dyoplosaurus [37], and Nodocephalosaurus [38]. It is unknown whether tail clubs were present or not in the basal ankylosaurids Crichtonsaurus [8], [9], Cedarpelta [20], Gobisaurus [39], and Shamosaurus, but the ankylosaurids Minmi, Liaoningosaurus, and Zhongyuansaurus lack tail clubs [7], [40], [41]. The likely absence of a tail club in Chuanqilong chaoyangensis adds further support to the hypotheses that the tail club is a derived feature that appears only in derived, and currently only Late Cretaceous taxa.
Pectoral Girdle and Forelimb
Scapula
Both scapulae are preserved. The scapula and coracoid are not co-ossified, contrary to the condition in most ankylosaurians (e.g., Ankylosaurus: [28]), but this may be another indication of specimen immaturity [42]. The scapula and coracoid are unfused in all known juvenile ankylosaurs, including Liaoningosaurus [7], an indeterminate nodosaurid hatchling from the Paw Paw Formation of Texas, juvenile Pinacosaurus [43], and Anoplosaurus [42], but they are fused in most adult or sub-adult individuals ( ). The scapula blade is slender, deflected caudoventrally, and has a rhomboidal outline ( , ). The dorsal margin of the scapula blade is relatively straight, whereas its ventral margin is concave. The caudal margin expands dorsoventrally. The shaft of the scapula blade is narrowest caudal to the glenoid fossa. A transverse flange is positioned along the craniodorsal margin of the scapula, as in ankylosaurids, whereas in nodosaurids the acromion is positioned ventrally, near the glenoid, and overhangs the coracoid [21]. There is no distinct enthesis present on the ventral edge of the scapula, which probably marks the insertion of the M. triceps longus caudalis (See [28]), and has been reported in derived ankylosaurids, including Crichtonsaurus [9], Ankylosaurus [28], and Euoplocephalus [31]. The absence of a distinct enthesis also suggests that CJPM 001 is not fully grown (K. Carpenter, pers. comm.). The glenoid fossa is large, oval in outline, and faces ventromedially. Both of the coracoids are missing or concealed.
Table 2
TaxonSpecimen numberFemur lengthTibia lengthHumerus lengthScapula & coracoidRatio H/FRatio T/FReference Liaoningosaurus paradoxus IVPP V125602.52.52.5unfused11.00 [7] nodosaurid scuteling from Paw Paw FormationSMU 724447.3–6.84unfused0.94– [52] Anodontosaurus lambei AMNH 526625.5190–unknown–0.75 [31] Crichtonsaurus benxiensis BXGMV0012-1322923fused0.720.91 [9] Crichtonsaurus bohlini LPM 10134.3–24unfused0.700.00 [8] cf. Pinacosaurus MPC 100/13053820.326.5–0.700.53 [66] Chuanqilong chaoyangensis CJPM 001403635unfused0.880.90this study Pinacosaurus granger PIN 614402730unknown0.750.68 [66] Animantarx ramaljonesi CEUM 6288R41.5–29.8fused0.72– [20] Gargoyleosaurus parkpinoorum DMNH 2772646.5–29.2unknown0.63– [19] Talarurus plicatospineus PIN 557-34724.833.5fused0.710.53 [33] Hungarosaurus tormai MTM 2007.2549–45.5fused0.93– [55] Euoplocephalus tucki UALVP 3151.5–37.7probablyunfused0.73– [31] Euoplocephalus tucki AMNH 540453.542.140.3fused0.750.79 [31] Polacanthus foxii NHMUK R1755334.5–unknown–0.65 [53] Dyoplosaurus acutosquameus ROM 78456.2––unknown–– [37] Scolosaurus cutleri NHMUK nR51616041.544fused0.730.69 [14] Ankylosaurus magniventris AMNH 521467–54.2fused0.81– [28] Sauropelta edwardsi AMNH 303670–49.5fused0.71– [67]
Humerus
Both humeri are well preserved, except that the ventral part of the deltopectoral crest is damaged on the left humerus. The left humerus is exposed in cranial view, and the right humerus in lateral view ( , ). The humerus is short and robust. The deltopectoral crest is large and rounded in outline in cranial view, unlike in Crichtonsaurus benxiensis which possesses a straight lateral margin [9] ( ). The deltopectoral crest and the transverse axis through the distal condyles are in the same plane, and the deltopectoral crest extends for more than half of the length of the humerus as in ankylosaurids, but unlike the condition in nodosaurids, which possess a relatively short deltopectoral crest [21]. There is no distinct separation between the humeral head and the deltopectoral crest as in most ankylosaurians, but in contrast to Cedarpelta and Ankylosaurus in which the dorsal surface of the deltopectoral crest is lower than the humeral head [28], [44] ( ). The width of the proximal end is much greater than the distal width as in most ankylosaurids except Zhongyuansaurus, in which both ends are of equal width [40]. The laterally placed radial condyle is oval and more prominent than the medial ulna condyle. The lateral epicondylar ridge is not well developed.
Ulna and Radius
Both of the ulnae and radii are complete ( , ). The olecranon process of the ulna is low and wedge-shaped as in Liaoningosaurus [7] and juvenile specimens of Pinacosaurus [45], whereas it is tall and strongly developed in most large ankylosaurians, such as Pelorolites and Cedapelta [44] ( ). The low olecranon process may represent an ontogenetically variable character of ankylosaurians [7]. The humeral notch is moderately developed. The radius is slender in comparison with the ulna. It is rod-like with a flat proximal articular surface and a rugose, convex distal end. The distal end is wider transversely than the proximal end.
Manus
The left manus contains four complete but disarticulated metacarpals and their identifications are based on the well preserved metacarpals of Peloroplites [44] and Pinacosaurus [45], [46] ( ). All of the preserved metacarpals are slender, as in Pinacosaurus [45]. Metacarpal III is the longest. Metacarpals I and II are sub-equal in length. Metacarpal IV is significantly shorter than other metacarpals. Metacarpal I is the most robust of the metacarpals, as in other ankylosaurids [1]. Metacarpals II and IV are more slender than metacarpals I and III. All of the metacarpals have expanded proximal and distal ends. The proximal articular surfaces are slightly concave, whereas the distal articular surfaces are strongly convex. There are no distinct ginglymi at the distal end. The phalanges are proximodistally short and transversely wide. The ungual phalanges are triangular in outline with sharp point in dorsal view. Their ventral surfaces are flattened and their proximal surfaces have a round outline and are slightly concave.
Pelvic girdle and hind limb
Ilium
Both ilia are well preserved and exposed in ventral view ( , ). As in other ankylosaurs, the preacetabular process rotated medially, making the ‘original’ lateral surface face dorsally, whereas the postacetabular process rotated in apposition and the original surface faces ventrally [47]. The preacetabular process is very long and transversely wide, and diverges laterally from the vertebral column at an angle of approximately 45°. The lateral margin of the preacetabular process is straight in ventral view, as in ankylosaurids, but unlike the condition in most nodosaurids, such as Sauropelta [25], Struthiosaurus [48], and Zhejiangosaurus [49], in which it is laterally curved. The postacetabular process is subtriangular in outline. It is very short, with a length less than that of the acetabulum, as in ankylosaurids [1]. The acetabulum is imperforate with a concave articular surface for accepting the femoral head, as in all ankylosaurians except Mymoorapelta [18]. The pubic peduncle is well developed with a sub-rounded profile and is dorsoventrally compressed. The ischial peduncle is rudimentary.
Ischium
The ischium is long, slender, and mediolaterally compressed ( ). The proximal end is expanded craniocaudally and contributes to half of the medial wall of the shallow acetabulum. There is no obturator process, which is absent in all ankylosaurians. The shaft of the ischium is slender and slightly curved ventrally, as in the ankylosaurid Zhongyuansaurus [40], whereas it straight in most ankylosaurids [28] and significantly curved ventrally near the distal end in nodosaurids [1]. The shaft of the ischium is unique in being narrower in its mid-shaft region and widening towards to the distal end, prior to tapering again further distally, whereas in other ankylosaurians the shaft either tapers distally along the whole shaft (e.g. Ankylosaurus [28], Sauropelta, Edmontonia [48]) or remains sub-equal in size along the whole shaft (e. g., Euoplocephalus: [31]), or is just slightly expanded at the distal end (e.g., Cedarpelta: [44]) ( ). The proximal end of the ischium is straight in lateral view. This is unlike the convex and fan-like ischium in the ankylosaurids Ankylosaurus [50] and Euoplocephalus [51], and also unlike the concave proximal ischia of the nodosaurids Struthiosaurus and Edmontonia [48]. The proximal end of the ischium lacks the medial wall present in the basal ankylosaurid Cedarpelta [20], [44].
Femur
The femur is robust and straight, as in other ankylosaurians ( , ). The femoral head is robust and expanded forming a spherical articular surface. It forms an angle of about 145° with the long axis of the femur. Both the cranial and greater trochanters are present, and they are separated from the femoral head by a prominent constriction. The cranial trochanter is slender, finger-like, and separated from the greater trochanter by a vertical cleft. The cranial trochanter is present in juveniles, such as the Paw Paw nodosaurid scuteling and Anoplosaurus, but fused with the greater trochanter in most adult ankylosaurs [52]. However, the cranial trochanter is also present in some large primitive nodosaurids, such as Polacanthus [53] and Texasetes [54]. So the presence of a cranial trochanter is likely to be a primitive character of ankylosaurids, as well as being under ontogenetic control in some taxa. The fourth trochanter is a prominent rugosity that is located distal to femoral mid-length, as in typical ankylosaurids [21].
Distally, a shallow cranial intercondylar fossa is present. A deep caudal intercondylar groove divides the medial (tibia) and lateral (fibula) condyles, and the former is slightly larger than the latter. The ratio of humerus to femur length is 0.83, similar to the condition in Ankylosaurus, but lower than the ratios in Liaoningosaurus, the indeterminate juvenile nodosaurid from Paw Paw Formation, and Hungarosaurus, and greater than those of other known ankylosaurians ( ; ). Juveniles may have proportionally longer forelimbs than adults [52]. The juvenile Liaoningosaurus and indeterminate Paw Paw nodosaurid have humerus to femur length ratios of 1.0 and 0.93, respectively, whereas the ratio is about 0.7 in most large ankylosaurians ( ). However, this ratio is also substantially higher in adult Hungarosaurus (0.92: [55]), Ankylosaurus (AMNH 5214, 0.81, [28]), and in the late juvenile Chuanqilong (0.88) ( ). This suggests that the ratio of humerus to femur length may represent a valid taxonomic difference.
Tibia
The tibia is shorter than the femur ( , ). The ratio of tibia to femur length is approximately 0.9. This is similar to the ratio in Crichtonsaurus benxiensis (0.91: [9]), Europelta carbonensis (0.91: [56]), Liaoningosaurus paradoxus (0.95: pers. observ.) and greater than in all other known ankylosaurians ( ; ). The tibia is straight, robust, and greatly expanded mediolaterally both proximally and distally. The transverse expansion of the proximal end is relatively weaker than that of the distal end in cranial view.
Fibula
The fibula is slender and slightly shorter than the tibia. The proximal end is expanded craniocaudally and compressed laterally. The whole shaft is relatively equal in size and oval in cross-section. The distal end is slightly expanded mediolaterally with a flattened cranial surface.
Proximal tarsals
The calcaneum and astragalus are not preserved, and they are inferred to have remained unfused to the distal end of the tibia. The calcaneum and astragalus are usually fused with the distal end of the tibia in most ankylosaurians [1], but they are unfused in juveniles of Anodontosaurus lambei (AMNH 5266; [31]) Liaoningosaurus [7], and Pinacosaurus [46], suggesting that this was under ontogenetic control in ankylosaurians. However, the astragalus is not fused to the distal end of the tibia in the early ankylosaurians Mymoorapelta (DMNH 15162: [57]), Peloroplites (CEUM 11319: [44]), and possibly Hylaeosaurus (NHMUK R2615: [53]), which indicates they may have been unfused primitively in adult ankylosaurians.
Pes
Metatarsals II, III, and IV are well preserved and in articulation in the right foot ( , ). The possible presence of metatarsals I and V cannot not be excluded due to the preservation of the specimen. Metatarsal III is much longer (187.5% of the length) and more robust than metacarpal III. This ratio is similar to that seen in the primitive ankylosaurid Gargoyleosaurus (184.4%: [19]), greater than in most ankylosaurians, such as Pinacosaurus grangeri (113.5%: [45]) and Talarurus plicatospineus (132.8%: [45]), but smaller than that in Liaoningosaurus, which has even longer metatarsals (more than 200%: [7]). Metatarsals II and IV are sub-equal in length, and metatarsal III is longer and more robust than the other two metatarsals. They all have expanded proximal and distal ends. The proximal end of metatarsal II is dorsoventrally deeper than it is wide transversely and it has a concave and oval articular surface for the distal tarsals. Metatarsal III has a square cross-section proximally, and metatarsal IV is transversely wider than deep craniocaudally. The distal ends of all metatarsals are transversely expanded and bear no, or very weak, ginglymi.
The unguals are robust and sub-triangular in outline with sub-rounded distal ends in dorsal view. This is unlike the pedal unguals of Liaoningosaurus, which have much sharper distal ends [7]. Coombs [51] noted that in ankylosaurs the pedal unguals are widest at a point approximately one-third of the distance from their proximal ends in juveniles, whereas in adults they are widest proximally. However, in juveniles like Liaoningosaurus and Chuanqilong, the pedal unguals are widest proximally and taper distally, contrary to this observation. Ungual shape may, therefore, be a useful character for taxonomic or systematic purposes, and similar sub-triangular unguals are also known in Dyoplosaurus [37].
Armor
Cranial armor is not visible on the slab. The cervical armor usually comprises two cervical half rings in ankylosaurids and three cervical half rings in nodosaurids. These half rings consist of a superficial layer of osteoderms fused to an underlying band of bone. The osteoderms are usually pitted and rugose, similar to body osteoderms, whereas the connecting band is usually smooth and plate-like [58]. In Chuanqilong, only one cervical half ring is present in ventral view. The band appears to be fused into a single large plate, as in ankylosaurids [58]. However, it is compressed dorsoventrally, damaged and separated into four sections ( ). The right three sections are thickened, arched dorsally, and subrectangular in outline with smooth ventral surfaces. The left section has a wide medial edge and tapers caudolaterally with a subtriangular profile, and this section has a straight craniolateral edge as also seen in Gargoyleosaurus [19], but which is unknown in other ankylosaurians. Two large armor plates are preserved in the shoulder region. They are thickened, subrectangular in outline, have flat, smooth surfaces, and are thicker along their margins than centrally. The larger plate is twice the length of the smaller one. Both of them are similar to the osteoderms of the cervical half ring, and they may represent separate cervical armor plates. A relatively small triangular armor plate is present between the proximal end of the left ulna and radius ( ). It is dorsoventrally compressed, wide at the base and tapers distally. One nearly complete oval armor plate is present near the left ischium in dorsal view, which is sharply keeled along its midline ( ). A variety of small irregular osteoderms and ossicles are preserved over the whole body in ventral view ( ), as in most ankylosaurians. Dermal armor is absent in the indeterminate Paw Paw nodosaurid ([52]: SMU 72444), the juvenile specimen of Anoplosaurus [42], and the hatchling dinosaur Propanolosaurus [52], [59]. However, dermal armor is present in the small specimen Liaoningosaurus, suggesting that the ability to produce dermal armor had already appeared by this early growth stage [7], although armor is absent in the even smaller specimen of Propanolosaurus [59].
Discussion
Ankylosauria is traditionally divided into two families, Ankylosauridae and Nodosauridae, which are distinct in many features [21]. A third group, Polacanthinae [60] or Polacanthidae [5], has been proposed, and it is normally defined as all ankylosaurians more closely related to Gastonia than to either Edmontonia or Euoplocephalus [5]. However, some phylogenetic analyses, including ours (see below) do not recover this group as a separate clade [1], [10], [26], and here we follow traditional ankylosaurian taxonomy in our discussion.
Chuanqilong chaoyangensis possesses many ankylosaurid features, including cheek teeth with a strongly swollen tooth crown with a weak cingulum, a long deltopectoral crest that extends for more than half of humeral length, a straight lateral margin of the preacetabular process, a very short postacetabular process that is shorter than the length of the acetabulum, a slender ischial shaft that is curved slightly ventrally, and a distally located fourth trochanter. However, it lacks several features shared by derived ankylosaurids, such as the presence of a tail club. This character combination suggests that Chuanqilong chaoyangensis represents a basal ankylosaurid.
In order to confirm our hypothesis regarding the systematic position of Chuanqilong chaoyangensis, we conducted a phylogenetic analysis by adding Chuanqilong chaoyangensis to a recently published dataset on ankylosaurian phylogenetic relationships [10]. Our analysis produced 15902 most parsimonious trees (MPTs), with tree lengths of 542 steps (Consistency Index = 0.34, Retention Index = 0.66). The strict consensus tree (not shown) of these 15902 MPTs lacks resolution, with the only clear result being recovery of ankylosaurid monophyly. A reduced consensus tree was calculated a posteriori which excluded seven wildcard taxa (Zhejiangosaurus, Niobrarasaurus, Hungarosaurus, Antarctopelta, Anoplosaurus, Polacanthus rudgwickensis, and Stegopelta) [61], and this shows considerably greater resolution ( ).
Our results indicate that Chuanqilong chaoyangensis is a basal ankylosaurid and that it is the sister taxon of the sympatric Liaoningosaurus. However, only two unambiguous synapomorphies support this relationship: presence of an antorbital fossa or fenestra ([10]: character 1) and scapula glenoid oriented ventrally ([10]: character 121).
It should be noted that both Chuanqilong chaoyangensis and Liaoningosaurus paradoxus are represented by specimens at a relatively early ontogenetic stage, although the much larger size, relatively smaller orbit, and higher tooth count of Chuanqilong suggest that the latter is at a more advanced ontogenetic stage. Although Chuanqilong and Liaoningosaurus are sister taxa, they can be distinguished on the basis of the following characters, which are probably not ontogenetically variable:
Cheek tooth crown morphology. In Chuanqilong, the cheek teeth are relatively small compared to the skull, and there are more than 20 maxillary teeth, whereas in Liaoningosaurus, the cheek teeth are significantly larger in comparison to the skull, and there are only approximately 10 maxillary teeth. Although Xu et al. [7] noted that the low tooth number of Liaoningosaurus may be due to its juvenile status, the ontogenetic variation in teeth number among all known ankylosaurians is less than 10. For example, the variation in Euoplocephalus and Pinacosaurus cheek teeth number is five and three, respectively [28]. Additionally, the cheek tooth crowns of Chuanqilong bear small denticles and cusps, with approximately 12 denticles per tooth, whereas in Liaoningosaurus, the denticles are large cusps that are also relatively large with respect to the size of the tooth crown, and there are approximately seven denticles per tooth. The tooth crowns of Liaoningosaurus are similar to those of another ankylosaurid from Liaoning Province, Crichtonsaurus bohlini, but differ from those of Chuanqilong.
The proximal end of the humerus is strongly expanded in comparison to humeral length in Chuanqilong (the ratio of proximal width to whole length is 0.51), whereas it is only moderately expanded in Liaoningosaurus (the ratio of proximal width to whole length is 0.38). Additionally, the distal end of the deltopectoral crest extends for more than half of the length of the humerus in Chuanqilong, as in typical ankylosaurids, whereas in Liaoningosaurus, the deltopectoral crest is less developed and does not extend to the mid-length of the humerus, as in other nodosaurids.
The lateral edge of the ilium is straight or slightly convex in ventral view in Chuanqilong, whereas it is slightly concave above the acetabulum in Liaoningosaurus.
The ischial shaft of Chuanqilong has a constriction at mid-length and tapers distally, whereas the ischial shaft is relatively equal in width along the whole length and slightly expanded at the distal end in Liaoningosaurus.
The ratios of metatarsus to metacarpus length in Chuanqilong is substantially less than that of Liaoningosaurus ( ). The metatarsus is more than twice the length of the metacarpus in Liaoningosaurus and this is probably an autapomorphy of this taxon [7]. It is unknown whether the ratio of metatarsus to metacarpus length changes during ontogeny. However, the metatarsus is less than twice the length of the metacarpus in the hatchling Propanoplosaurus ([59]: ).
The pedal unguals of Chuanqilong are widest at a point approximately one-third of the distance from the proximal end and are slightly constricted at the proximal end, whereas the pedal unguals are sub-triangular and widest at the proximal end in Liaoningosaurus [7] and Dyoplosaurus [37].
Several juvenile ankylosaurians have been recognized and provide important ontogenetic information [7], [26], [42], [51], [52], [62]. These studies indicate that some features used for species diagnosis are probably under ontogenetic control, such as some fusion characters, including fusion of the scapula and coracoid, fusion of the calcaneum and astragalus, and fusion of the cranial and greater trochanters.
Ontogenetic variation may affect phylogenetic reconstruction (e.g. [63]). Many derived features found in adult specimens are rudimentary or undeveloped in juvenile specimens, making the latter appear more basal than adult individuals in phylogenetic analyses. Therefore, ideally, ontogenetically variable characters should be excluded from phylogenetic analysis or such analyses should be based upon adult specimens only. However, many ankylosaurians are only partially preserved and it has been difficult to document their ontogenetic variation. Euoplocephalus and Pinacosaurus, which are known from multiple individuals, may provide more insights into this problem, but the taxonomy of Euoplocephalus has been controversial and many formerly referred specimens are now thought to represent other distinct taxa [31]. In order to retain as many taxa in our analysis as possible, we were unable to exclude ontogenetic characters from our phylogenetic analysis. Further ontogenetic precision could be gained from aging individuals using bone histology, which has not been widely applied to ankylosaurians. As Liaoningosaurus and Chuanqilong are represented by juvenile specimens only, more material, especially adult specimens, will help to further elucidate their phylogenetic relationships.
Chuanqilong was moderate in size compared with other known ankylosaurians ( ). However, it still larger than adult Jurassic ankylosaurians, including Mymoorapelta and Gargoyleosaurus [18], [19]. The juvenile Chuanqilong is similar in size to most Cretaceous ankylosaurians, including adult Hungarosaurus [55] and Europelta [56], but is smaller than Cedarpelta (7.5–8.5 m: [20]) and Polacanthus (5–7 m: [53]). However, as the holotype of Chuangqilong is not fully-grown, based on the above-mentioned features, this suggests that the adults of this taxon may have been among the largest ankylosaurians. This suggests in turn that ankylosaurs has already evolved large size by the late Early Cretaceous. Bone histology should be used in future to gain a more accurate understanding of the ontogenetic age of this specimen.
Supporting Information
Text S1
Updated character scores for Chuanqilong , and additional scores for Liaoningosaurus .
(DOC)
Acknowledgments
We thank Haijun Li for his invitation to work on the material and for his hospitality in Chaoyang, Liaoning province. We thank Hailong Zang for providing the photographs of this specimen. We thank Xulong Lai and Richard Butler for their useful comments. Many thanks to editor Peter Dodson, and reviewers Victoria Arbour and Kenneth Carpenter for their helpful and constructive reviews of an earlier version of this article.
Funding Statement
This project was supported by the National Natural Science Foundation of China (41120124002; 41172026) and 973 program (2012CB821900). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
Data Availability
The authors confirm that all data underlying the findings are fully available without restriction. All relevant data are within the paper and its Supporting Information files.
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[] |
[] |
[
""
] | null |
[
"Bryanna West",
"tcu.academia.edu"
] |
2021-12-28T00:00:00
|
Big Bend National Park is known for its unique Late Cretaceous fauna, such as Alamosaurus sanjuanensis and Quetzalcoatlus northropi. Most major groups of dinosaurs are represented in the Late Cretaceous strata, which ranges from the Early Campanian
|
https://www.academia.edu/66199999/Campanian_Maastrichtian_Ankylosaurs_of_West_Texas
|
"A new, small ankylosaurid, Ahshislepelta minor, from the upper Campanian Kirtland Formation (Hunter Wash Member), San Juan Basin, New Mexico, consists of shoulder girdle and forelimb elements, vertebral fragments, and numerous osteoderms. Ahshislepelta minor differs from other ankylosaurids on the basis of a prominent dorsolateral overhang of the acromion and its osteoderm texture. It ranks as one of the most complete ankylosaur specimens known from New Mexico and adds to our understanding of ankylosaurid paleobiogeography, stratigraphy, and taxonomy."
Isolated bones and osteoderms of ankylosaurian dinosaurs recovered from Late Cretaceous sediments of northern Coahuila, northeastern Mexico, have been identified as remains of nodosaurids. Here, we summarize these discoveries and provide a review on Mexican Ankylosauria from a taxonomic perspective. We also present a new taxon, Acantholipan gonzalezi gen. et sp. nov. from the Pen Formation and provide a phylogenetic analysis integrating the new taxon. A. gonzalezi is the first named ankylosaur from Mexico that adds to the currently rare nodosaurid diversity from southern Laramidia.
Nodosaurid ankylosaur remains from the Upper Cretaceous of Mexico are summarized. The specimens are from the El Gallo Formation of Baja California, and the Pen and Aguja Formations of northwestern Coahuila, Mexico. These specimens show significant differences from other known nodosaurids, including ulna with very well developed olecranon and prominent humeral notch, the distal end of the femur not flaring to the extent seen in other nodosaurids, and a horn-like spine with vascular grooves on one side. The specimens are important because they are the southern-most occurrences in North America, and provide an important biogeographical link between nodosaurids of the United States and Canada on the one hand, and Argentina and Antarctica on the other.
Fossil evidences of the presence of ankylosaurian dinosaurs in Gondwana are scarce but consistent, being found in Antarctica, Oceania and South America. In spite that there are no nominated species in South America, the ankylosaur fossil record has increased in the last years. Indeterminate nodosaurid specimens, some isolated osteoderms and many trackways are known from the Upper Cretaceous of South America. The aim of the present contribution is to report new ankylosaurian remains from the Allen Formation (Campanian-Maastrichtian) at the Salitral Moreno locality, Northern Patagonia, Argentina. These osteoderms are small and conical, and includes thoracic, sacral and caudal scutes. The thoracic and sacral pieces are similar to those belonging to nodosaurids. The caudal osteoderm is a new element for the record of South American ankylosaurs. It resembles the caudal plates of Kunbarrasaurus and some ankylosaurids. The scutes show a mixture of characters so it is not possible to assign these pieces to a nodosaurid-like or ankylosaurid ankylosaur. These elements are consistent with the previously known ankylosaur fossil record of the Upper Cretaceous of Argentina, being a new sample of the diversity of the latest Cretaceous from South America.
|
|||||
622
|
dbpedia
|
2
| 2
|
https://dinotoyblog.com/zhejiangosaurus-vitae/
|
en
|
Zhejiangosaurus (Vitae) – Dinosaur Toy Blog
|
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2020-09-11T17:33:47+00:00
|
en
|
https://dinotoyblog.com/zhejiangosaurus-vitae/
|
While a lot of dinosaur names are quite a challenge for the laymen to be read and correctly spelled, the ones inspired by Chinese locations and names may even be a serious challenge to the dinosaur expert. Zhejiangosaurus comes as one of the easier names, but maybe one you do not really need to remember…..
The fossils related to that name were found in 2007 near Lishui, Zhejiang Province, in the subtropical parts of China. While there`s an offical holotype announced, no diagnostic features are specified, rendering the species a nomen dubium. As far as the description goes, the fossil is identified as a nodosaurid, the same family amongst Ankylosauria to which the more popular Sauropleta and Edmontonia belong. Zhejiangosaurus grew up to 4.5 metres and may have weighed around 1.5 metric tons.
Vitae had a short but glamorous life as a company and left the collectors with few figures, but these of comparable high quality. While some models were cast from resin, others are made from PVC and can even serve as toys. The Zhejiangosaurus counts amongst these.
The figure measures 17 cm in direct length, stands 4 cm high above the hips and is 4.5 cm wide at the hips, 5.5 cm including the spikes. The pose gives the figure a quite agile look, walking in a wide stride as if it has a certain target it wants to reach in time, like a mating partner or a contendor to fend off. The detailing is very fine, all spikes and scutes are reasonably sharp and pointed. There are five digits in the front feet, and four toes in the hind feet. The most prominent feature of this figure are the absurdly wide hips under a massive bony plate, it just looks so ankylosauriangood. Light chocolate brown is the main color, the belly is light tan, a modest pattern of white and reddish brown decorates the flanks.
You may recognize that the head is not original in my figure. I customized it before I thought of writing a review, because it was the only downer for me. The original face is deeply set into the skull and looks somewhat emaciated and off. I filled it, sculpted new eyes and added two pairs of scutes to the nose and brows. I am unaware and oblivious about the (possible) scientific accuracy of this feature, but I like the figure better this way.
If you want to add this nice little thyreophoran to your collection, hurry and grab one from Aliexpress. With Vitae out of business since a year or so, who knows how long supplies lasts?!
|
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622
|
dbpedia
|
0
| 11
|
https://www.desertcart.com.om/products/161513121-vitae-zhejiangosaurus-dinosaur-model-toy-collectable-art-figure
|
en
|
Buy Vitae ZHEJIANGOSAURUS Dinosaur Model Toy Collectable Art Figure Online at desertcart OMAN
|
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Shop Vitae ZHEJIANGOSAURUS Dinosaur Model Toy Collectable Art Figure online at best prices at desertcart - the best international shopping platform in OMAN. âFREE Delivery Across OMAN. âEASY Returns & Exchange.
|
en
|
https://cdn.desertcart.com/favicon.ico
|
https://www.desertcart.com.om/products/161513121-vitae-zhejiangosaurus-dinosaur-model-toy-collectable-art-figure
|
Disclaimer: The price shown above includes all applicable taxes and fees. The information provided above is for reference purposes only. Products may go out of stock and delivery estimates may change at any time. desertcart does not validate any claims made in the product descriptions above. For additional information, please contact the manufacturer or desertcart customer service. While desertcart makes reasonable efforts to only show products available in your country, some items may be cancelled if they are prohibited for import in United Arab Emirates. For more details, please visit our Support Page.
|
|||||
622
|
dbpedia
|
1
| 9
|
https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4580109/
|
en
|
Ankylosaurid dinosaur tail clubs evolved through stepwise acquisition of key features
|
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[] |
[] |
[
""
] | null |
[
"Victoria M Arbour",
"Philip J Currie"
] |
2015-10-10T00:00:00
|
Ankylosaurid ankylosaurs were quadrupedal, herbivorous dinosaurs with abundant dermal ossifications. They are best known for their distinctive tail club composed of stiff, interlocking vertebrae (the handle) and large, bulbous osteoderms (the knob), which ...
|
en
|
https://www.ncbi.nlm.nih.gov/coreutils/nwds/img/favicons/favicon.ico
|
PubMed Central (PMC)
|
https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4580109/
|
Results
The anterior caudal vertebrae of ankylosaurs (e.g. Arbour & Currie, 2013a: fig. 9; Arbour et al. 2009: fig. 9) are wider than long, with amphiplatyan centra, and transverse processes set at about the midheight of the centrum. The prezygapophyses are separate (unlike those of the dorsal vertebrae, which are joined at the midline), and are finger-like projections from the neural arch; the postzygapophyses do not extend far past the posterior border of the neural spine. The neural spine in ankylosaurids is usually mediolaterally thin (e.g. Euoplocephalus, Arbour & Currie, 2013a; Talarurus, Maleev, 1956), and in nodosaurids the neural spine can be more robust and with a substantial distal mediolateral expansion (e.g. Polacanthus, Blows, 1987: figs 2–3). The haemal arch often fuses to the posterior ventral edge of the centrum, and the haemal spine is about as long as the neural spine.
The distal caudal vertebrae in basal ankylosaurs (e.g. Mymoorapelta maysi Kirkland & Carpenter, 1994; Fig. ), basal ankylosaurids (e.g. Gastonia), and nodosaurids (e.g. Nodosaurus textilis Marsh, 1889; Sauropelta edwardsorum Ostrom, 1970) tend to be longer than wide, and dorsoventrally compressed relative to anterior caudals. The neural spine is reduced relative to the anterior caudals, and the haemal spine takes on a rounded, hatchet-shaped appearance (e.g. Hungarosaurus tormai Ősi, 2005: fig. 10C). The prezygapophyses do not extend past the anterior edge of the centrum by more than about 25% of the centrum length (e.g. Nodosaurus, Lull, 1921; Mymoorapelta MWC 5819, Fig. ). The distal tail of these ankylosaurs would have been flexible.
The distal caudal vertebrae of ankylosaurines are unique among dinosaurs. In basal nodosaurids and basal ankylosaurids, the morphological transition from anterior to posterior caudal vertebrae is smooth, but in ankylosaurines, this transition is abrupt and occurs at about the midpoint of the caudal series (e.g. Arbour et al. 2009: Fig. , Parks, 1924: Pl. 1). In contrast, the distal caudal vertebrae of ankylosaurines interlock tightly, forming a series of interlocking Vs in dorsal view (Fig. ). The prezygapophysis of each caudal overlaps the adjacent anterior vertebra by at least 50% of its length, unlike in basal ankylosaurids, nodosaurids, and basal ankylosaurs, where the overlap is only about 25% of the centrum length (e.g. Mymoorapelta MWC 5819, Fig. ). The prezygapophyses are dorsoventrally deep, mediolaterally flattened, and with vertically oriented articular surfaces; the modified neural spine and postzygapophyses of the preceding vertebra completely fill the space between the prezygapophyses. Transverse processes are absent on most vertebrae in this region, although small bumps may be present on the first few handle vertebrae (Arbour et al. 2009). The haemal arches are similarly modified into a tightly interlocking series. The haemal spine is dorsoventrally short but anteroposteriorly long, and has a boat-like shape (e.g. Maleev, 1956: fig. 35; Maryańska, 1977: fig. 10). Anteriorly it is bifurcated, and posteriorly it tapers to a point. Flexibility in the distal portion of the tail in ankylosaurines was highly reduced, and where fusion of the vertebral centra occurred, there would have been almost no flexibility whatsoever.
In ankylosaurines, knob osteoderms completely envelop and obscure the distalmost vertebrae. Two laterally positioned osteoderms (the major osteoderms, sensu Coombs, 1995) form the bulk of the knob and are usually keeled and dorsoventrally flattened rather than hemispherical. Two or more smaller osteoderms form the terminus of the knob, and the boundaries between these minor knob osteoderms can be indistinct (Arbour & Currie, 2013a). Only a few specimens preserve osteoderms along the more proximal portions of the tail, and the best example is MPC 100/1305, a large ankylosaurid tentatively referred to Pinacosaurus (Arbour & Currie, 2013b). In this specimen, the lateral osteoderms are triangular and sharply pointed in the anterior and middle portions of the tail, and become smaller and less sharply pointed posteriorly (Carpenter et al. 2011: fig. 15; Arbour & Currie 2013b: fig. 1). The penultimate pair of lateral osteoderms anterior to the knob are similar to the major knob osteoderms, with rounded lateral edges, but they are not as dorsoventrally deep and do not envelop the handle vertebrae to the same degree as the knob osteoderms.
We find direct evidence for two ankylosaurs with a tail club handle but not a tail club knob: Gobisaurus domoculus and Liaoningosaurus paradoxus. HGM 41HIII-0002 (the holotype of Zhongyuansaurus luoyangensis Xu et al. 2007; but referred to Gobisaurus by Arbour & Currie, in press ) clearly preserves the handle of a tail club (contra Xu et al. 2007 and Carpenter et al. 2008), even though knob osteoderms are not present (Fig. , ). The vertebrae are indistinguishable from those of more derived ankylosaurine ankylosaurs, with elongated prezygapophyses and neural spines that interlock tightly together. The tail club of HGM 41HIII-0002 appears to preserve the distalmost caudal vertebra; the last three vertebrae in the handle abruptly shorten, and the terminal vertebra is rounded at the distal end, similar to what was observed in CT scans (Fig. , ) of an Albertan tail club (UALVP 16247, Arbour, 2009). The tail club of HGM 41HIII-0002 is unusual compared with other ankylosaurid tail clubs because it preserves no evidence for the large terminal knob osteoderms. No other ankylosaurid specimen preserves the distal end of the handle without at least some of the knob preserved, because the knob osteoderms envelop and are tightly appressed to the vertebrae and associated ossified tendons. This suggests that either a large terminal knob was not present in HGM 41HIII-0002, or that the knob osteoderms were smaller or more loosely associated with the handle vertebrae.
The second ankylosaur that preserves tail club handle vertebrae without a tail club knob is Liaoningosaurus. IVPP V12560, the holotype of Liaoningosaurus paradoxus, is one of the smallest known ankylosaur skeletons, at only about 33 cm in length (Fig. ). Unfused neural arches, small size, and the absence of osteoderms posterior to the cervical/pectoral region (as in juvenile Pinacosaurus grangeri, Burns et al. 2011) suggest that IVPP V12560 is a juvenile individual. Liaoningosaurus paradoxus does not possess an obvious tail club, but close observation of the distal caudal vertebrae shows a close similarity to the handle vertebrae of ankylosaurines. The neural arches of the distal tail vertebrae interlock, and the prezygapophyses overlap the adjacent vertebra by at least 50% of the centrum length, as in ankylosaurines (Fig. ). Therefore, Liaoningosaurus paradoxus appears to have possessed a tail club handle but does not appear to have had a tail club knob. However, osteoderms are only preserved in the cervical/pectoral region (Fig. ) and it is possible that the full complement of osteoderms had not yet developed in the holotype specimen IVPP V12560. Intriguingly, Chuanqilong does not appear to have modified handle-like vertebrae in its distal tail (Han et al. 2014: fig. 3) despite its similar geologic age and provenance and possible close relationship to Liaoningosaurus. Han et al. (2014) recovered Chuanqilong as the sister taxon to Liaoningosaurus, although Arbour & Currie (in press) found Chuanqilong as the sister taxon to Cedarpelta).
Ancestral state reconstruction provides additional information on the origin of elongated prezygapophyses that overlap at least 50% of the preceding vertebral centrum length, and on the origin of enlarged knob osteoderms (Fig. ). Elongated prezygapophyses were present in the ancestor of all ankylosaurines more derived than Crichtonpelta (proportional likelihood = 1.000), and were most likely present in the ancestor of Ankylosaurinae+Shamosaurinae (proportional likelihood = 0.963). Whether the ancestor of the clade containing all ankylosaurids more derived than Ahshislepelta and Gastonia had elongated prezygapophyses is equivocal (proportional likelihood 0.501) because the base of this clade includes a polytomy that has taxa with elongated prezygapophyses (Liaoningosaurus) and taxa that do not (Aletopelta, Chuanqilong). Knob osteoderms have a more restricted phylogenetic distribution: a tail club knob was most likely present in the ancestor of all ankylosaurines more derived than Crichtonpelta (proportional likelihood = 0.977) but not in the ancestor of Ankylosaurinae+Shamosaurinae (proportional likelihood = 0.023).
Discussion
Ankylosaurid tail clubs are complex structures involving contributions from both the vertebral series and the dermal skeleton. Our results suggest that the tail club evolved in a stepwise fashion, in which modifications to the distal caudal vertebrae preceded modifications to the terminal osteoderms (Fig. ).
Bonebed material at the DMNH of Gastonia burgei, the oldest ankylosaurid in this study (although some other analyses recover this taxon as a basal nodosaurid, e.g. Thompson et al. 2012), includes a large sample of caudal vertebrae of many sizes and positions within the vertebral series, and none has the distinctive morphology of handle vertebrae.
Isolated osteoderms that could be identified as disarticulated knob osteoderms are unknown in any formations prior to the Campanian. A tail club was described for Tianchisaurus nedegoapeferima from the Middle Jurassic of China (Dong, 1993), which would make this the earliest occurrence of a tail club in the fossil record. However, the ‘tail club’ of IVPP V10614 does not appear to represent a tail club knob. The putative knob appears subdivided by deep grooves into three sections, with two larger sections flanking a small triangular area. In most ankylosaurid knobs, the major osteoderms are clearly separated at the midline in dorsal and ventral view, and the terminal end of the knob is made up of more than one osteoderm (e.g. Arbour & Currie, 2013a: Fig. 14). It is unclear what the putative knob of Tianchisaurus nedegoapeferima represents, but it is unlikely that it is a true tail club knob, and so this should not be considered the first occurrence of an ankylosaurine-like tail club in the fossil record. Another putative tail club-like structure was reported for Polacanthus foxii Owen vide Anonymous, 1865, from the Barremian Wessex Formation of England (Blows & Honeysett, 2014). Blows (1987) described a ‘caudal end mass’ consisting of osteoderms, caudal vertebrae, and ossified tendons in NHMUK R175, and considered that this represented the terminus of the tail and fusion of these elements. Later, Blows (2001) suggested that the presence of ossified tendons in the caudal region of Polacanthus (which are also present in ankylosaurines with a tail club) may have been an adaptation for lateral tail swinging even in the absence of a fully developed tail club. Pereda-Suberbiola (1994) and Carpenter & Kirkland (1998) considered the ‘caudal end mass’ of NHMUK R175 to represent a more anterior portion of the tail, and suggested that it does not represent an incipient tail club. We agree that this structure does not represent the distal end of the tail or an incipient tail club.
The oldest ankylosaur to possess either of the two modifications present in derived ankylosaurid tail clubs (distal caudal vertebrae modified to form a handle, or terminal osteoderms enlarged and enveloping the tail terminus) is the holotype of Liaoningosaurus paradoxus (122 Ma, Aptian; Xu & Norell, 2006; Fig. ). In Liaoningosaurus, the prezygapophyses of the distal caudal vertebrae overlap the preceding vertebra by at least 50% of its length, a feature found only in more derived ankylosaurids with complete tail clubs (Fig. ); this feature is not present in more basal ankylosaurids such as Gastonia, nodosaurids such as Sauropelta, or basal ankylosaurs such as Mymoorapelta. Liaoningosaurus lacks knob osteoderms but the holotype (and only published specimen to date) is a very small juvenile and likely had not developed its full complement of osteoderms before it died. This makes it difficult to determine whether Liaoningosaurus had a tail club knob in addition to the modified distal caudal vertebrae. However, one specimen of Gobisaurus (HGM 41HIII-0002) preserves a tail club handle without a knob. This specimen includes the terminal caudal vertebrae, so the absence of the knob is not because the end of the tail is missing. Although the skull for HGM 41HIII has some cranial sutures visible (Arbour & Currie, in press), which suggests that the specimen is not fully mature, it is still a relatively large individual, and several post-cervical osteoderms were associated with it (Xu et al. 2007). Ontogeny does not seem to be the best explanation for the absence of knob osteoderms in this specimen. It is possible that knob osteoderms were present in life and disarticulated from the handle after death. However, in isolated tail club knobs from more derived ankylosaurids, there are often some fragments of the distal caudal vertebrae or ossified tendons associated with the knob or knob osteoderms; this is most likely because of the close association between these elements in the living animal (e.g. UALVP 16247, CMN 2251). Thompson et al. (2012) considered HGM 41HIII-0002 (as Zhongyuansaurus) to be the first known ankylosaurid in which the tail club was definitively absent, although this was in reference to a ‘fully developed’ tail club consisting of a handle and knob. However, HGM 41HIII-0002 clearly preserves a tail club handle. After Liaoningosaurus paradoxus, Gobisaurus domoculus is the next youngest ankylosaurid known to have possessed a tail club, with an age of no more than 92 Ma (Turonian; Kobayashi & Lü, 2003, Fig. ). Given the close anatomical similarity between the overlapping elements of Gobisaurus and Shamosaurus, it seems likely that Shamosaurus also had a tail club handle; ancestral state reconstruction also suggests that the ancestor of Gobisaurus and Ankylosaurinae had a tail club handle. Ancestral state reconstruction suggests that Tsagantegia and ‘Zhejiangosaurus’, for which caudal material is unknown, most likely each had a tail club handle. Cedarpelta, from the Mussentuchit Member of the Cedar Mountain Formation (∼104–98 Ma, Chure et al. 2010; Cifelli et al. 1997), is the oldest North American ankylosaurid besides Gastonia and has been considered closely related to Gobisaurus and Shamosaurus (Carpenter et al. 2008). Unfortunately, no distal caudal vertebrae are known for Cedarpelta, and ancestral state reconstruction was ambiguous about the presence or absence of a tail club handle at this level in the phylogeny (Fig. ). At present, no pre-Campanian North American ankylosaurids appear to have had a tail club handle.
The oldest and most basal ankylosaur known to possess terminal osteoderms that envelop the end of the tail is Pinacosaurus, from the Campanian (Dashzeveg et al. 2005) of Mongolia and China (Fig. ); Talarurus (PIN 557) is geologically older but phylogenetically more derived, and the handle is incomplete. All ankylosaurid ankylosaurs that are more derived than Pinacosaurus either are known to have had a tail club (e.g. Ankylosaurus, Euoplocephalus) or occur in formations in which disarticulated tail clubs are known but cannot be attributed to a specific taxon (e.g. Tarchia, Arbour et al. 2014a; Ziapelta, Arbour et al. 2014b). Several ankylosaurid specimens from Mongolia (PIN 614 and MPC 100/1305, both tentatively assigned to Pinacosaurus grangeri, Arbour & Currie, 2013b, and ZPAL MgD I/113, an indeterminate ankylosaurid from the Nemegt Formation, Arbour et al. 2013) demonstrate that osteoderms were present along the length of the handle, not just at the terminus. Interestingly, in MPC 100/1305 the penultimate lateral osteoderms are rounded and similar in shape to the major knob osteoderms, although they are not as dorsoventrally deep.
Crichtonpelta is the earliest and most basal ankylosaurine (Fig. ), but no caudal material has been described for this taxon. An undescribed mounted skeleton on display at the Sihetun Fossil Museum (Liaoning, China) is presented with a tail club, but it is unclear whether this has been sculpted or represents real fossil material, and the tail vertebrae have not yet been described or figured. Ancestral state reconstruction suggests that Crichtonpelta probably had a tail club handle, but was ambiguous about whether Crichtonpelta was likely to have a tail club knob (Fig. ). Ankylosaurines more derived than Crichtonpelta were very likely to have a knob, but the ancestor of Shamosaurinae and Ankylosaurinae was unlikely to have had a knob.
Understanding trends within the evolution of the tail club among ankylosaurines is complicated by the dearth of tail clubs that can be referred to different species; for example, it is difficult to associate isolated tail club knobs from the Baruungoyot and Nemegt formations of Mongolia with any of the known ankylosaurids from those formations (Saichania, Tarchia, and Zaraapelta) because no specimens preserving a skull and tail club have been described in detail yet (Arbour et al. 2014a). Nevertheless, two patterns merit further investigation as more specimens are collected. First, the maximum size of tail club knobs seems to increase from the earliest known knob to later knobs in the late Campanian and Maastrichtian. The largest tail club knob from the Djadokhta Formation (on MPC 100/1305, ?Pinacosaurus), the stratigraphically oldest formation with tail club knobs, is 146 mm wide. The largest knob from the younger Nemegt, Dinosaur Park, and Horseshoe Canyon formations are 620 mm wide (ZPAL MgD I/43), 572 mm wide (ROM 788), and 593 mm wide (AMNH 5245), respectively, and the only known tail club from the youngest unit, the Scollard Formation, is ∼450 mm wide (AMNH 5214, a subadult Ankylosaurus). Testing this apparent trend is complicated by the lack of precise age estimates for the Djadokhta, Baruungoyot, and Nemegt formations, and the absence of information about ankylosaur knob osteoderm ontogeny. Secondly, although the morphology of the handle vertebrae is consistent across most species of ankylosaurines, it diverges significantly in two taxa. In most ankylosaurines, the edges of the neural spines diverge at an angle of about 22° in dorsal view, forming the distinctive interlocking V morphology. In ZPAL MgD I/113, an ankylosaur of unknown affinity from the Nemegt Formation, this angle is approximately 35°, and in Ankylosaurus (AMNH 5214) it is approximately 60° (Arbour et al. 2009). In Ankylosaurus, this results in a U-shaped rather than a V-shaped neural spine in the handle vertebrae. Why these two taxa diverged from the basal condition found in other ankylosaurines is not clear, but it is worth noting that these two specimens are among the largest of all known ankylosaurines, suggesting that overall body size increases may have necessitated a change in tail club morphology.
The absence of enlarged terminal osteoderms in taxa without handle vertebrae, and the absence of isolated knob osteoderms in formations without taxa that had a tail club handle, suggests that the hypothesis that the tail club knob evolved before the handle (Fig. ) can be rejected. Biomechanically, a large knob of dermal bone at the end of a flexible tail (analogous to a flail, rather than a club) could result in damage to the ankylosaur if wielded as a weapon; the rotational inertia of a large mass at the end of the tail could lead to tearing of the soft tissues between the vertebrae, and twisting stresses could break the vertebrae. The absence of knob osteoderms in Gobisaurus suggests that the handle-first hypothesis (Fig. ) may best explain the evolution of the ankylosaurid tail club. This is also supported by the results from ancestral state reconstruction (Fig. ), which reconstruct a later and more derived first appearance of the tail club knob relative to the first appearance of elongated prezygapophyses. However, we cannot reject the hypothesis that the handle and knob evolved in tandem (Fig. ), as the absence of knob osteoderms in the known specimens of Gobisaurus or Liaoningosaurus might be attributed to ontogenetic and/or taphonomic changes. Basal thyreophorans such as Scelidosaurus harrisonii Owen, 1861 had spiky lateral tail osteoderms that would certainly have been effective weapons if the tails were swung from side to side, even if they were not being used to deliver forceful impacts. Although early ankylosaurids and shamosaurines may not have had the enlarged knob osteoderms found in later taxa, the stiffened end of the tail may still have been an effective bat-like weapon.
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https://dokumen.pub/the-princeton-field-guide-to-dinosaurs-course-booknbsped-9781400836154.html
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en
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The Princeton Field Guide to Dinosaurs [Course Book ed.] 9781400836154
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This lavishly illustrated volume is the first authoritative dinosaur book in the style of a field guide. World-renowned...
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dokumen.pub
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https://dokumen.pub/the-princeton-field-guide-to-dinosaurs-course-booknbsped-9781400836154.html
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Table of contents :
CONTENTS
Preface
Introduction
Group and Species Accounts
Dinosaurs
Theropods
Sauropodomorphs
Ornithischians
Additional Reading
Index
Citation preview
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https://handwiki.org/wiki/Biology:Nodosauridae
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Biology:Nodosauridae
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Nodosauridae is a family of ankylosaurian dinosaurs, from the Late Jurassic to the Late Cretaceous period in what is now North America, South America, Europe, and Asia.[1]
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Short description: Extinct family of dinosaurs
Nodosaurids
Gargoyleosaurus skeleton cast Scientific classification Domain: Eukaryota Kingdom: Animalia Phylum: Chordata Clade: Dinosauria Clade: †Ornithischia Clade: †Thyreophora Suborder: †Ankylosauria Clade: †Euankylosauria Family: †Nodosauridae
Marsh, 1890 Subgroups
Acanthopholis
Anoplosaurus
Dongyangopelta
Gastonia
Gargoyleosaurus
Glyptodontopelta
Horshamosaurus
Hylaeosaurus?
Invictarx
Mymoorapelta?
Priconodon?
Propanoplosaurus
Rhadinosaurus
Sauroplites
Polacanthinae
Nodosaurinae
Synonyms
Acanthopholididae Nopcsa, 1902
Acanthopholidae Nopcsa, 1917
?Hylaeosauridae Nopcsa, 1902
Palaeoscincidae Nopcsa, 1918
Panoplosauridae Nopcsa, 1929
Struthiosauridae Kuhn, 1966
Nodosauridae is a family of ankylosaurian dinosaurs, from the Late Jurassic to the Late Cretaceous period in what is now North America, South America, Europe, and Asia.[1]
Description
Nodosaurids, like their close relatives the ankylosaurids, were heavily armored dinosaurs adorned with rows of bony armor nodules and spines (osteoderms), which were covered in keratin sheaths. All nodosaurids, like other ankylosaurians, were medium-sized to large, heavily built, quadrupedal, herbivorous dinosaurs, possessing small, leaf-shaped teeth. Unlike ankylosaurids, nodosaurids lacked mace-like tail clubs, instead having flexible tail tips. Many nodosaurids had spikes projecting outward from their shoulders. One particularly well-preserved nodosaurid "mummy", the holotype of Borealopelta markmitchelli, preserved a nearly complete set of armor in life position, as well as the keratin covering and mineralized remains of the underlying skin, which indicate reddish dorsal pigments in a countershading pattern.[2][3]
Classification
The family Nodosauridae was erected by Othniel Charles Marsh in 1890, and anchored on the genus Nodosaurus.[4][5]
The clade Nodosauridae was first informally defined by Paul Sereno in 1998 as "all ankylosaurs closer to Panoplosaurus than to Ankylosaurus," a definition followed by Vickaryous, Teresa Maryańska, and Weishampel in 2004. Vickaryous et al. considered two genera of nodosaurids to be of uncertain placement (incertae sedis): Struthiosaurus and Animantarx, and considered the most primitive member of the Nodosauridae to be Cedarpelta.[6] Following the publication of the PhyloCode, Nodosauridae needed to be formally defined following certain parameters, including that the type genus Nodosaurus was required as an internal specifier. In formally naming Nodosauridae, Madzia and colleagues followed the previously established use for the clade, defining it as the largest clade including Nodosaurus textilis but not Ankylosaurus magniventris. As all phylogenies referenced included both Panoplosaurus and Nodosaurus within the same group relative to Ankylosaurus, the addition of another internal specifier was deemed unnecessary. The 2018 phylogenetic analysis of Rivera-Sylva and colleagues was used as the primary reference for Panoplosaurini by Madzia et al., in addition to the supplemental analyses of Thompson et al. (2012), Arbour and Currie (2016), Arbour et al. (2016), and Brown et al. (2017).[7][8][9][10][11][12]
Nodosauridae
Sauroplites
Mymoorapelta
Dongyangopelta
Gastonia
Gargoyleosaurus
Polacanthinae
Hoplitosaurus
Polacanthus
Nodosaurinae
Peloroplites
Taohelong
Sauropelta
Acantholipan
Nodosaurus
Niobrarasaurus
Ahshislepelta
Tatankacephalus
Silvisaurus
CPC 273
Panoplosaurini
Animantarx
Panoplosaurus
Argentinian ankylosaur (Patagopelta)
Texasetes
Denversaurus
Edmontonia longiceps
Edmontonia rugosidens
Struthiosaurini
Hungarosaurus
Europelta
Pawpawsaurus
Borealopelta markmitchelli
Stegopelta
Struthiosaurus languedocensis
Struthiosaurus transylvanicus
Struthiosaurus austriacus
The highly isolated Antarctopelta, from the late Cretaceous of Antarctica, was previously thought to be the most basal nodosaurid, but a 2021 study found it to belong to the Parankylosauria, a separate basal lineage of ankylosaurs restricted to the Southern Hemisphere.[13] However, the 2022 description of Patagopelta, a nodosaurine from South America, suggests that true nodosaurids also inhabited Gondwana, having colonized South America during a biotic interchange from North America during the Campanian.[14]
Biogeography
The near simultaneous appearance of nodosaurids in both North America and Europe is worthy of consideration. Europelta is the oldest nodosaurid from Europe, it is derived from the lower Albian Escucha Formation. The oldest western North American nodosaurid is Sauropelta, from the lower Albian Little Sheep Mudstone Member of the Cloverly Formation, at an age of 108.5±0.2 million years. Eastern North American fossils seem older. Teeth of Priconodon crassus from the Arundel Clay of the Potomac Group of Maryland, which dates near the Aptian–Albian boundary. The Propanoplosaurus hatchling from the base of the underlying Patuxent Formation, dating to the upper Aptian, is the oldest known nodosaurid.[4]
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</timeline>
See also
Timeline of ankylosaur research
References
Further reading
Carpenter, K. (2001). "Phylogenetic analysis of the Ankylosauria." In Carpenter, K., (ed.) 2001: The Armored Dinosaurs. Indiana University Press, Bloomington & Indianapolis, 2001, pp. xv-526
Osi, Attila (2005). Hungarosaurus tormai, a new ankylosaur (Dinosauria) from the Upper Cretaceous of Hungary. Journal of Vertebrate Paleontology 25(2):370-383, June 2003.
Alberta oilsands discovery of 2011
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https://en.wikipedia.org/wiki/Zhejiangosaurus
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Zhejiangosaurus
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https://en.wikipedia.org/wiki/Zhejiangosaurus
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Zhejiangosaurus lishuiensis on display at the Zhejiang Museum of Natural History Scientific classification Domain: Eukaryota Kingdom: Animalia Phylum: Chordata Clade: Dinosauria Clade: †Ornithischia Clade: †Thyreophora Clade: †Ankylosauria Clade: †Euankylosauria Genus: †Zhejiangosaurus
Lü et al., 2007 Species:
†Z. lishuiensis
Binomial name †Zhejiangosaurus lishuiensis
Lü et al., 2007
Zhejiangosaurus (meaning "Zhejiang lizard") is an extinct genus of ankylosaurian dinosaur from the Upper Cretaceous (Cenomanian stage) of Zhejiang, eastern China. It was first named by a group of Chinese authors Lü Junchang, Jin Xingsheng, Sheng Yiming and Li Yihong in 2007 and the type species is Zhejiangosaurus lishuiensis ("from Lishui", where the fossil was found).[1] It has no diagnostic features, and thus is a nomen dubium.[2]
Description
[edit]
Zhejiangosaurus could grow up to 4.5 m (17 ft) in length and was 1.4 metric tons in weigh.[3]
Material
[edit]
Material for Zhejiangosaurus consists of the holotype, ZNHM M8718, a partial skeleton which has preserved a sacrum with eight vertebrae, a complete right ilium and partial left ilium, a complete right pubis, the proximal end of the right ischium, two complete hindlimbs, fourteen caudal vertebrae, and some unidentified bones. These remains come from Liancheng, in the Chinese administrative unit of Lishui on the province of Zhejiang and they were collected from the Cenomanian-age Chaochuan Formation.[1]
Systematics
[edit]
On the species description, Lü et al. (2007) found Zhejiangosaurus to belong to the ankylosaurian family Nodosauridae.[1][4]
Zhejiangosaurus in a cladogram after Pond et al. (2023):[5]
See also
[edit]
Dinosaurs portal
China portal
Timeline of ankylosaur research
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Scolosaurus is an extinct genus of ankylosaurid dinosaurs within the subfamily Ankylosaurinae. It is known from the lower levels of the Dinosaur Park Formation ...
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Scolosaurus was named by Franz Nopcsa von Felső-Szilvás in 1928, based on holotype NHMUK PV R.5161, a nearly complete specimen that preserves the entire skeleton except for the distal end of the tail, the right forelimb, the right hindlimb, and the skull. The rare preservation of osteoderms and skin impression are also present. The fossil skeleton was discovered by William Edmund Cutler, an independent fossil collector in 1914 at Quarry 80 of the Deadlodge Canyon locality.[2][4] It was collected from the bottom of the Dinosaur Park Formation in fine-grained sandstone and fine-grained claystone sediments that were deposited during the Campanian stage of the Late Cretaceous period, approximately 76.5 million years ago.[5] The holotype specimen is housed in the collection of the Natural History Museum in London, England.
In 2013, Arbour and Currie reassigned specimen MOR 433, upon which the genus Oohkotokia was based, to Scolosaurus. This specimen consists of a partial skull, both humeri, a caudal vertebra and several osteoderms and was recovered in the Upper Member of the Two Medicine Formation, in Montana, which has been dated at approximately 74 million years.[6] The remains were collected in 1986-1987 in grey siltstone that was deposited during the Campanian stage of the Cretaceous period.[5] The specimen is housed in the collection of the Museum of the Rockies in Bozeman, Montana.
The generic name Scolosaurus means "pointed stake lizard" and is derived from the Greek words skolos (σκῶλος) meaning "pointed stake", and saûros (σαύρα) meaning "lizard".[7] The specific name, cutleri, honours its discoverer and the collector of the holotype, W. E. Cutler,[4] who was seriously injured when the specimen fell on him as he was excavating it.[8]
In 1928, Nopcsa assigned the specimen to the family Ankylosauridae, and drew morphological comparisons with the fossil material known from Dyoplosaurus. In 1971, Walter Coombs concluded that there was only one species of ankylosaurid during the Campanian age of the Late Cretaceous of North America. He synonymized the species Anodontosaurus lambei, Dyoplosaurus acutosquameus, and Scolosaurus cutleri with Euoplocephalus tutus but did not provide any justification for these synonymies.[9] The synonymization of Scolosaurus cutleri and Euoplocephalus tutus was generally accepted and thus NHMUK R.5161 was assigned to E. tutus. However, a redescription of Scolosaurus published in 2013 in the Canadian Journal of Earth Sciences by Paul Penkalski and William T. Blows suggested that the genus is a valid taxon. They concluded that Scolosaurus can be distinguished from Euoplocephalus by the form of their cervical armour, the details of other armour and the structure of the forelimb. They also concluded that Scolosaurus and Dyoplosaurus are distinct, due to differences noted in the pelvis and armour.[2] Due to its completeness, the holotype of Scolosaurus has formed the basis for most Euoplocephalus reconstructions since 1971; therefore, most images of Euoplocephalus actually depict Scolosaurus instead.
A 2013 study found that the ankylosaurine Oohkotokia was indistinguishable from Scolosaurus, and was therefore considered a junior synonym.[10] However, this synonymization is contentious as Oohkotokia was subsequently recognized as valid.[11] Thus, much of the material illustrated as belonging to Scolosaurus may actually pertain to Oohkotokia.
The following cladogram is based on a 2015 phylogenetic analysis of the Ankylosaurinae conducted by Victoria Arbour and Phillip J. Currie. The cladogram follows the biogeographical family tree provided by that study, which is a fusion of the study's 50% majority rule tree as well as the maximum agreement subtree. The study's 50% majority rule tree was a cladogram formed by a collection of clades, although it only included clades that appear in more than 50% of the family trees found during the analysis. The maximum agreement subtree is the cladogram that results from an algorithm which attempts to maximize the amount of taxa included in the result while also retaining the fundamental shape of all other trees in the sample. Some controversial taxa thus had to be omitted by the subtree in order for the resulting cladogram to fulfill the second requirement. The biogeographical tree (i.e. the following cladogram) is basically the 50% majority rule tree, except with some of the polytomies resolved according to the results of the maximum agreement subtree:[12]
Ankylosaurinae
Crichtonpelta
Tsagantegia
Zhejiangosaurus
Pinacosaurus
Saichania
Tarchia
Zaraapelta
Ankylosaurini
Dyoplosaurus
Talarurus
Nodocephalosaurus
Ankylosaurus
Anodontosaurus
Euoplocephalus
Scolosaurus
Ziapelta
The following cladogram is based on a 2017 phylogenetic analysis of the Ankylosaurinae conducted by Victoria Arbour and David Evans. The cladogram depicts the majority rule (average result) of 10 most parsimonious trees, which each are considered to have the fewest evolutionary steps, thus being the most accurate under the principle of Occam's razor:[13]
Ankylosaurinae
Zhejiangosaurus luoyangensis
Pinacosaurus grangeri
Pinacosaurus mephistocephalus
Tsagantegia longicranialis
Talarurus plicatospineus
Nodocephalosaurus kirtlandensis
Saichania chulsanensis
Zaraapelta nomadis
Tarchia kielanae
Ankylosaurini
Ziapelta sanjuanensis
Euoplocephalus tutus
Zuul crurivastator
Scolosaurus cutleri
Dyoplosaurus acutosquameus
Anodontosaurus lambei
Ankylosaurus magniventris
Referred material
In 1874, G. M. Dawson excavated specimen USNM 7943 at the Milk River locality of the Frenchman Formation in Alberta. It was collected from terrestrial sediments that are considered to be from the Maastrichtian stage of the Late Cretaceous, approximately 70.6 to 66 million years old. The specimen consisted of a partial first cervical ring, which is part of the dinosaur's neck. In 2013, this material was assigned to Scolosaurus by Arbour and Currie who conducted a detailed phylogenetic analysis of the ankylosauridae.[14] It is currently housed at the Smithsonian Institution in Washington, DC.
In 1928, George F. Sternberg, collected specimen USNM 11892, from the Montanazhdarcho holotype locality, high up in the Two Medicine Formation in Glacier County, Montana.[15] The material, a partial skull, was recovered from channel sandstone sediments that were deposited during the Campanian stage, approximately 74 million years ago. This is also housed at the Smithsonian Institution.
Other referred specimens include FPDM V-31, NSM PV 20381 and TMP 2001.42.9. FPDM V-31 and TMP 2001.42.9 are both skulls, in various states of preservation. NSM PV 20381 includes a skull, dorsal vertebrae, caudal vertebrae, ribs, both scapulae, both ilia, partial ischia, and both femora, both tibiae and fibulae.
Distinguishing anatomical features
A differential diagnosis is a statement of the anatomical features of an organism (or group) that collectively distinguish it from all other organisms. Some, but not all, of the features in a diagnosis are also autapomorphies. An autapomorphy is a distinctive anatomical feature that is unique to a given organism.
According to Arbour and Currie (2013), Scolosaurus (including the Two Medicine material) can be distinguished from other ankylosaurines based on the following characteristics:
the squamosal horns are proportionately longer, are backswept, and have distinct apices (unlike Anodontosaurus lambei and Euoplocephalus tutus)
the presence of a small circular caputegula at the bases of the squamosal and quadratojugal bones (unlike Euoplocephalus tutus)
the postacetabular process of the ilium is proportionately longer (compared to Anodontosaurus lambei, Dyoplosaurus acutosquameus and Euoplocephalus tutus)
the presence of proportionately large circular medial osteoderms with low central prominences, and compressed, half-moon shaped lateral/distal osteoderms on the cervical half rings (unlike Anodontosaurus lambei and Euoplocephalus tutus)
the sacral ribs are laterally-directed (unlike Dyoplosaurus acutosquameus)
the osteoderms are conical, with centrally positioned apices on the lateral sides of the anterior portion of the tail (unlike Dyoplosaurus acutosquameus)
the tail club knob appears circular in dorsal view, unlike that of Anodontosaurus, which appears wider than it is long or that of Dyoplosaurus, which appears longer than it is wide
the presence of anteriorly-directed nares, and the absence of a continuous keel between the squamosal horn and the supraorbital bones (unlike Ankylosaurus magniventris)
Habitat
Argon-argon radiometric dating indicates that the Two Medicine Formation was deposited between 83.5 and 70.6 million years ago, during the Campanian stage of the Late Cretaceous period, in what is now northwestern Montana.[16] If Oohkotokia is the same as Scolosaurus it would mean that Scolosaurus existed for around 3 million years. The Two Medicine Formation correlates to the Belly River Group in southwest Alberta, and the Pakowki Formation eastward. The Two Medicine Formation was deposited by rivers and deltas between the western shoreline of the Western Interior Seaway and the eastward advancing margin of the Cordilleran Overthrust Belt. Since the mid-Cretaceous, North America had been divided in half by this seaway, with much of Montana and Alberta below the surface of the water. However, the uplift of the Rocky Mountains forced the seaway to retreat eastwards and southwards. Rivers flowed down from the mountains and drained into the seaway, carrying sediment that formed the Two Medicine Formation and the Judith River Group. About 73 million years ago, the seaway began to advance westwards and northwards again, and the entire region was covered by the Bearpaw Sea, now preserved throughout the Western US and Canada by the massive Bearpaw Shale, which overlies the Two Medicine.[17][18] Below this formation are the nearshore deposits of the Virgelle Sandstone. Lithologies, invertebrate faunas, and plant and pollen data support that the Two Medicine Formation was deposited in a seasonal, semi-arid climate with possible rainshadows from the Cordilleran highlands. This region experienced a long dry season and warm temperatures. The extensive red beds and caliche horizons of the upper Two Medicine are evidence of at least seasonally arid conditions.
Paleofauna
Scolosaurus shared its paleoenvironment with other dinosaurs, such as the duck-billed hadrosaurs Hypacrosaurus, Acristavus, Gryposaurus, Brachylophosaurus, Glishades, Prosaurolophus and Maiasaura, and the ankylosaur Edmontonia.[19] Volcanic eruptions from the west periodically blanketed the region with ash, resulting in large-scale mortality, while simultaneously enriching the soil for future plant growth. Fluctuating sea levels also resulted in a variety of other environments at different times and places within the Judith River Group, including offshore and nearshore marine habitats, coastal wetlands, deltas and lagoons, in addition to the inland floodplains. The Two Medicine Formation was deposited at higher elevations farther inland than the other two formations.[20] A large variety of ceratopsians coexisted in this region, which included Achelousaurus, Brachyceratops, Cerasinops, Einiosaurus, Prenoceratops and Rubeosaurus. Carnivores included an unnamed troodontid, possibly Stenonychosaurus, the dromaeosaurs Bambiraptor and Saurornitholestes, and the large tyrannosaurids Daspletosaurus and Gorgosaurus.[21]
The excellent vertebrate fossil record of Two Medicine and Judith River rocks resulted from a combination of abundant animal life, periodic natural disasters, and the deposition of large amounts of sediment. Many types of freshwater and estuarine fish are represented, including sharks, rays, sturgeons, gars and others. This region preserves the remains of many aquatic amphibians and reptiles, including bivalves, gastropods, frogs, salamanders, turtles, Champsosaurus and crocodilians. Terrestrial lizards, including whiptails, skinks, monitors and alligator lizards have also been discovered. Pterosaurs like Montanazhdarcho and Piksi as well as birds like Apatornis and Avisaurus flew overhead. Several varieties of mammals, such as the multituberculate Cimexomys coexisted with dinosaurs in the Two Medicine Formation and the various other formations that make up the Judith River wedge. Fossilized eggs belonging to a dromaeosaur have been recovered here. When water was plentiful, the region could support a great deal of plant and animal life, but periodic droughts often resulted in mass mortality.[22]
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Anoplosaurus (meaning "unarmored or unarmed lizard") is an extinct genus of herbivorous nodosaurid dinosaur, from the late Albian-age Lower Cretaceous Cambridge Greensand of Cambridgeshire, England. It has in the past been classified with either the armored dinosaurs or the ornithopods, but current thought has been in agreement with the "armored dinosaur" interpretation, placing it in the Ankylosauria.
History
Harry Govier Seeley named this genus in 1879 for a disarticulated partial postcranial skeleton that had been uncovered at Reach, Cambridgeshire, composed of a left dentary fragment, numerous vertebrae from the neck, back, and sacrum, parts of the pectoral girdle, humerus fragments, part of the left femur, left tibia, foot bones, ribs, and other fragments. He regarded it as possibly juvenile, due to its small size,[1] with a length of about five feet. The type species is Anoplosaurus curtonotus. The generic name, derived from the Greek hoplo~, a word element used in combinations, with the meaning of "armed", refers to the fact no armour plates had been discovered. The specific name is derived from Latin curtus, "short", and Greek νῶÏον, noton, "back".
A second species, Anoplosaurus major, "the larger one", was named by Seeley in 1879 for a neck vertebra and three partial caudal vertebrae he removed from the material previously referred to Acanthopholis stereocercus, from the same formation as the type species.[1] This species now appears to be chimeric, the neck vertebra coming from an ankylosaur, the caudals from an indeterminate iguanodont.[2][3][4]
Although Seeley assigned Anoplosaurus to a general Dinosauria, he understood its possible affinities with Scelidosaurus or Polacanthus, as shown by the genus name, and other workers began to see it as an armored dinosaur.[5][6] In 1902, Baron Franz Nopcsa referred both species to Acanthopholis, creating a Acanthopholis curtonotus and a Acanthopholis major.[7] In 1923 Nopcsa suggested that, while some of the remains belonged to Acanthopholis, other remains, which he removed from that genus, belonged to a camptosaurid.[8] This suggestion led to considerable confusion, with some authors beginning to classify Anoplosaurus under the Camptosauridae,[9] a practice that was continued over several decades (with modifications as iguanodontian taxonomy changed over the years).[10]
In 1964, Oskar Kuhn renamed Syngonosaurus macrocercus Seeley 1879 into Anoplosaurus macrocercus. In 1969, Rodney Steel renamed Eucercosaurus tanyspondylus Seeley 1879 into Anoplosaurus tanyspondylus. Both Syngonosaurus and Eucercosaurus are today seen as nomina dubia and these last two Anoplosaurus species are hereby equally invalid.[2]
In 1998, Xabier Pereda-Suberbiola and Paul Barrett reexamined the material of Anoplosaurus curtonotus. They wrote that it all belonged to a "primitive" or basal member of the Nodosauridae, the lack of armor possibly due to the young age of the animal at death. The basal position would be indicated by the long tooth row and the low sacral vertebrae count. Seeley had never indicated a holotype among the syntype series. Pereda-Superbiola & Barrett therefore selected specimen SMC B55731, a right scapula piece, as the lectotype. Its ankylosaurian affinities would be proven by a high acromion process. The other nodosaurid fossils found at Reach, specimens SMC B55670 - 55742, were assigned as paralectotypes. Pereda-Superbiola & Barrett considered it possible that the discovery had in fact not been made in the Cambridge Green Sand but in the, also Albian, Upper Gault Clay, because the skeletal elements seemed to have belonged to a single individual which might preclude a provenance from the very reworked marine Green Sand deposits. Anoplosaurus curtonotus was by them considered a possibly valid taxon.[2] Reviews since then have followed this interpretation of the genus as an armored dinosaur belonging to the Ankylosauria.[11][3]
Palaeobiology
As a possible nodosaurid, Anoplosaurus would have been a quadrupedal, low-slung herbivore, with armour on its body for protection.[3]
See also
Timeline of ankylosaur research
References
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Peloroplites (from Greek pelor "monster", and hoplites, "armoured soldier") is a genus of nodosaurid armored dinosaur from Lower Cretaceous rocks of Utah, United States. It is known from a partial skull and partial postcranial remains from the base of the Mussentuchit Member of the Cedar...
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Peloroplites Scientific classification
Peloroplites (from Greek pelor "monster", and hoplites, "armoured soldier") is a genus of nodosaurid armored dinosaur from Lower Cretaceous rocks of Utah, United States. It is known from a partial skull and partial postcranial remains from the base of the Mussentuchit Member of the Cedar Mountain Formation, deposited during the Albian-Cenomanian boundary, about 104.46 to 98.37 million years ago, and was found in Emery County, Utah. It was named in 2008 by Kenneth Carpenter and colleagues.
Peloroplites was about 5 to 5.5 meters (16 to 18 ft) long, comparable to its approximate contemporary Sauropelta. It is one of the largest known nodosaurids, and came from a time when ankylosaurians in general were attaining larger sizes.
Discovery and Naming[]
In 2001, a skeleton of a large nodosaurid from the Cedar Mountain Formation in Emery County, Utah was mentioned by Burge and Bird in a publication about the faunal composition of the Price River II quarry. More material was obtained and was subsequently described in 2008 by Kenneth Carpenter, Jeff Bartlett, John Bird and Reese Barrick. The Price River II quarry was previously reported as occurring in the Ruby Ranch Member by Burge and Bird (2001) but was later reported as occurring in the base of the Mussentuchit Member due to the dark, carbonaceous nature of mudstones of the strata. The Price River II quarry has also produced specimens pertaining to four individuals of a new brachiosaurid, an iguanodontid, associated cranial and postcranial material of Cedarpelta, a turtle and a pterosaur. The holotype specimen, CEUM 26331, consists of a partial skull. Additional specimens were assigned to Peloroplites that consist of cervical vertebrae, dorsal vertebrae, synsacrums, caudal vertebrae, chevron, scapula-coracoids, humeri, radii, ulnae, ilia, pubis, ischium, femora, tibiae, fibulae, metacarpals, metatarsal, metapodials, phalanges, unguals, osteoderms and various bone fragments. The holotype and assigned specimens are currently housed at the College of Eastern Utah, Prehistoric Museum, Utah.[1]
The generic name, Peloroplites, is derived from the Greek words “peloros” (monstrous, gigantic) and “hoplites” (heavily armed), and as a subjunctive, a heavily armed soldier. The specific name, cedrimontanus, is derived from the Latin words “cedrus” (Cedar) and “mont-“ (mountain), in reference to the Cedar Mountain Formation.[1]
Carpenter et al. (2008) suggested that some of the large nodosaurid material from the Ruby Ranch Member of the Cedar Mountain Formation that has been questionably identified as Sauropelta may actually belong to Peloroplites. If the material does belong to Peloroplites, then it would extend the stratigraphic range based on a specimen described by Warren and Carpenter (2004). However, one specimen tentatively referred to Sauropelta cannot be assigned to either that genus or to Peloroplites. The specimen was obtained from the Poison Strip Sandstone Member of the Cedar Mountain Formation and assigned to Hoplitosaurus by Bodily (1969) based on the morphology of the spines. The compressed, triangular spines of the specimen are characteristic of polacanthines, which also includes Hoplitosaurus. Carpenter et al. (2008) considered that the specimen probably represents an unnamed large polacanthine.[1]
Description[]
Size and distinguishing traits[]
Carpenter et al. (2008) originally gave Peloroplites an estimated length of 5-5.5 metres (16-18 feet).[1] However, Gregory S. Paul in 2016 gave a higher estimate of 6 metres (20 feet) and a weight of 2 tonnes (4,410 lbs).[2]
Carpenter et al. (2008) diagnosed Peloroplites based on the lack of premaxillary teeth, occiput sloping forwards and towards the back, the absence of a prominent lateral temporal notch towards the back as in Sauropelta, small and blunt squamosal horns, paroccipital process projecting from the sides, a vertical quadrate that isn’t anteriorly bowed or sloped on the underside of the front, a very short odontoid, a short axis centrum which is as long as it is tall, and similar coracoid to scapula proportions to Animantarx and Edmontonia.[1]
Cranium[]
The skull of Peloroplites has an estimated length of 56 cm and a maximum width of 35.5 cm between the dorsal orbital rims, which is about the same width as Sauropelta. The snout tapers towards the front and ends at a relatively broad premaxillary beak, as compared to Silvisaurus. The premaxillae are fused along their mid-line and are dorsoventrally thick, unlike Gastonia whereas they are thin. Although the side of the left premaxilla is damaged, the width of the premaxillary beak is estimated to be 18 cm. The upper side of the premaxillae is rugose for the keratinous beak and arched in front view. In addition, the beak has a broad, inverted U-shaped notch. A groove is present near the lower margins of the beak in front view and continues to the palatal side, defining the side edge of the tomial ridge. Both the prefrontals and lachrymals are fused and the presence of the lachrymals is inferred from the lachrymal foramen seen within the front orbital wall. Both sets of prefrontal-lachrymals are triangular in upper and side view and have rugose sculpturing external surfaces that are composed of irregular pits which is especially prominent over the orbits. The front of the orbit has a faint, shallow groove which extends onto the upper surface of the prefrontal and probably outline the margins between adjoining keratinous scales, a feature also similarly seen in other nodosaurids such as Edmontonia. The prefrontal-lachrymals are divided by the front orbital wall towards the middle, which separates the orbit from the nasal cavity. The orbit has a concave upper surface. The postorbital, squamosal, jugal, quadratojugal, and quadrate are coossified on both sides of the skull. The postorbital horncores are conical structures that are very low that project dorsolaterally, which are much less prominent than those of Pawpawsaurus, Sauropelta and Gastonia. The jugal-quadratojugal horncores appear as low, localized thickening of bone as they are not prominent on the skull unlike Gastonia and Animantarx. The jugal probably composes the ventral rim of the orbit and forms a medial-lateral narrow floor to the orbit. As in other dinosaurs, the posterior rim of the orbit is composed of the postorbital and jugal. The orbit is widely separated from the lateral temporal fenestra on the sides of the skull, as in Edmontonia and Pawpawsaurus. The squamosal is fused to the head of the quadrate and the quadrates are slightly bowed towards the front. The frontoparietal region is slightly domed and is moderately arched towards the sides in posterior view. The paroccipital process faces obliquely downwards, similar to Edmontonia and Animantarx. As in other nodosaurids, the supraoccipital crest is weakly developed. The proatlas has a facet that is seen on the right side, although it is partially damaged. The exoccipital is also damaged on the left side while the exoccipital-basioccipital suture of the right side is fused. The occipital condyle has the typical shape of nodosaurids, and the condyle neck has an upper surface that is slightly concave. The basioccipital-basisphenoid suture is fused on the underside and the basioccipital is twice as long as the basisphenoid. The posterior pterygoid plate is present anterior to the parasphenoid and is concave as in Edmontonia. A tooth from a maxillary fragment is similar to some teeth referred to Priconodon and has an extensive wear facet that extends the entire face of the crown as seen in ankylosaurids.[1]
Only the rear portions of the left and right mandibles are preserved and include the articular, angular, surangular and prearticular. The posterior surface is remodelled on the sides and gives the appearance of a coossified osteoderm. However, the internal surface of the mandible is revealed by a crack on the right portion and does not support such interpretation. The adductor fossa is large and lateral-medially and is separated from the articular by a wall, both are features that are not seen in Edmontonia or Animantarx. The articular cotyle is deeper relative to the size of the bone and may correspond to the deeper and thicker retroarticular as well. The large size of the adductor fossa, along with the large teeth, suggest a strong correlation that might relate to a diet of tougher forage.[1]
Postcrania[]
The total vertebral count of Peloroplites is unknown and the atlas has not been found. The axis is nearly complete but lacks the neural spine and postzygapophyses. The fragment of neural spine is more recumbent than in Sauropelta and the centrum is almost as tall as it is long, unlike Sauropelta. Additionally, the sides of the centrum are concave, and the odontoid is short rather than long as in Sauropelta. The prezygapophysis is seen on the right side of the atlas, near the neural canal. The diapophysis is short and located at the end of the midline on the centrum which is contrast to Sauropelta as it occurs on the neural arch instead. The post-atlas cervical vertebrae consist mostly of centra except for one nearly complete anterior cervical. The centrum of this vertebra is short and slopes to the point where the posterior articular face is lower than the anterior face, a feature seen in Edmontonia but not Sauropelta or Cedarpelta. Unlike Cedarpelta, Sauropelta and Edmontonia, the articular face of Peloroplites is circular, rather than heart-shaped, hexagonal or horizontally ellipsoid. The neural spine of the anterior cervical was anteroposteriorly narrow and the anterior margin of the spine extends down between the prezygapophyses as a ridge. The neural arch is anteroposteriorly short, tall and erect which results in the postzygapophyses not reaching the level of the posterior face of the centrum. The prezygapophysis is short and angled upwards while the diapophysis is long and steeply angled towards the ventral and posterior sides. The dorsal vertebrae are represented by several vertebrae. The centra are amphiplatyan, lack nodochordal projections and are shorter than tall, specifically in the centra of the anterior dorsal vertebrae. The centrum of Peloroplites is strongly concave in the ventral margin and the prezygapophyses are steeply angled. The transverse processes are angled upwards and end in subtriangular diapophyses. In addition, a ridge extends along the transverse process ventrally to the parapophysis located on a short neural arch. As typical for other ankylosaurs, the ribs are coossified with the vertebrae. The synsacrum of Peloroplites consists of six coossified vertebrae, as in Silvisaurus. The vertebrae consist of three probably true sacrals, a dorsal and two caudal vertebrae. All the synsacral vertebrae are missing neural spines and most of the neural arches which may have been lost prior to burial. The sacral ribs are damaged with the exception for the right second rib which retains a portion of the acetabular facet. The caudal vertebrae are represented by different parts of the tail. The anterior caudal has a centrum that is wider than tall and is proportionally longer relative to height. The caudal ribs are fused to the centrum and project laterally unlike Edmontonia or Sauropelta. The distal ends of the ribs are expanded and lack a developed dorsal process. The caudal ribs also project laterally and ventrally. The caudal neural spines were expanded towards the sides. The neural arch encloses a subcircular neural canal. The centrum of the distal caudal vertebrae elongated relative to its height.[1]
The right scapula and coracoid are coossified and are almost complete. The scapula has a damaged acromion process, which appear to be pathological due to avulsion of the deltoideus clavicularis. The remains of the acromion process indicates that it had a similar position on scapula as Edmontonia. The scapular blade is intermediate in shape between curved and straight. The posterior margin of the scapula is rounded and, like most ankylosaurs, is large and deep. The coracoid almost as long as the scapula and is pierced by the coracoid foramen. Parts of the humeri are known and enough of the crest remains to show that the humeral shaft is elongated in a similar condition to Sauropelta, Edmontonia, Animantarx, and Gastonia. The radial and ulnar condyles are widely separated. The radius more closely resembles that of Edmontonia as the radius lacks the extreme expanded ends seen in Sauropelta. The ulna is long and straight unlike the bowed ulna of Sauropelta or the short and massive of Cedarpelta and Gastonia. The olecranon process partially overhangs the humeral notch. The carpals are unknown and a partial manus was found in loose association with some of the forelimb material. The manus has a complete set of metacarpals. The metacarpals have proximal ends that are faceted and fit close against one another. Metacarpal I is sub-rectangular and has a subtriangular proximal end, a condition similar to that of ceratopsians. In addition, metacarpal I is the largest in the manus and no distal condyles. The rest of the metacarpals are hourglass shaped and have expanded proximal and distal ends. Metacarpal II has a subtriangular proximal end and weakly separated distal condyles. As in other ankylosaurs, metacarpal III is the most robust and longest metacarpal in the manus. The distal condyles of metacarpal III are separate unlike Sauropelta and Nodosaurus. Metacarpal IV is roughly the same length as metacarpal I and is subpentagonal. The distal condyles of metacarpal IV form a single surface rather than being separate. The smallest metacarpal in the series is metacarpal V, although it is slightly more robust than metacarpal IV. Metacarpal V has a rounded distal end, with no development of condyles that are separated. All the phalanges are anteroposteriorly short, have very shallow proximal articular surfaces and lack distal condyles. Due to the shortness of the phalanges, the lateral collateral ligament pits are absent. Phalanx I-1 is the largest and the longest of the phalanxes. The distal unguals are disc-shapes as they are wider than long and rounded.[1]
A 2011 study by Philip J. Senter suggested that the metacarpal configuration of Peloroplites and other ankylosaurs were positioned in a vertical semi-tube configuration, similar to that of sauropods, as the metacarpals are wedge-shaped in proximal view which fit tightly in such position without gaps between the proximal gaps and proximal surfaces.[3] A right ilium and left pubis represent the only material of the pelvis. The preacetabular process of the pelvis diverges 55° unlike Sauropelta where it diverges 39° and 28° in Edmontonia. However, the degree of divergence might possibly be an artefact of not having a complete medial surface. The postacetabular process is broad and short. Additionally, the lateral surface is almost straight rather than concave. The pubis is robust and has a lateral face that forms the anterior wall of the acetabulum. The preacetabular process is short, straight and thick, while the postpubic process is short and angled posteroventrally. Both the left and right femora are complete and are relatively straight shafted. The head of the femora are set at a slight upward angle. The left femur preserves an oblique transverse ridge that is present below the greater trochanter but is damaged and better seen on the right femur. The cnemial crest of the tibia is short and rounded in profile. The tibial shaft is thick throughout its length and astragalus is not fused to the distal end of the tibia. Only a right metatarsal and ungual are known of the hindfoot. The metatarsal is proportionally shorter and more robust than the metacarpals of the manus. As in Sauropelta, the proximal end of the metatarsal is sloped laterally in anterior view and the distal condyles are well developed. The ungual is broad throughout its length.[1]
Phylogeny[]
Carpenter et al. (2008) originally placed Peloroplites within Nodosauridae but did not conduct a phylogenetic analysis to determine its exact relationships within the clade.[1] Thompson et al. (2012) recovered Peloroplites as sister taxon to Polacanthus, a position also recovered by Chen et al. (2013).[4][5] However, Yang et al. (2013) found Peloroplites to be sister taxon to both Taohelong and Polacanthus, while Zheng et al. (2018) found it to be sister taxon to Taohelong and a large clade containing more nested taxa such as Nodosaurus, Edmontonia, Struthiosaurus and Europelta.[6] Rivera-Sylva et al. (2018) placed Peloroplites as sister taxon to Sauropelta, Taohelong and a clade containg more nested taxa.[7]
A phylogenetic analysis conducted by Rivera-Sylva et al. (2018) and modified by Madzia et al. (2021) is reproduced below.[7][8]
Nodosauridae
Sauroplites Mymoorapelta Dongyangopelta
Gastonia
Gargoyleosaurus
Polacanthinae
Hoplitosaurus Polacanthus
Nodosaurinae
Peloroplites
Taohelong Sauropelta
Acantholipan Nodosaurus
Niobrarasaurus Ahshislepelta
Tatankacephalus
Silvisaurus
CPC 273
Panoplosaurini
Animantarx
Panoplosaurus
’’Patagopelta’’ Texasetes Denversaurus Edmontonia longiceps Edmontonia rugosidens
Struthiosaurini
Hungarosaurus Europelta
Pawpawsaurus Stegopelta Struthiosaurus languedocensis
Struthiosaurus transylvanicus Struthiosaurus austriacus
The results of an earlier analysis by Thompson et al. (2012) are reproduced below.[4]
Nodosauridae
Antarctopelta
Mymoorapelta
Hylaeosaurus Anoplosaurus
Tatankacephalus Horshamosaurus Polacanthinae
Gargoyleosaurus Hoplitosaurus
Gastonia
Peloroplites Polacanthus
Struthiosaurus Zhejiangosaurus
Hungarosaurus
Animantarx
Niobrarasaurus Nodosaurus Pawpawsaurus Sauropelta Silvisaurus Stegopelta Texasetes
Edmontonia Panoplosaurus
Paleobiology[]
Peloroplites is known from the uppermost part of the Cedar Mountain Formation, a layer known as the Mussentuchit Member. The layer was originally interpreted as being of Aptian to Albian age (∼109–116 Ma) using radiometric dating.[1] However, other radioistopic datings places the Mussentuchit Member in the Cenomanian to early Turonian age (98.2 ± 0.6 to 93 Ma).[9] The Mussentuchit Member has been interpreted as a either a wet, lacustrine environment or a fluvial environment like a distal delta system at the western margin of the Western Interior Seaway.[9]
Peloroplites was contemporaneous with the basal hadrosauromorph Eolambia,[10] the brachiosaurid sauropod Abydosaurus,[11] the tyrannosauroid Moros,[12] the carcharodontosaurian Siats, the indeterminate coelurosaur Richardoestesia,[13] the ankylosaurs Animantarx and Cedarpelta,[14][1] the thescelosaurid cf. Zephyrosaurus,[15] an indeterminate neoceratopsian,[15] an indeterminate pachycephalosaurid,[15] an indeterminate velociraptorine[15] and an indeterminate dromaeosaurine.[15] Non-dinosaur taxa contemporaneous with Peloroplites include the crocodylomorphs Dakotasuchus,[16] cf. Bernissartia[15] and Machimosaurus,[15] the lizards Bicuspidon, Dimekodontosaurus, cf. Pseudosaurillus, Bothriagenys, Harmodontosaurus, Dicothodon and Primaderma,[17] the snake Coniophis,[17] the amphibian Albanerpeton,[15] the turtles Glyptops and Naomichelys,[15] the mammals Astroconodon, Spalacotherium, Symmetrodontoides, Paracimexomys and Kokopellia,[15] the batoids Ischyrhiza and cf. Baibisha,[15] the hybodonts Polyacrodus, Lissodus and Hybodus,[15] and the lungfish Ceratodus.[15]
In the Media[]
Peloroplites appeared in Jurassic World: Fallen Kingdom, but only a carcass and skeleton seen on Isla Nublar and Lockwood Manor.
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Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago, although the exact origin and timing of the evolution of dinosaursis the subject of active research. They became the dominant terrestrial...
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Prehistoric Wiki
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https://prehistoric-wiki.fandom.com/wiki/List_of_dinosaur_genera
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Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago, although the exact origin and timing of the evolution of dinosaursis the subject of active research. They became the dominant terrestrial vertebrates after the Triassic–Jurassic extinction event 201.3 million years ago; their dominance continued throughout the Jurassic and Cretaceousperiods. The fossil record demonstrates that birds are modern feathered dinosaurs, having evolved from earlier theropods during the Late Jurassic epoch. Birds were therefore the only dinosaur lineage to survive the Cretaceous–Paleogene extinction event approximately 66 million years ago. Dinosaurs can be divided into avian dinosaurs (birds) and non-avian dinosaurs, which are all dinosaurs other than birds. Birds are feathered theropod dinosaursand constitute the only known living dinosaurs. [[null|link=https://en.m.wikipedia.org/wiki/File:Field_dinos_2.jpg%7Cthumb%7C260x260px%7CMounted skeletons of Tyrannosaurus(left) and Apatosaurus (right) at the AMNH]] This list of dinosaurs is a comprehensive listing of all genera that have ever been considered to be non-avian dinosaurs, but also includes some dinosaurs of disputed status (avian? or non-avian?, where "avian" refers to the clade Avialae), as well as purely vernacular terms.
The list includes all commonly accepted genera, but also genera that are now considered invalid, doubtful (nomen dubium), or were not formally published (nomen nudum), as well as junior synonyms and genera that are no longer considered dinosaurs. Many listed names have been reclassified as everything from true birds to crocodilians to petrified wood. The list contains 1764 names, of which approximately 1328 are considered either valid dinosaur genera or nomina dubia.
Scope and terminology[]
There is no official, canonical list of all non-avian dinosaur genera. The closest is the Dinosaur Genera List, compiled by biological nomenclature expert George Olshevsky, which was first published online in 1995 and was regularly updated until June 2021. The most authoritative general source in the field is the second (2004) edition of The Dinosauria. The vast majority of names listed below are sourced to Olshevsky's list, and all subjective determinations (such as junior synonymy or non-dinosaurian status) are based on The Dinosauria, except where they conflict with primary literature. These exceptions are noted.
Naming conventions and terminology follow the International Code of Zoological Nomenclature. Technical terms used include:
Junior synonym: A name which describes the same taxon as a previously published name. If two or more genera are formally designated and the type specimens are later assigned to the same genus, the first to be published (in chronological order) is the senior synonym, and all other instances are junior synonyms. Senior synonyms are generally used, except by special decision of the ICZN (see Tyrannosaurus), but junior synonyms cannot be used again for a different genus, even if deprecated. Junior synonymy is often subjective, unless the genera described were both based on the same type specimen.
Nomen nudum (Latin for "naked name"): A name that has appeared in print but has not yet been formally published by the standards of the ICZN. Nomina nuda (the plural form) are invalid, and are therefore not italicized as a proper generic name would be. If the name is later formally published, that name is no longer a nomen nudum and will be italicized on this list. Often, the formally published name will differ from any nomina nuda that describe the same specimen.
Nomen oblitum (Latin for "forgotten name"): A name that has not been used in the scientific community for more than fifty years after its original proposal.
Nomen manuscriptum (Latin for "manuscript name"): A name that appears in manuscript of a formal publication that has no scientific backing.
Preoccupied name: A name that is formally published, but which has already been used for another taxon. This second use is invalid (as are all subsequent uses) and the name must be replaced. Preoccupied names are not valid generic names.
Nomen dubium (Latin for "dubious name"): A name describing a fossil with no unique diagnostic features. As this can be an extremely subjective and controversial designation (see Hadrosaurus), no genera should be marked as such on this list.
A[]
Aachenosaurus – subsequently found to be a piece of petrified wood
Aardonyx
"Abdallahsaurus" – nomen nudum, synonym of Giraffatitan
Abdarainurus
Abditosaurus
Abelisaurus
Abrictosaurus
Abrosaurus
Abydosaurus
Acantholipan
Acanthopholis
Achelousaurus
Acheroraptor
Achillesaurus
Achillobator
Acristavus
Acrocanthosaurus
[[null|link=https://en.m.wikipedia.org/wiki/File:Allosaurus_SDNHM.jpg%7Calt=%7Cthumb%7CReplica of an Allosaurus skeleton.]]
Acrotholus
Actiosaurus – subsequently found to be a choristoderan
Adamantisaurus
Adasaurus
Adelolophus
Adeopapposaurus
Adratiklit
Adynomosaurus
Aegyptosaurus
Aeolosaurus
Aepisaurus
Aepyornithomimus
Aerosteon
Aetonyx – junior synonym of Massospondylus
Afromimus
Afrovenator
Agathaumas – possible synonym of Triceratops
Aggiosaurus – subsequently found to be a metriorhynchid crocodilian
Agilisaurus
Agnosphitys – possibly non-dinosaurian
Agrosaurus – probably a junior synonym of Thecodontosaurus
Agujaceratops
Agustinia
Ahshislepelta
"Airakoraptor" – nomen nudum; Kuru
Ajancingenia – synonym of Heyuannia
Ajkaceratops
Ajnabia
Akainacephalus
Alamosaurus
Alaskacephale
Albalophosaurus
Albertaceratops
Albertadromeus
Albertavenator
Albertonykus
Albertosaurus
Albinykus
Albisaurus – subsequently found to be a non-dinosaurian reptile
Alcovasaurus – possible junior synonym of Miragaia
Alectrosaurus
Aletopelta
[[null|link=https://en.m.wikipedia.org/wiki/File:Amargasaurus_Reconstruction_Fred_Wierum.png%7Calt=%7Cthumb%7CArtist's reconstruction of Amargasaurus.]]
Algoasaurus
Alioramus
Aliwalia – junior synonym of Eucnemesaurus
Allosaurus
Almas
Alnashetri
Alocodon
Altirhinus
Altispinax
Alvarezsaurus
Alwalkeria
Alxasaurus
Amanasaurus – possibly non-dinosaurian
Amanzia
Amargasaurus
"Amargastegos" – nomen nudum
Amargatitanis
Amazonsaurus
Ambopteryx
Ammosaurus – junior synonym of Anchisaurus
Ampelognathus
Ampelosaurus
Amphicoelias
"Amphicoelicaudia" – nomen nudum; synonym of Huabeisaurus
"Amphisaurus" – preoccupied name, now known as Anchisaurus
Amtocephale
Amtosaurus – possibly a junior synonym of Talarurus
Amurosaurus
Amygdalodon
Anabisetia
Analong
Anasazisaurus
Anatosaurus – junior synonym of Edmontosaurus
Anatotitan – junior synonym of Edmontosaurus
Anchiceratops
Anchiornis
Anchisaurus
Andesaurus
[[null|link=https://en.m.wikipedia.org/wiki/File:Anzu_wyliei.jpg%7Calt=%7Cthumb%7CArtist's reconstruction of Anzu.]]
"Andhrasaurus" – nomen nudum
Angaturama – possible junior synonym of Irritator
"Angloposeidon" – nomen nudum
Angolatitan
Angulomastacator
Anhuilong
Aniksosaurus
Animantarx
Ankistrodon – subsequently found to be a proterosuchid archosauriform
Ankylosaurus
Anodontosaurus
Anomalipes
Anoplosaurus
Anserimimus
Antarctopelta
Antarctosaurus
Antetonitrus
Anthodon – subsequently found to be a pareiasaur
Antrodemus – possibly a synonym of Allosaurus
Anzu
Aoniraptor
Aorun
Apatodon – possibly a junior synonym of Allosaurus
Apatoraptor
Apatosaurus
Appalachiosaurus
Aquilarhinus
Aquilops
Arackar
Aragosaurus
Aralosaurus
Aratasaurus
"Araucanoraptor" – nomen nudum; Neuquenraptor
Archaeoceratops
Archaeodontosaurus
[[null|link=https://en.m.wikipedia.org/wiki/File:Archaeoceratops_BW.jpg%7Calt=%7Cthumb%7CArtist's restoration of Archaeoceratops.]]
Archaeopteryx – possibly a bird
Archaeoraptor – a chimaera of the birdYanornis and the dromaeosaur Microraptor
Archaeornis – junior synonym of Archaeopteryx
Archaeornithoides
Archaeornithomimus
Arcovenator
Arctosaurus – subsequently found to be a non-dinosaurian reptile
Arcusaurus
Arenysaurus
Argentinosaurus
Argyrosaurus
Aristosaurus – junior synonym of Massospondylus
Aristosuchus
Arizonasaurus – subsequently found to be a rauisuchian
Arkansaurus
Arkharavia
Arrhinoceratops
Arrudatitan
Arstanosaurus
Asfaltovenator
Asiaceratops
Asiamericana – a fish
Asiatosaurus
Asilisaurus – possibly non-dinosaurian
Astrodon
Astrodonius – junior synonym of Astrodon
Astrodontaurus – junior synonym of Astrodon
Astrophocaudia
Asylosaurus
Atacamatitan
Atlantosaurus
Atlasaurus
Atlascopcosaurus
Atrociraptor
Atsinganosaurus
[[null|link=https://en.m.wikipedia.org/wiki/File:Austroraptor_Restoration.png%7Calt=%7Cthumb%7CArtist's restoration of Austroraptor.]]
Aublysodon
Aucasaurus
"Augustia" – preoccupied name, now known as Agustinia
Augustynolophus
Auroraceratops
Aurornis
Australodocus
Australotitan
Australovenator
Austrocheirus
Austroposeidon
Austroraptor
Austrosaurus
Avaceratops
"Avalonia" – preoccupied name, now known as Avalonianus
Avalonianus – subsequently found to be a non-dinosaurian archosaur
Aviatyrannis
Avimimus
Avipes – probably a non-dinosaurian dinosauromorph
Avisaurus – subsequently found to be an enantiornithine bird
Azendohsaurus – subsequently found to be a non-dinosaurian archosauromorph
B[]
Baalsaurus
Bactrosaurus
Bagaceratops
Bagaraatan
Bagualia
Bagualosaurus
Bahariasaurus
Bainoceratops
Bajadasaurus
"Bakesaurus" – nomen nudum; Bactrosaurus
Balaur – possibly a bird
"Balochisaurus" – nomen nudum
Bambiraptor
Banji
Bannykus
[[null|link=https://en.m.wikipedia.org/wiki/File:Baryonyx_walkeri_mount_NMNS.jpg%7Calt=%7Cthumb%7CReconstructed skeletal mount of Baryonyx at the National Museum of Nature and Science, Tokyo.]]
Baotianmansaurus
Barapasaurus
Barilium
Barosaurus
Barrosasaurus
Barsboldia
Baryonyx
Bashanosaurus
Bashunosaurus
Basutodon – subsequently found to be a non-dinosaurian archosaur
Bathygnathus – a pelycosaur, Dimetrodon
Batyrosaurus
Baurutitan
Bayannurosaurus
"Bayosaurus" – nomen nudum
Becklespinax – junior synonym of Altispinax
"Beelemodon" – nomen nudum
Beg
Beibeilong
Beipiaognathus – chimera of several unnamed dinosaurs
Beipiaosaurus
Beishanlong
Bellusaurus
Belodon – subsequently found to be a phytosaur
Berberosaurus
Berthasaura
Betasuchus
Bicentenaria
Bienosaurus
"Bihariosaurus" – nomen nudum
"Bilbeyhallorum" – nomen nudum; Cedarpelta
Bissektipelta
Bistahieversor
Bisticeratops
"Blancocerosaurus" – nomen nudum, synonym of Giraffatitan
Blasisaurus
Blikanasaurus
Bolong
Bonapartenykus
Bonapartesaurus
Bonatitan
Bonitasaura
Borealopelta
Borealosaurus
Boreonykus
Borogovia
Bothriospondylus
Brachiosaurus
Brachyceratops
Brachylophosaurus
Brachypodosaurus
Brachyrophus – junior synonym of Camptosaurus
[[null|link=https://en.m.wikipedia.org/wiki/File:Borealopelta_NT.jpg%7Calt=%7Cthumb%7CArtist's restoration of Borealopelta.]]
Brachytaenius – subsequently found to be a metriorhynchid; junior objective synonym of Dakosaurus or Geosaurus
Brachytrachelopan
Bradycneme
Brasileosaurus – subsequently found to be a non-dinosaurian archosaur
Brasilotitan
Bravasaurus
Bravoceratops
Breviceratops
Brighstoneus
Brohisaurus
Brontomerus
"Brontoraptor" – nomen nudum, synonym of Torvosaurus
Brontosaurus
Bruhathkayosaurus
Bugenasaura – junior synonym of Thescelosaurus
Buitreraptor
Burianosaurus
Buriolestes
Bustingorrytitan
"Byranjaffia" – nomen nudum; Byronosaurus
Byronosaurus
C[]
Caenagnathasia
Caenagnathus
Caieiria
Caihong
Calamosaurus
"Calamospondylus" – preoccupied name, now known as Calamosaurus
Calamospondylus
Callovosaurus
Calvarius
Camarasaurus
Camarillasaurus
Camelotia
Camposaurus
"Camptonotus" – preoccupied name, now known as Camptosaurus
Camptosaurus
"Campylodon" – preoccupied name, now known as Campylodoniscus
Campylodoniscus
Canardia
"Capitalsaurus" – nomen nudum
Carcharodontosaurus
Cardiodon
Carnotaurus
Caseosaurus
Cathartesaura
Cathetosaurus — possibly a species of Camarasaurus [[null|link=https://en.m.wikipedia.org/wiki/File:Centrosaurus.JPG%7Calt=%7Cthumb%7CCentrosaurus skull.]]
Caudipteryx
Caudocoelus – junior synonym of Teinurosaurus
Caulodon – junior synonym of Camarasaurus
Cedarosaurus
Cedarpelta
Cedrorestes
Centemodon – subsequently found to be a phytosaur
Centrosaurus
Cerasinops
Ceratonykus
Ceratops
Ceratosaurus
Ceratosuchops
Cetiosauriscus
Cetiosaurus
Chakisaurus
Changchunsaurus
"Changdusaurus" – nomen nudum
Changmiania
Changyuraptor
Chaoyangsaurus
Charonosaurus
Chasmosaurus
Chassternbergia – junior synonym of Edmontonia
Chebsaurus
Chenanisaurus
Cheneosaurus – junior synonym of Hypacrosaurus
Chialingosaurus
Chiayusaurus
Chienkosaurus – possible junior synonym of Szechuanosaurus
"Chihuahuasaurus" – nomen nudum; Sonorasaurus
Chilantaisaurus
Chilesaurus
Chindesaurus
Chingkankousaurus
Chinshakiangosaurus
Chirostenotes
Choconsaurus
Chondrosteosaurus [[null|link=https://en.m.wikipedia.org/wiki/File:Ceratosaurus_nasicornis_DB.jpg%7Calt=%7Cthumb%7CArtist's restoration of Ceratosaurus.]]
Choyrodon
Chromogisaurus
Chuandongocoelurus
Chuanjiesaurus
Chuanqilong
Chubutisaurus
Chucarosaurus
Chungkingosaurus
Chuxiongosaurus
"Cinizasaurus" – nomen nudum
Cionodon
Citipati
Citipes
Cladeiodon – subsequently found to be a non-dinosaurian rauisuchian; synonym of Teratosaurus
Claorhynchus – possibly Triceratops
Claosaurus
Clarencea – subsequently found to be a sphenosuchian; synonym of Sphenosuchus
Clasmodosaurus
Clepsysaurus – subsequently found to be a phytosaur, possibly Palaeosaurus
Coahuilaceratops
Coelophysis
"Coelosaurus" – preoccupied genus name, species "Coelosaurus" antiquus
Coeluroides
Coelurosauravus – subsequently found to be a primitive diapsid
Coelurus
Colepiocephale
"Coloradia" – preoccupied name, now known as Coloradisaurus
Coloradisaurus
"Colossosaurus" – nomen nudum; Pelorosaurus
Comahuesaurus
"Comanchesaurus" – nomen nudum
Compsognathus [[null|link=https://en.m.wikipedia.org/wiki/File:Coelophysis_bauri_mount.jpg%7Calt=%7Cthumb%7CCoelophysis mounted skeleton at the Cleveland Museum of Natural History.]]
Compsosuchus
Concavenator
Conchoraptor
Condorraptor
Convolosaurus
Coronosaurus
Corythoraptor
Corythosaurus
Craspedodon
Crataeomus – junior synonym of Struthiosaurus
Craterosaurus
Creosaurus – junior synonym of Allosaurus
Crichtonpelta
Crichtonsaurus
Cristatusaurus
Crittendenceratops
Crosbysaurus – subsequently found to be a non-dinosaurian archosauriform
Cruxicheiros
Cryolophosaurus
Cryptodraco – junior synonym (unneeded replacement name) of Cryptosaurus
"Cryptoraptor" – nomen nudum
Cryptosaurus
Cryptovolans – junior synonym of Microraptor
Cumnoria
D[]
Daanosaurus
Dacentrurus
"Dachongosaurus" – nomen nudum
Daemonosaurus
Dahalokely
Dakosaurus – subsequently found to be a metriorhynchid crocodilian
Dakotadon
Dakotaraptor
Daliansaurus
"Damalasaurus" – nomen nudum
Dandakosaurus
Danubiosaurus – junior synonym of Struthiosaurus
"Daptosaurus" – nomen nudum; early manuscript name for Deinonychus
Darwinsaurus – junior synonym of Hypselospinus or Mantellisaurus
Dashanpusaurus
Daspletosaurus
Dasygnathoides – subsequently found to be a non-dinosaurian archosaur; possible junior synonym of Ornithosuchus
"Dasygnathus" – preoccupied name, now known as Dasygnathoides
Datai
Datanglong [[null|link=https://en.m.wikipedia.org/wiki/File:Hypothetical_Deinocheirus.jpg%7Calt=%7Cthumb%7CArtist's restoration of Deinocheirus.]]
Datonglong
Datousaurus
Daurlong
Daurosaurus – synonym of Kulindadromeus
Daxiatitan
Deinocheirus
Deinodon – possibly Gorgosaurus
Deinonychus
Delapparentia – junior synonym of Iguanodon
Deltadromeus
Demandasaurus
Denversaurus
Deuterosaurus – subsequently found to be a therapsid
Diabloceratops
Diamantinasaurus
Dianchungosaurus – subsequently found to be a crocodilian
"Diceratops" – preoccupied name, now known as Nedoceratops
Diceratus – junior synonym of Nedoceratops
Diclonius
Dicraeosaurus
Didanodon – synonym of Lambeosaurus; possibly a nomen nudum
Dilong
Dilophosaurus
Diluvicursor
Dimodosaurus – junior synonym of Plateosaurus
Dineobellator
Dinheirosaurus – possible junior synonym of Supersaurus
Dinodocus
"Dinosaurus" – preoccupied name for a junior synonym of Brithopus; now a junior synonym of Plateosaurus [[null|link=https://en.m.wikipedia.org/wiki/File:Diamantinasaurus.png%7Calt=%7Cthumb%7CArtist's restoration of Diamantinasaurus.]]
Dinotyrannus – junior synonym Tyrannosaurus or some other tyrannosaurid
Diodorus – possibly non-dinosaurian
Diplodocus
Diplotomodon
Diracodon – junior synonym of Stegosaurus
Dolichosuchus
Dollodon – junior synonym of Mantellisaurus
"Domeykosaurus" – nomen nudum, synonym of Arackar
Dongbeititan
Dongyangopelta
Dongyangosaurus
Doratodon – subsequently found to be a crocodilian
Doryphorosaurus – junior synonym (unneeded replacement name) of Kentrosaurus
Draconyx
Dracopelta
Dracoraptor
Dracorex – junior synonym of Pachycephalosaurus
Dracovenator
Dravidosaurus – possibly non-dinosaurian
Dreadnoughtus
Drinker – junior synonym of Nanosaurus
Dromaeosauroides
Dromaeosaurus
Dromiceiomimus
Dromicosaurus – junior synonym of Massospondylus
Drusilasaura
Dryosaurus [[null|link=https://en.m.wikipedia.org/wiki/File:Dryosaurus_lettowvorbecki_skeleton.jpg%7Calt=%7Cthumb%7CA Dysalotosaurus skeleton.]]
Dryptosauroides
Dryptosaurus
Dubreuillosaurus
"Duranteceratops" – nomen nudum
Duriatitan
Duriavenator
Dynamosaurus – junior synonym of Tyrannosaurus
Dynamoterror
Dyoplosaurus
Dysalotosaurus
Dysganus
Dyslocosaurus
Dystrophaeus
Dystylosaurus – junior synonym of Supersaurus
Dzharaonyx
Dzharatitanis
E[]
Echinodon
Edmarka – junior synonym of Torvosaurus
Edmontonia
Edmontosaurus
Efraasia [[null|link=https://en.m.wikipedia.org/wiki/File:Eoraptor_Japan.jpg%7Calt=%7Cthumb%7CReplica of an Eoraptor skeleton.]]
Einiosaurus
Ekrixinatosaurus
Elachistosuchus – subsequently found to be a rhynchocephalian
Elaltitan
Elaphrosaurus
Elemgasem
Elmisaurus
Elopteryx
Elosaurus – junior synonym of Brontosaurus
Elrhazosaurus
"Elvisaurus" – nomen nudum; Cryolophosaurus
Emausaurus
Embasaurus
Enigmosaurus
Eoabelisaurus
Eobrontosaurus – junior synonym of Brontosaurus
Eocarcharia
Eoceratops – junior synonym of Chasmosaurus
Eocursor
Eodromaeus
"Eohadrosaurus" – nomen nudum; Eolambia
Eolambia
Eomamenchisaurus
Eoneophron
"Eoplophysis" – nomen nudum
Eoraptor
Eosinopteryx
Eotrachodon
Eotriceratops
Eotyrannus
Eousdryosaurus
Epachthosaurus
Epanterias – may be Allosaurus
"Ephoenosaurus" – nomen nudum; Machimosaurus (a crocodilian)
Epicampodon – subsequently found to be a proterosuchid archosauriform, Ankistrodon
Epichirostenotes
Epidendrosaurus – synonym of Scansoriopteryx
Epidexipteryx
Equijubus
Erectopus
Erketu [[null|link=https://en.m.wikipedia.org/wiki/File:Euoplocephalus_BW.jpg%7Calt=%7Cthumb%7CLife restoration of Euoplocephalus.]]
Erliansaurus
Erlikosaurus
Erythrovenator
Eshanosaurus
"Euacanthus" – nomen nudum; junior synonym of Polacanthus
Eucamerotus
Eucentrosaurus – junior synonym (unneeded replacement name) of Centrosaurus
Eucercosaurus
Eucnemesaurus
Eucoelophysis – possibly non-dinosaurian
"Eugongbusaurus" – nomen nudum
Euhelopus
Euoplocephalus
Eupodosaurus – subsequently found to be a nothosaur synonymous with Lariosaurus
"Eureodon" – nomen nudum; Tenontosaurus
Eurolimnornis – subsequently found to be a pterosaur
Euronychodon
Europasaurus
Europatitan
Europelta
Euskelosaurus
Eustreptospondylus
F[]
Fabrosaurus – possibly Lesothosaurus
Falcarius
"Fendusaurus" – nomen nudum
"Fenestrosaurus" – nomen nudum; Oviraptor
Ferganasaurus
"Ferganastegos" – nomen nudum
Ferganocephale
Ferrisaurus
Foraminacephale
Fosterovenator
Fostoria
Frenguellisaurus – junior synonym of Herrerasaurus
[[null|link=https://en.m.wikipedia.org/wiki/File:Fruitadens_NT.jpg%7Calt=%7Cthumb%7CLife restoration of Fruitadens.]]
Fruitadens
Fujianvenator
Fukuiraptor
Fukuisaurus
Fukuititan
Fukuivenator
Fulengia
Fulgurotherium
Furcatoceratops
Fushanosaurus
"Fusinasus" – nomen nudum; Eotyrannus
Fusuisaurus
"Futabasaurus" – nomen nudum; not to be confused with the formally named plesiosaurFutabasaurus
Futalognkosaurus
Fylax
G[]
"Gadolosaurus" – nomen nudum
Galeamopus
Galesaurus – subsequently found to be a therapsid
Galleonosaurus
Gallimimus
Galtonia – subsequently found to be a pseudosuchian; possibly a junior synonym of Revueltosaurus
Galveosaurus – synonym of Galvesaurus
Galvesaurus
Gamatavus — possibly non-dinosaurian
Gandititan
Gannansaurus
"Gansutitan" – nomen nudum; Daxiatitan
Ganzhousaurus
Gargoyleosaurus
Garrigatitan
Garudimimus
Garumbatitan
Gasosaurus [[null|link=https://en.m.wikipedia.org/wiki/File:Giganotos_Db.jpg%7Calt=%7Cthumb%7CArtist's restoration of Giganotosaurus.]]
Gasparinisaura
Gastonia
"Gavinosaurus" – nomen nudum; Eotyrannus
Geminiraptor
Genusaurus
Genyodectes
Geranosaurus
Gideonmantellia
Giganotosaurus
Gigantoraptor
"Gigantosaurus" – preoccupied name, now known as Tornieria
Gigantosaurus
Gigantoscelus – Probable junior synonym of Euskelosaurus
Gigantspinosaurus
Gilmoreosaurus
"Ginnareemimus" – nomen nudum; Kinnareemimus
Giraffatitan
Glacialisaurus
Glishades
Glyptodontopelta
Gnathovorax
Gobiceratops – possibly a junior synonym of Bagaceratops
Gobihadros
Gobiraptor
Gobisaurus
Gobititan
Gobivenator
"Godzillasaurus" – nomen nudum; Gojirasaurus
Gojirasaurus
Gondwanatitan
Gongbusaurus
Gongpoquansaurus
Gongxianosaurus
Gonkoken
Gorgosaurus
Goyocephale [[null|link=https://en.m.wikipedia.org/wiki/File:Giraffatitan_DB.jpg%7Calt=%7Cthumb%7CArtist's reconstruction of Giraffatitan.]]
Graciliceratops
Graciliraptor
Gracilisuchus – subsequently found to be a non-dinosaurian archosaur
Gravitholus
Gremlin
Gresslyosaurus – possible junior synonym of Plateosaurus
Griphornis – junior synonym of Archaeopteryx
Griphosaurus – junior synonym of Archaeopteryx
Gryphoceratops
Gryponyx
Gryposaurus
"Gspsaurus" – nomen nudum
Guaibasaurus
Gualicho
Guanlong
Guemesia
Gwyneddosaurus – subsequently found to be a tanystrophid
Gyposaurus – possibly a junior synonym of Massospondylus
H[]
"Hadrosauravus" – nomen nudum; junior synonym of Gryposaurus
Hadrosaurus
Haestasaurus
Hagryphus
Hallopus – subsequently found to be a crocodylomorph
Halszkaraptor
Halticosaurus
Hamititan
Hanssuesia
"Hanwulosaurus" – nomen nudum
Haplocanthosaurus
"Haplocanthus" – preoccupied name, now known as Haplocanthosaurus
Haplocheirus
Harpymimus
Haya
Hecatasaurus – junior synonym of Telmatosaurus
"Heilongjiangosaurus" – nomen nudum
Heishansaurus
Helioceratops [[null|link=https://en.m.wikipedia.org/wiki/File:Huaxiagnathus_orientalis.JPG%7Calt=%7Cthumb%7CHuaxiagnathus fossil displayed in Hong Kong Science Museum]]
"Helopus" – preoccupied name, now known as Euhelopus
Heptasteornis
Herbstosaurus – subsequently found to be a pterosaur
Herrerasaurus
Hesperonychus
Hesperonyx
Hesperornithoides
Hesperosaurus
Heterodontosaurus
Heterosaurus – possible synonym of Mantellisaurus
Hexing
Hexinlusaurus
Heyuannia
Hierosaurus
Hikanodon – junior synonym of Iguanodon
Hippodraco
"Hironosaurus" – nomen nudum
"Hisanohamasaurus" – nomen nudum
Histriasaurus
Homalocephale
"Honghesaurus" – nomen nudum later described as Yandusaurus; name later used for a genus of marine reptile
Hongshanosaurus – junior synonym of Psittacosaurus
Hoplitosaurus
Hoplosaurus – junior synonym of Struthiosaurus
Horshamosaurus
Hortalotarsus – possible junior synonym of Massospondylus
Huabeisaurus
Hualianceratops
Huallasaurus
Huanansaurus
Huanghetitan
Huangshanlong
Huaxiagnathus
Huaxiaosaurus – junior synonym of Shantungosaurus
"Huaxiasaurus" – nomen nudum; Huaxiagnathus
Huayangosaurus
Hudiesaurus
Huehuecanauhtlus [[null|link=https://en.m.wikipedia.org/wiki/File:Hypsilophodon_foxii_1.jpg%7Calt=%7Cthumb%7CSkeleton of Hypsilophodon.]]
Huinculsaurus
Hulsanpes
Hungarosaurus
Huxleysaurus – junior synonym of Hypselospinus
Hylaeosaurus
Hylosaurus – junior synonym of Hylaeosaurus
Hypacrosaurus
Hypselorhachis – subsequently found to be a ctenosauriscid
Hypselosaurus
Hypselospinus
Hypsibema
Hypsilophodon
Hypsirhophus
I[]
Iani
Iberospinus
Ibirania
"Ichabodcraniosaurus" – nomen nudum; Shri
Ichthyovenator
Igai
Ignavusaurus
Ignotosaurus – possibly non-dinosaurian
Iguanacolossus
Iguanodon
"Iguanoides" – nomen nudum; Iguanodon
"Iguanosaurus" – nomen nudum; Iguanodon
Iliosuchus
Ilokelesia
Imperobator
Inawentu
Incisivosaurus
Indosaurus
Indosuchus
"Ingenia" – preoccupied name, now known as Heyuannia yanshini
Ingentia
Inosaurus
Invictarx
Irisosaurus
[[null|link=https://en.m.wikipedia.org/wiki/File:Iguanodon_NT.jpg%7Calt=%7Cthumb%7CArtist's reconstruction of Iguanodon.]]
Irritator
Isaberrysaura
Isanosaurus
Isasicursor
Ischioceratops
Ischisaurus – junior synonym of Herrerasaurus
"Ischyrosaurus" – preoccupied genus name, species Ischyrosaurus manseli
Isisaurus
"Issasaurus" – nomen nudum; Dicraeosaurus
Issi
Itapeuasaurus
Itemirus
Iuticosaurus
Iyuku
J[]
Jaculinykus
Jainosaurus
Jakapil
Jaklapallisaurus
Janenschia
Jaxartosaurus
Jeholosaurus
Jenghizkhan – junior synonym of Tarbosaurus
"Jensenosaurus" – nomen nudum; Supersaurus
Jeyawati
Jianchangosaurus
"Jiangjunmiaosaurus" – nomen nudum; Monolophosaurus
Jiangjunosaurus
Jiangshanosaurus [[null|link=https://en.m.wikipedia.org/wiki/File:Jinfengopteryx_wiki.jpg%7Calt=%7Cthumb%7CLife restoration of Jinfengopteryx.]]
Jiangxisaurus
Jiangxititan
Jianianhualong
Jinbeisaurus
Jinfengopteryx
"Jingia" – preoccupied name, has not yet been renamed
Jingshanosaurus
Jintasaurus
Jinyunpelta
Jinzhousaurus
Jiutaisaurus
Jobaria
Jubbulpuria – possible junior synonym of Laevisuchus
Judiceratops
Jurapteryx – junior synonym of Archaeopteryx
"Jurassosaurus" – nomen nudum; Tianchisaurus
Juratyrant
Juravenator
K[]
Kaatedocus
"Kagasaurus" – nomen nudum
Kaijiangosaurus
Kaijutitan
Kakuru
Kamuysaurus
Kangnasaurus
Kansaignathus
Karongasaurus
Katepensaurus
"Katsuyamasaurus" – nomen nudum
Kayentavenator
Kazaklambia
Kelmayisaurus
Kelumapusaura
Kemkemia – subsequently found to be a crocodyliform
Kentrosaurus
Kentrurosaurus – junior synonym (unneeded replacement name) of Kentrosaurus
Kerberosaurus
[[null|link=https://en.m.wikipedia.org/wiki/File:Berlin_Naturkundemuseum_Dino_Eingangshalle.jpg%7Calt=%7Cthumb%7CKentrosaurus skeleton.]]
Khaan
"Khetranisaurus" – nomen nudum
Kholumolumo
Khulsanurus
Kileskus
Kinnareemimus
"Kitadanisaurus" – nomen nudum; Fukuiraptor
"Kittysaurus" – nomen nudum; Eotyrannus
Klamelisaurus
Kol
Koparion
Koreaceratops
Koreanosaurus
"Koreanosaurus" – nomen nudum; name later used formally for a genus of ornithopod
Koshisaurus
Kosmoceratops
Kotasaurus
Koutalisaurus – possible junior synonym of Pararhabdodon
Kritosaurus
Kryptops
Krzyzanowskisaurus – probably a pseudosuchian (Revueltosaurus?)
Kukufeldia – junior synonym of Barilium
Kulceratops
Kulindadromeus
Kulindapteryx – synonym of Kulindadromeus
Kunbarrasaurus
Kundurosaurus
"Kunmingosaurus" – nomen nudum
Kuru
Kurupi
Kuszholia – subsequently found to be a bird
Kwanasaurus – possibly non-dinosaurian
L[]
Labocania
Labrosaurus – junior synonym of Allosaurus
[[null|link=https://en.m.wikipedia.org/wiki/File:Lambeosaurus_magnicristatus_DB.jpg%7Calt=%7Cthumb%7CArtist's restoration of Lambeosaurus.]]
"Laelaps" – preoccupied name, now known as Dryptosaurus
Laevisuchus
Lagerpeton – subsequently found to be a non-dinosaurian pterosauromorph
Lagosuchus – subsequently found to be a non-dinosaurian dinosauromorph
Laiyangosaurus
Lajasvenator
Lamaceratops – possible junior synonym of Bagaceratops
Lambeosaurus
Lametasaurus
Lamplughsaura
Lanasaurus – junior synonym of Lycorhinus
"Lancangosaurus" – variant spelling of "Lancanjiangosaurus"
"Lancanjiangosaurus" – nomen nudum
Lanzhousaurus
Laosaurus
Lapampasaurus
Laplatasaurus
Lapparentosaurus
Laquintasaura
Latenivenatrix – possible junior synonym of Stenonychosaurus
Latirhinus
Lavocatisaurus
Leaellynasaura
Ledumahadi
Leinkupal
Leipsanosaurus – junior synonym of Struthiosaurus
"Lengosaurus" – nomen nudum; Eotyrannus
Leonerasaurus
Lepidocheirosaurus — junior synonym of Kulindadromeus
Lepidus
[[null|link=https://en.m.wikipedia.org/wiki/File:Limusaurus_runner.jpg%7Calt=%7Cthumb%7CArtist's restoration of Limusaurus.]]
Leptoceratops
Leptorhynchos
Leptospondylus – junior synonym of Massospondylus
Leshansaurus
Lesothosaurus
Lessemsaurus
Levnesovia
Lewisuchus – possibly non-dinosaurian
Lexovisaurus
Leyesaurus
Liaoceratops
Liaoningosaurus
Liaoningotitan
Liaoningvenator
"Liassaurus" – nomen nudum; possible synonym of Sarcosaurus
Libycosaurus – subsequently found to be an anthracothere mammal
Ligabueino
Ligabuesaurus
"Ligomasaurus" – nomen nudum, synonym of Giraffatitan
"Likhoelesaurus" – nomen nudum; possibly non-dinosaurian
Liliensternus
Limaysaurus
"Limnornis" – preoccupied name, now known as Palaeocursornis ( a pterosaur)
"Limnosaurus" – preoccupied name, now known as Telmatosaurus
Limusaurus
Lingwulong
Lingyuanosaurus
Linhenykus
Linheraptor
Linhevenator
[[null|link=https://en.m.wikipedia.org/wiki/File:Linhenykus_monodactylus.jpg%7Calt=%7Cthumb%7CLife reconstruction of two individuals of Linhenykusin their arid Campanian-aged living environment.]]
Lirainosaurus
Lisboasaurus – subsequently found to be a crocodilian
Liubangosaurus
Llukalkan
Lohuecotitan
Loncosaurus
Longisquama – subsequently found to be a non-dinosaurian reptile
Longosaurus – junior synonym of Coelophysis
Lophorhothon
Lophostropheus
Loricatosaurus
Loricosaurus
Losillasaurus
Lourinhanosaurus
Lourinhasaurus
Luanchuanraptor
"Luanpingosaurus" – nomen nudum; Psittacosaurus
Lucianosaurus – subsequently found to be a non-dinosaurian archosauriform
Lucianovenator
Lufengosaurus
Lukousaurus – possibly a crurotarsan
Luoyanggia
Lurdusaurus
Lusitanosaurus
Lusotitan
Lusovenator
Lutungutali – possibly non-dinosaurian
Lycorhinus
Lythronax
M[]
Macelognathus – subsequently found to be a sphenosuchian crocodilian
Machairasaurus
Machairoceratops
Macrocollum
Macrodontophion – subsequently found to be a member of Lophotrochozoa
Macrogryphosaurus
Macrophalangia – junior synonym of Chirostenotes [[null|link=https://en.m.wikipedia.org/wiki/File:Massospondylus_reconstruction.png%7Calt=%7Cthumb%7CArtist's reconstruction of Massospondylus.]]
"Macroscelosaurus" – nomen nudum; junior synonym of Tanystropheus
Macrurosaurus
"Madsenius" – nomen nudum, Allosaurus
Magnamanus
Magnapaulia
Magnirostris – possible junior synonym of Bagaceratops
Magnosaurus
"Magulodon" – nomen nudum
Magyarosaurus
Mahakala
Mahuidacursor
Maiasaura
Maip
Majungasaurus
Majungatholus – junior synonym of Majungasaurus
Malarguesaurus
Malawisaurus
Maleevosaurus – junior synonym of Tarbosaurus
Maleevus
Malefica
Mamenchisaurus
Mandschurosaurus
Manidens
Manospondylus – synonym of Tyrannosaurus
Mansourasaurus
Mantellisaurus
Mantellodon – junior synonym of Mantellisaurus
"Maojandino" – nomen nudum
Mapusaurus
Maraapunisaurus
Marasuchus – subsequently found to be a non-dinosaurian dinosauromorph
"Marisaurus" – nomen nudum
Marmarospondylus
Marshosaurus
Martharaptor
Masiakasaurus
Massospondylus [[null|link=https://en.m.wikipedia.org/wiki/File:Microraptor_Restoration.png%7Calt=%7Cthumb%7CArtist's reconstruction of Microraptor with colouration based on fossilised melanosomes.]]
Matheronodon
Maxakalisaurus
Mbiresaurus
Medusaceratops
"Megacervixosaurus" – nomen nudum
"Megadactylus" – preoccupied name, now known as Anchisaurus
"Megadontosaurus" – nomen nudum; Microvenator
Megalosaurus
Megapnosaurus – possible junior synonym of Coelophysis
Megaraptor
Mei
Melanorosaurus
Mendozasaurus
Menefeeceratops
Menucocelsior
Meraxes
Mercuriceratops
Meroktenos
Metriacanthosaurus
"Microcephale" – nomen nudum
"Microceratops" – preoccupied name, now known as Microceratus
Microceratus
Microcoelus
"Microdontosaurus" – nomen nudum
Microhadrosaurus
Micropachycephalosaurus
Microraptor
Microvenator
Mierasaurus
"Mifunesaurus" – nomen nudum
Migmanychion
Minimocursor
Minmi
Minotaurasaurus
Minqaria
Miragaia
Mirischia
Mnyamawamtuka
Moabosaurus
Mochlodon
"Mohammadisaurus" – nomen nudum; Tornieria [[null|link=https://en.m.wikipedia.org/wiki/File:Muttaburrasaurus_skel_QM_email.jpg%7Calt=%7Cthumb%7CCast of a Muttaburrasaurusskeleton.]]
Mojoceratops – junior synonym of Chasmosaurus
Mongolosaurus
Mongolostegus
Monkonosaurus
Monoclonius
Monolophosaurus
"Mononychus" – preoccupied name, now known as Mononykus
Mononykus
Montanoceratops
Morelladon
Morinosaurus
Moros
Morosaurus – junior synonym of Camarasaurus
Morrosaurus
Mosaiceratops
"Moshisaurus" – nomen nudum; possibly Mamenchisaurus
"Mtapaiasaurus" – nomen nudum, synonym of Giraffatitan
"Mtotosaurus" – nomen nudum; Dicraeosaurus
Murusraptor
Mussaurus
Muttaburrasaurus
Muyelensaurus
Mymoorapelta
N[]
Naashoibitosaurus
Nambalia
Nankangia
Nanningosaurus
Nanosaurus
Nanotyrannus – junior synonym of Tyrannosaurus
Nanshiungosaurus
Nanuqsaurus
Nanyangosaurus
Napaisaurus
Narambuenatitan
Narindasaurus
Nasutoceratops
Natovenator
"Natronasaurus" – invalid name, either Alcovasaurus or Miragaia
[[null|link=https://en.m.wikipedia.org/wiki/File:Neimongosaurus.jpg%7Calt=%7Cthumb%7CLife restoration of Neimongosaurus.]]
Navajoceratops
Nebulasaurus
"Nectosaurus" – preoccupied name, now known as Kritosaurus
Nedcolbertia
Nedoceratops – possible junior synonym of Triceratops
Neimongosaurus
"Nemegtia" – preoccupied name, now known as Nemegtomaia
Nemegtomaia
Nemegtonykus
Nemegtosaurus
"Neosaurus" – preoccupied name; renamed Parrosaurus, which is now Hypsibema
Neosodon
Neovenator
Neuquenraptor
Neuquensaurus
Nevadadromeus
"Newtonsaurus" – nomen nudum, possibly Zanclodon
"Ngexisaurus" – nomen nudum
Ngwevu
Nhandumirim
Nicksaurus – nomen manuscriptum
Niebla
Nigersaurus
Ningyuansaurus
Ninjatitan
Niobrarasaurus
Nipponosaurus
Noasaurus
Nodocephalosaurus
Nodosaurus
Nomingia – possible junior synonym of Elmisaurus
Nopcsaspondylus
Normanniasaurus
[[null|link=https://en.m.wikipedia.org/wiki/File:Nemegtomaia_barsboldi_profile1.jpg%7Calt=%7Cthumb%7CRestored head of Nemegtomaia.]]
Notatesseraeraptor
Nothronychus
Notoceratops
Notocolossus
Notohypsilophodon
Nqwebasaurus
"Nteregosaurus" – nomen nudum; Janenschia
Nullotitan
"Nurosaurus" – nomen nudum
Nuthetes
Nyasasaurus — possibly non-dinosaurian
"Nyororosaurus" – nomen nudum; Dicraeosaurus
O[]
Oblitosaurus
Oceanotitan
Ohmdenosaurus
Ojoceratops – possible synonym of Eotriceratops
Ojoraptorsaurus
Oksoko
Oligosaurus – possible synonym of Mochlodon
Olorotitan
Omeisaurus
"Omosaurus" – preoccupied name, now known as Dacentrurus
Ondogurvel
Onychosaurus – junior synonym of Zalmoxesor Rhabdodon, or an ankylosaurian
Oohkotokia
Opisthocoelicaudia
Oplosaurus
"Orcomimus" – nomen nudum
Orinosaurus – junior synonym (unneeded replacement name) of Orosaurus
Orkoraptor
Ornatops
Ornatotholus – junior synonym of Stegoceras
Ornithodesmus
"Ornithoides" – nomen nudum; Saurornithoides
Ornitholestes
Ornithomerus – possible synonym of Mochlodon
Ornithomimoides
Ornithomimus
Ornithopsis
[[null|link=https://en.m.wikipedia.org/wiki/File:Omeisaurus_tianfuensis34.jpg%7Calt=%7Cthumb%7CArtist's reconstruction of Omeisaurus.]]
Ornithosuchus – subsequently found to be a non-dinosaurian archosaur
Ornithotarsus – junior synonym of Hadrosaurus
Orodromeus
Orosaurus
Orthogoniosaurus
Orthomerus
Oryctodromeus
"Oshanosaurus" – nomen nudum
Osmakasaurus
Ostafrikasaurus
Ostromia
Othnielia – junior synonym of Nanosaurus
Othnielosaurus – junior synonym of Nanosaurus
Otogosaurus — possibly a nomen nudum
Ouranosaurus
Overoraptor
Overosaurus
Oviraptor
"Ovoraptor" – nomen nudum; Velociraptor
Owenodon
Oxalaia – possible junior synonym of Spinosaurus
Ozraptor
P[]
Pachycephalosaurus
Pachyrhinosaurus
Pachysauriscus – junior synonym of Plateosaurus
Pachysaurops – junior synonym of Plateosaurus
"Pachysaurus" – preoccupied name, now known as Pachysauriscus; junior synonym of Plateosaurus
Pachyspondylus – junior synonym of Massospondylus
Pachysuchus
Padillasaurus
"Pakisaurus" – nomen nudum
Palaeoctonus – subsequently found to be a phytosaur
Palaeocursornis – subsequently found to be an azhdarchoid pterosaur
"Palaeolimnornis" – nomen nudum; Palaeocursornis, pterodactyloid pterosaur belonging to Azhdarchoidea
Palaeopteryx – possibly a bird
Palaeosauriscus – junior synonym of Palaeosaurus
Palaeosaurus – subsequently found to be a non-dinosaurian reptile
"Palaeosaurus" – preoccupied name, now known as Sphenosaurus and considered to be a non-dinosaurian procolophonid
Palaeoscincus
Paleosaurus – subsequently found to be a non-dinosaurian archosaur; junior synonym (unneeded replacement name) of Palaeosaurus
Paludititan
Paluxysaurus – junior synonym of Sauroposeidon
Pampadromaeus
[[null|link=https://en.m.wikipedia.org/wiki/File:Pachycephalosaurus_Reconstruction.jpg%7Calt=%7Cthumb%7CLife restoration of Pachycephalosaurus.]]
Pamparaptor
Panamericansaurus
Pandoravenator
Panguraptor
Panoplosaurus
Panphagia
Pantydraco – possible synonym of Thecodontosaurus
Papiliovenator
"Paraiguanodon" – nomen nudum; Bactrosaurus
Paralitherizinosaurus
Paralititan
Paranthodon
Pararhabdodon
Parasaurolophus
Paraxenisaurus
Pareiasaurus – subsequently found to be a pareiasaur
Pareisactus
Parksosaurus
Paronychodon
Parrosaurus – now known as Hypsibema missouriensis
Parvicursor
Patagonykus
Patagopelta
Patagosaurus
Patagotitan
Pawpawsaurus
Pectinodon
Pedopenna
Pegomastax
Peishansaurus
Pekinosaurus – subsequently found to be a pseudosuchian; junior synonym of Revueltosaurus
Pelecanimimus
Pellegrinisaurus
Peloroplites
Pelorosaurus
"Peltosaurus" – preoccupied name, now known as Sauropelta
Pendraig
Penelopognathus
Pentaceratops
[[null|link=https://en.m.wikipedia.org/wiki/File:Plateosaurus_BW.jpg%7Calt=%7Cthumb%7CArtist's restoration of Plateosaurus.]]
Perijasaurus
Petrobrasaurus
Phaedrolosaurus
Philovenator
Phuwiangosaurus
Phuwiangvenator
Phyllodon
Piatnitzkysaurus
Picrodon – possibly non-dinosaurian
Pilmatueia
Pinacosaurus
Pisanosaurus — possibly non-dinosaurian
Pitekunsaurus
Piveteausaurus
Planicoxa
Plateosauravus
Plateosaurus
Platyceratops – possible junior synonym of Bagaceratops
Platypelta
Platytholus
Plesiohadros
Pleurocoelus – possible junior synonym of Astrodon
Pleuropeltus – junior synonym of Struthiosaurus
Pneumatoarthrus – subsequently found to be a turtle
Pneumatoraptor
Podokesaurus
Poekilopleuron
Polacanthoides – chimera of Hylaeosaurusand Polacanthus
Polacanthus
Polyodontosaurus
Polyonax
Ponerosteus – subsequently found to be a non-dinosaurian archosaur
Poposaurus – subsequently found to be a non-dinosaurian archosaur
Portellsaurus
Postosuchus – subsequently found to be a rauisuchian
Powellvenator
Pradhania
Prenocephale
Prenoceratops
Priconodon
Priodontognathus
[[null|link=https://en.m.wikipedia.org/wiki/File:Prosaurolophus_maximus2.JPG%7Calt=%7Cthumb%7CProsaurolophus skull.]]
Proa
Probactrosaurus
Probrachylophosaurus
Proceratops – junior synonym (unneeded replacement name) of Ceratops
Proceratosaurus
Procerosaurus – subsequently found to be a tanystropheid protorosaur, Tanystropheus
"Procerosaurus" – preoccupied name, now known as Ponerosteus
Procheneosaurus – junior synonym of Lambeosaurus
Procompsognathus
Prodeinodon
"Proiguanodon" – nomen nudum; Iguanodon
Propanoplosaurus
Proplanicoxa – junior synonym of Mantellisaurus
Prosaurolophus
Protarchaeopteryx
Protathlitis
Protecovasaurus – subsequently found to be a non-dinosaurian archosauriform
Protiguanodon – junior synonym of Psittacosaurus
Protoavis – described as a bird, probably a chimera including theropod dinosaur bones
Protoceratops
Protognathosaurus
"Protognathus" – preoccupied name, now known as Protognathosaurus
Protohadros
"Protorosaurus" – preoccupied name, now known as Chasmosaurus
Protorosaurus – subsequently found to be a non-dinosaurian reptile
"Proyandusaurus" – nomen nudum; Hexinlusaurus.
Pseudolagosuchus – possibly non-dinosaurian; a junior synonym of Lewisuchus
Psittacosaurus
Pteropelyx
Pterospondylus
Puertasaurus
Pukyongosaurus
Pulanesaura
Punatitan
Pycnonemosaurus
Pyroraptor
Q[]
Qantassaurus
Qianlong
[[null|link=https://en.m.wikipedia.org/wiki/File:Qiupalong_Restoration.png%7Calt=%7Cthumb%7CArtist's reconstruction of Qiupalong.]]
Qianzhousaurus
Qiaowanlong
Qijianglong
Qingxiusaurus
Qinlingosaurus
Qiupalong
Qiupanykus
Quaesitosaurus
Quetecsaurus
Quilmesaurus
R[]
Rachitrema – subsequently found to be a chimera primarily based on ichthyosaurfossils
Rahiolisaurus
"Rahona" – preoccupied name, now known as Rahonavis
Rahonavis – possibly a bird
Rajasaurus
Rapator
Rapetosaurus
Raptorex – possible junior synonym of Tarbosaurus
Ratchasimasaurus
Rativates
Rayososaurus
Razanandrongobe – subsequently found to be a crocodylomorph
Rebbachisaurus
Regaliceratops
Regnosaurus
Revueltosaurus – subsequently found to be a pseudosuchian
Rhabdodon
Rhadinosaurus – may be non-dinosaurian, possibly crocodilian
Rhinorex – possible synonym of Gryposaurus
Rhodanosaurus – junior synonym of Struthiosaurus
Rhoetosaurus
Rhomaleopakhus
Rhopalodon – subsequently found to be a synapsid
Riabininohadros
Richardoestesia [[null|link=https://en.m.wikipedia.org/wiki/File:Ruyangosaurus_skeleton.jpg%7Calt=%7Cthumb%7CRuyangosaurus skeleton.]]
"Rileya" – preoccupied name, now known as Rileyasuchus
Rileyasuchus – subsequently found to be a phytosaur
Rinchenia
Rinconsaurus
Rioarribasaurus – junior synonym of Coelophysis
"Riodevasaurus" – nomen nudum; Turiasaurus
Riojasaurus
Riojasuchus – subsequently found to be a non-dinosaurian archosaur
Riojavenatrix
Riparovenator
Rocasaurus
"Roccosaurus" – nomen nudum; Melanorosaurus
"Ronaldoraptor" – nomen nudum
Rubeosaurus – junior synonym of Styracosaurus
Ruehleia
Rugocaudia
Rugops
Ruixinia
Rukwatitan
Ruyangosaurus
S[]
Sacisaurus – possibly non-dinosaurian
Sahaliyania
Saichania
"Saldamosaurus" – nomen nudum
"Salimosaurus" – nomen nudum, synonym of Giraffatitan
Saltasaurus
Saltopus – possibly non-dinosaurian
"Saltriosaurus" – nomen nudum
Saltriovenator
"Sanchusaurus" – nomen nudum, possible Gallimimus
"Sangonghesaurus" – nomen nudum, Tianchisaurus
Sanjuansaurus
Sanpasaurus
Santanaraptor
Sanxiasaurus
Sarahsaurus
Saraikimasoom – nomen manuscriptum
Sarcolestes
Sarcosaurus
Sarmientosaurus
Saturnalia
"Sauraechinodon" – nomen nudum; Echinodon
[[null|link=https://en.m.wikipedia.org/wiki/File:Scelidosaurus_harrisonii.png%7Calt=%7Cthumb%7CArtist's restoration of Scelidosaurus.]]
"Sauraechmodon" – nomen nudum; Echinodon
"Saurechinodon" – nomen nudum; Echinodon
Saurolophus
Sauroniops
Sauropelta
Saurophaganax
"Saurophagus" – preoccupied name, now known as Saurophaganax
Sauroplites
Sauroposeidon
Saurornithoides
Saurornitholestes
Savannasaurus
Scansoriopteryx
Scaphonyx – subsequently found to be a rhynchosaur, Hyperodapedon
Scelidosaurus
Schleitheimia
Scipionyx
Sciurumimus
Scleromochlus – subsequently found to be a non-dinosaurian avemetatarsalian
Scolosaurus
Scutellosaurus
Secernosaurus
Sefapanosaurus
Segisaurus
Segnosaurus
Seismosaurus – junior synonym of Diplodocus
Seitaad
Sektensaurus
"Selimanosaurus" – nomen nudum; Dicraeosaurus
Sellacoxa – junior synonym of Barilium
Sellosaurus – junior synonym of Plateosaurus
Serendipaceratops
Serikornis
Shamosaurus
Shanag
Shanshanosaurus – junior synonym of Tarbosaurus
Shantungosaurus
Shanxia
Shanyangosaurus
Shaochilong
[[null|link=https://en.m.wikipedia.org/wiki/File:Sinosaurus_triassicus.JPG%7Calt=%7Cthumb%7CSinosaurus skeleton, Museo delle Scienze of Trento, Italy.]]
Shenzhousaurus
Shidaisaurus
Shingopana
Shishugounykus
Shixinggia
Shri
Shuangbaisaurus – possible synonym of Sinosaurus
Shuangmiaosaurus
Shunosaurus
Shuvosaurus – subsequently found to be a rauisuchian
Shuvuuia
Siamodon
"Siamodracon" – nomen nudum
Siamosaurus
Siamotyrannus
Siamraptor
Siats
"Sibirosaurus" – nomen nudum, now known as Sibirotitan
Sibirotitan
Sidersaura
"Sidormimus" – nomen nudum
Sierraceratops
Sigilmassasaurus – possible junior synonym of Spinosaurus
Silesaurus – possibly non-dinosaurian
Siluosaurus
Silutitan
Silvisaurus
Similicaudipteryx
Sinankylosaurus
Sinocalliopteryx
Sinocephale
Sinoceratops
Sinocoelurus
"Sinopelta" – nomen nudum; synonym of Sinopeltosaurus
"Sinopeltosaurus" – nomen nudum
Sinopliosaurus – a pliosaur; one species, "S." fusuiensis, is actually a dinosaur that may be synonymous with Siamosaurus
Sinornithoides
[[null|link=https://en.m.wikipedia.org/wiki/File:Skorpiovenator_skull.jpg%7Calt=%7Cthumb%7CSkorpiovenator skull.]]
Sinornithomimus
Sinornithosaurus
Sinosauropteryx
Sinosaurus
Sinotyrannus
Sinovenator
Sinraptor
Sinusonasus
Sirindhorna
Skorpiovenator
"Smilodon" – preoccupied name, now known as Zanclodon
Smitanosaurus
Smok — possibly non-dinosaurian
Sonidosaurus
Sonorasaurus
Soriatitan
Soumyasaurus – possibly non-dinosaurian
Spectrovenator
Sphaerotholus
Sphenosaurus – subsequently found to be a non-dinosaurian reptile
Sphenospondylus – junior synonym of Mantellisaurus
Spiclypeus
Spicomellus
Spinophorosaurus
Spinops
Spinosaurus
Spinostropheus
Spinosuchus – subsequently found to be a non-dinosaurian reptile
Spondylosoma – subsequently found to be an aphanosaur
Squalodon – subsequently found to be a cetacean
Staurikosaurus
Stegoceras
Stegopelta
Stegosaurides
Stegosaurus
Stegouros
Stellasaurus
Stenonychosaurus
[[null|link=https://en.m.wikipedia.org/wiki/File:Stegosaurus_stenops_sophie_wiki_martyniuk.png%7Calt=%7Cthumb%7CArtist's restoration of Stegosaurus.]]
Stenopelix
Stenotholus – junior synonym of Stygimoloch, which is a possible junior synonym of Pachycephalosaurus
Stephanosaurus
"Stereocephalus" – preoccupied name, now known as Euoplocephalus
Sterrholophus – junior synonym of Triceratops
Stokesosaurus
Stormbergia – junior synonym of Lesothosaurus
Strenusaurus – junior synonym of Riojasaurus
Streptospondylus
Struthiomimus
Struthiosaurus
Stygimoloch – junior synonym of Pachycephalosaurus
Stygivenator – junior synonym of Tyrannosaurus
Styracosaurus
Succinodon – subsequently found to be fossilized mollusc borings
Suchomimus
Suchoprion – subsequently found to be a phytosaur
Suchosaurus – possible synonym of Baryonyx
"Sugiyamasaurus" – nomen nudum
"Sulaimanisaurus" – nomen nudum
Supersaurus
Suskityrannus
Suuwassea
Suzhousaurus
Symphyrophus – junior synonym of Camptosaurus
Syngonosaurus
"Syntarsus" – preoccupied name, sometimes assigned to Coelophysis or Megapnosaurus
Syrmosaurus – junior synonym of Pinacosaurus
Szechuanosaurus
T[]
Tachiraptor [[null|link=https://en.m.wikipedia.org/wiki/File:Thecondontosaurus_life_restoration_2018.jpg%7Calt=%7Cthumb%7CArtist's restoration of Thecodontosaurus.]]
Talarurus
Talenkauen
Talos
Tamarro
Tambatitanis
Tangvayosaurus
Tanius
Tanycolagreus
Tanystropheus – subsequently found to be a protorosaur
Tanystrosuchus
Taohelong
Tapinocephalus – subsequently found to be a therapsid
Tapuiasaurus
Tarascosaurus
Tarbosaurus
Tarchia
Tastavinsaurus
Tatankacephalus
Tatankaceratops – probable junior synonym of Triceratops
Tataouinea
Tatisaurus
Taurovenator
Taveirosaurus
Tawa
Tawasaurus – junior synonym of Lufengosaurus
Tazoudasaurus
Technosaurus – possibly non-dinosaurian
Tecovasaurus – subsequently found to be a non-dinosaurian archosauriform
Tehuelchesaurus
"Teihivenator" – nomen nudum
Teinurosaurus
Teleocrater – subsequently found to be a basal avemetatarsalian
Telmatosaurus
"Tenantosaurus" – nomen nudum; Tenontosaurus
"Tenchisaurus" – nomen nudum; an unpublished museum name for Tianchisaurus
Tendaguria
Tengrisaurus
Tenontosaurus
Teratophoneus
Teratosaurus – subsequently found to be a non-dinosaurian archosaur
Termatosaurus – subsequently found to be a phytosaur
Terminocavus [[null|link=https://en.m.wikipedia.org/wiki/File:Coast_watch_(1979)_(20472080340).jpg%7Calt=%7Cthumb%7CThescelosaurus fossil.]]
Tethyshadros
Tetragonosaurus – junior synonym of Lambeosaurus
Texacephale
Texasetes
Teyuwasu – possibly junior synonym of Staurikosaurus
Thanatotheristes
Thanos
Tharosaurus
Thecocoelurus
Thecodontosaurus
Thecospondylus
Theiophytalia
Therizinosaurus
Therosaurus – a synonym of Iguanodon
Thescelosaurus
Thespesius
"Thotobolosaurus" – nomen nudum; Kholumolumo
Thyreosaurus
Tianchiasaurus – alternate spelling of Tianchisaurus
Tianchisaurus
"Tianchungosaurus" – nomen nudum; Dianchungosaurus (crocodilian)
Tianyulong
Tianyuraptor
Tianzhenosaurus
Tichosteus
Tienshanosaurus
Tietasaura
Timimus
Timurlengia
Titanoceratops
Titanomachya
Titanosaurus
"Titanosaurus" – preoccupied name, now known as Atlantosaurus
Tlatolophus
Tochisaurus
"Tomodon" – preoccupied name, now known as Diplotomodon
Tonganosaurus
Tongtianlong
"Tonouchisaurus" – nomen nudum
Torilion – junior synonym of Barilium
Tornieria
Torosaurus [[null|link=https://en.m.wikipedia.org/wiki/File:Triceratops_Specimen_at_the_Houston_Museum_of_Natural_Science.JPG%7Calt=%7Cthumb%7CSkeleton of Triceratops at the Houston Museum of Natural Science.]]
Torvosaurus
Tototlmimus
Trachelosaurus – subsequently found to be a basal archosauromorph
Trachodon
Tralkasaurus
Transylvanosaurus
Tratayenia
Traukutitan
Trialestes – subsequently found to be a basal crocodylomorph
"Triassolestes" – preoccupied name, now known as Trialestes
Tribelesodon – junior synonym of Tanystropheus, a protorosaur
Triceratops
Trierarchuncus
Trigonosaurus
Trimucrodon
Trinisaura
Triunfosaurus
Troodon
Tsaagan
Tsagantegia
Tsintaosaurus
Tuebingosaurus
Tugulusaurus
Tuojiangosaurus
Turanoceratops
Turiasaurus
Tylocephale
Tylosteus – synonym of Pachycephalosaurus
Tyrannomimus
Tyrannosaurus
Tyrannotitan
U[]
Uberabatitan
"Ubirajara" – nomen nudum
Udanoceratops
Udelartitan
Ugrosaurus – junior synonym of Triceratops
Ugrunaaluk – junior synonym of Edmontosaurus
Uintasaurus – junior synonym of Camarasaurus
Ultrasauros – junior synonym of Supersaurus
"Ultrasaurus" – preoccupied name, renamed Ultrasauros which is now a junior synonym of Supersaurus
Ultrasaurus
Ulughbegsaurus
"Umarsaurus" – nomen nudum; Barsboldia
Unaysaurus
Unenlagia
Unescoceratops
"Unicerosaurus" – nomen nudum, subsequently found to be a fish
Unquillosaurus
Urbacodon
Utahceratops
Utahraptor
Uteodon
V[]
Vagaceratops
Vahiny
Valdoraptor – possible synonym of Thecocoelurus
Valdosaurus
Vallibonavenatrix
Variraptor
Vayuraptor
Vectaerovenator
Vectensia – junior synonym of Polacanthusor Hylaeosaurus
Vectidromeus
Vectipelta
Vectiraptor
[[null|link=https://en.m.wikipedia.org/wiki/File:Velociraptor_Restoration.png%7Calt=%7Cthumb%7CLife restoration of Velociraptor.]]
Vectisaurus – junior synonym of Mantellisaurus
Velafrons
Velocipes
Velociraptor
Velocisaurus
Venaticosuchus – subsequently found to be a non-dinosaurian archosaur
Venenosaurus
Vespersaurus
Veterupristisaurus
Viavenator
"Vitakridrinda" – nomen nudum
"Vitakrisaurus" – nomen nudum
Volgatitan
Volkheimeria
Vouivria
Vulcanodon
W[]
Wadhurstia – junior synonym of Hypselospinus
Wakinosaurus
Walgettosuchus – possible synonym of Rapator
"Walkeria" – preoccupied name, now known as Alwalkeria
"Walkersaurus" – nomen nudum; Duriavenator
Wamweracaudia
"Wangonisaurus" – nomen nudum, synonym of Giraffatitan
Wannanosaurus
Weewarrasaurus
[[null|link=https://en.m.wikipedia.org/wiki/File:%D0%9D%D0%BE%D0%B2%D0%B0%D1%8F_%D1%80%D0%B5%D0%BA%D0%BE%D0%BD%D1%81%D1%82%D1%80%D1%83%D0%BA%D1%86%D0%B8%D1%8F_%D0%9C%D0%BE%D0%BD%D1%81%D1%82%D1%80%D0%B0_%D0%B8%D0%B7_%D0%9C%D0%B8%D0%BD%D0%B4%D0%B5%D0%BD%D0%B0.jpg%7Calt=%7Cthumb%7CArtist's restoration of Wiehenvenator.]]
Wellnhoferia – subsequently found to be a bird, possible junior synonym of Archaeopteryx
Wendiceratops
Wiehenvenator
Willinakaqe
Wintonotitan
Wuerhosaurus
Wulagasaurus
Wulatelong
Wulong
Wyleyia – subsequently found to be a bird
"Wyomingraptor" – nomen nudum, synonym of Allosaurus
X[]
Xenoceratops
Xenoposeidon
Xenotarsosaurus
Xianshanosaurus
Xiaosaurus
[[null|link=https://en.m.wikipedia.org/wiki/File:Xiongguanlong_NT.jpg%7Calt=%7Cthumb%7CArtist's reconstruction of Xiongguanlong.]]
Xiaotingia
Xingtianosaurus
Xingxiulong
Xinjiangovenator
Xinjiangtitan
Xiongguanlong
Xixianykus
Xixiasaurus
Xixiposaurus
Xiyunykus
Xuanhanosaurus
Xuanhuaceratops
"Xuanhuasaurus" – nomen nudum; Xuanhuaceratops
Xunmenglong
Xuwulong
Y[]
Yaleosaurus – junior synonym of Anchisaurus
Yamaceratops
Yamanasaurus
Yamatosaurus
Yanbeilong
Yandusaurus
Yangchuanosaurus
Yaverlandia
Yehuecauhceratops
"Yezosaurus" – nomen nudum; subsequently found to be a junior synonym of the mosasaur Taniwhasaurus
Yi
"Yibinosaurus" – nomen nudum [[null|link=https://en.m.wikipedia.org/wiki/File:Yangchuanosaurus_NT_small.jpg%7Calt=%7Cthumb%7CArtist's restoration of Yangchuanosaurus.]]
Yimenosaurus
Yingshanosaurus
Yinlong
Yixianosaurus
Yizhousaurus
Yongjinglong
Ypupiara
"Yuanmouraptor" – nomen nudum
Yuanmousaurus
Yueosaurus
Yulong
Yunganglong
Yunmenglong
Yunnanosaurus
"Yunxianosaurus" – nomen nudum
Yunyangosaurus
Yurgovuchia
Yutyrannus
Yuxisaurus
Yuzhoulong
Z[]
Zalmoxes
Zanabazar
Zanclodon – subsequently found to be non-dinosaurian
Zapalasaurus
Zapsalis
Zaraapelta
Zatomus – subsequently found to be a non-dinosaurian archosaur
Zby
Zephyrosaurus
Zhanghenglong
Zhejiangosaurus
Zhenyuanlong
[[null|link=https://en.m.wikipedia.org/wiki/File:Zby_NT_small.jpg%7Calt=%7Cthumb%7CLife restoration of Zby.]]
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https://www.academia.edu/7402314/JUVENILE_SAUROLOPHINE_SPECIMENS_DINOSAURIA_HADROSAURIDAE_FROM_THE_LATE_CAMPANIAN_OF_NORTHEASTERN_MEXICO
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en
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JUVENILE SAUROLOPHINE SPECIMENS (DINOSAURIA: HADROSAURIDAE) FROM THE LATE CAMPANIAN OF NORTHEASTERN MEXICO
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2014-06-19T00:00:00
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JUVENILE SAUROLOPHINE SPECIMENS (DINOSAURIA: HADROSAURIDAE) FROM THE LATE CAMPANIAN OF NORTHEASTERN MEXICO
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https://www.academia.edu/7402314/JUVENILE_SAUROLOPHINE_SPECIMENS_DINOSAURIA_HADROSAURIDAE_FROM_THE_LATE_CAMPANIAN_OF_NORTHEASTERN_MEXICO
|
The scientific publications of the Natural History Museum of Los Angeles County have been issued at irregular intervals in three major series; the articles in each series are numbered individually, and the numbers run consecutively, regardless of the subject matter. • Contributions in Science, a miscellaneous series of technical papers describing original research in the life and earth sciences. • Science Bulletin, a miscellaneous series of monographs describing original research in the life and earth sciences. This series was discontinued in 1978 with the issue of Numbers 29 and 30; monographs are now published by the museum in Contributions in Science. • Science Series, long articles and collections of papers on natural history topics. Copies of the publications in these series are sold through the Museum Book Shop. A catalog is available on request. The museum also publishes Technical Reports, a miscellaneous series containing information relative to scholarly inquiry and collect...
Abstract The entire catalogued paleontological collection of the New Mexico Museum of Natural History and Science (NMMNH), including 35,902+ fossils from New Mexico, is now online and searchable by the general public, avocational paleontologist, researcher, and geoscience educator. The Web site does not include sensitive geographic localities, but all aspects of the taxonomy, stratigraphy, and chronology of the specimens are viewable at http://164.64. 119.14/nmmnh/web/default. html.
Terrestrial vertebrate remains were recovered from sediments that lie on remnants of the lowest marine wave-cut platform between Point Buchon and Point San Luis. Uranium series ages of these samples, which range from 83 to about 49 ka suggest a correlation to late Pleistocene climatic and eustatic events associated with marine oxygen isotope substage Sa, and establish a maximum age of "'" 80 ka for the occurrence of terrestrial mammal fossils. The Point San Luis area assemblage appears typical of the late Pleistocene regional vertebrate paleo fauna from west-central California. Five mammalian taxa are added to the Pleis tocene record from San Luis Obispo County. Equus sp. cf. E. occidentalis, Ca mclops sp. cf. C. hesternus, and Bison antiquus were recovered from the Point San Luis area, and A1ammut americanum and B. latiffons from near Morro Bay and the Carrizo Plains in eastern San Luis Obispo County.
Paleontologic localities of significant scientific value occur on public lands in California. Some localities private land on are administered by the Bureau of Land Management (BLM) and the United States Forest Service (USFS) for their mineral resources. There is now an-opportunity to protect non-renewable palaeontologic resources through establishment of Areas of Critical Environmental Concern or through cooperative agreements with private institutions and other public agencies prior to these resources being lost, vandalized or developed for on-paleontological purposes. As a positive example, cooperation between governmental agencies and private institutions has resulted in the preservation and appropriate curation of palaeontologic resources from public land' These localities are important not only because they contain significant palaeontologic resources, but because they are field repositories offering insight to past community dynamics and structural activities of the crust, and offer data regarding rates and amounts of fault offset which directly effects the health and safety of California residents. The key to management of paleontological resources are (1) inventory, 2) cyclic prospecting for protection, and (3) curation of specimens into retrievable institutional storage to allow research and reporting.
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Plans to display Dinosaur Trackways in Washington D.C.
When asked to comment on dinosaur discoveries in the United States most experts may cite discoveries in the Badlands of Montana or the Cleveland-Lloyd Dinosaur quarry in Utah. Certainly, it is true to say that they are many fantastic Mesozoic fossil sites in the west of the USA but the eastern part of the United States, although perhaps a little under-represented in terms of fossil evidence, can still spring a few surprises.
Dinosaur Tracks
Now a new study of fossil trackways in Maryland, north-eastern USA has provided a glimpse into a thriving dinosaur based eco-system. Many of the trackways, have been found just a few miles drive out of Washington D.C. Trackways and footprints are called trace fossils. Trace fossils preserve evidence of the activity of animals such as their trackways, borings or burrows. The problem with most sets of footprints, even the very best preserved ones, is that, unless the animal is found fossilised at the end of the trackway, scientists can never be 100% certain as to the species or genus that actually left the prints.
Trace fossils such as footprints do have a significant advantage over other types of fossil such as fossil bones, most are direct in situ evidence of the environment at the time and place the organism was living.
Studying Trace Fossils
A total of over 900 fossilised footprints from a variety of dinosaurs all dated from the Cretaceous have been identified from the area. Theropods, ankylosaurs (Nodosauridae), sauropods and ornithopods are represented by the prints. Palaeontologists have estimated that the trackways were made between 121 and 98 million years ago.
Trace fossils of other animals have also been preserved in the this part of the USA, one trackway has been identified as a flying reptile, perhaps a pterosaur flew down to get a drink and its trail was preserved in the soft sediment. Mammal tracks have also been found, indeed one trackway indicates that some mammals were quite large, tracks of a quadrupedal mammal about the size of a large dog have been recorded.
Visit the Everything Dinosaur website: Everything Dinosaur.
Two Dozen Species of Dinosaur
“Based on the trace fossils, over two dozen species of dinosaurs were living in Maryland at that time,” co-author of the study, Ray Stanford commented. Ray specialises in studying fossil trackways, he began to discover tracks in the area whilst out looking for native Indian artifacts, in the stream-beds that criss cross the area.
He explained that as water and human development erode such beds, “floats” can result. These are pieces of track-bearing substrate that hydrodynamically dislodge from their natural stratigraphic context during stream bank flooding.
“This is one instance where building booms and storms can benefit science,” he said.
All of the discoveries were made either in Prince George’s county, near the capital, Washington D.C. or at the White Marsh Run area of Baltimore county.
An Illustration of the Track Made by an Ornithopod Dinosaur
Picture credit: Everything Dinosaur
Photographs show a number of footprint specimens, the peculiar, almost flower-shaped five-toed print in the foreground was most probably made by a nodosaur. Nodosaurs are members of the Ankylosauria, heavily built, slow-moving, plant-eaters with body armour and horns.
To read an early article about dinosaur tracks discovered on the North Yorkshire coast: Dinosaur Tracks Found by Young Boy.
Ray Stanford in conjunction with a Johns Hopkins University palaeontologist called Davide Weishampel hope to publish a journal paper on this new genus of nodosaur. The nodosaur print in the foreground is much smaller than the cast print in the very centre of the image (the print which the model nodosaur is facing), this indicates that some of the trackways may have been made by young, immature animals. This area may have provided a Cretaceous nursery for many species, a popular nesting and breeding ground for a variety of dinosaurs.
Providing an Insight into Dinosaur Behaviour
The scientists state that they may even have uncovered trackway evidence showing youngsters following adults, a possible insight into animal’s behavioural and social relationships.
So far, Stanford has described and published Maryland’s first dinosaur track species (called an ichnospecies which translates to ‘trace species’). It consists of both front and back footprints of a hypsilophodontid dinosaur. He named the new dinosaur footprint type or species Hypsiloichnus marylandicus, meaning “trace of a hypsilophodontid dinosaur from Maryland.”
An overview of these, and other, finds was recently published in the journal Ichnos.
Analysis of the region’s geology indicates that during that dinosaur era, fresh water sources and plant life would have been plentiful. Stanford has excavated fossilised pollen for ancient plants, along with fossilised wood for a large, now-extinct fern tree similar to today’s cycads.
The Smithsonian Museum of Natural History in Washington D.C. is investigating the possibility of putting some of the tracks on display in a special exhibition. There are certain obstacles to overcome, such as how best to present the casts so that their fine detail can be seen, but such an exhibit be popular with museum visitors. After all, it would give the residents of Washington D.C. an opportunity to learn more about some of the previous residents in the neighbourhood.
Everything Dinosaur stocks a wide selection of dinosaur models including replicas of ornithopods and nodosaurids: Everything Dinosaur Models and Replicas.
Earthquake shakes the Country – Epicentre 4km north of Market Rasen, Lincolnshire
At shortly before 1am this morning (GMT) an earthquake with a magnitude of 5.2 struck the United Kingdom. The epicentre (the point on the Earth’s surface directly above the centre of the earthquake), was 4 kilometres north of the town of Market Rasen, Lincolnshire. Reports have been received of a Market Rasen earthquake!
There is one report of an injury, the British Geological Society (BGS) had by 7am received over 1,400 reports from members of the public, the media and the emergency services. Some structural damage has been caused, chimneys falling off, walls collapsing close to the epicentral area, but this tremor was felt across a large part of the UK. Many residents in English and Wales towns were awoken by the shaking, the quake has been felt as far away as southern Scotland.
Market Rasen Earthquake
In this country we are not immune from earthquakes, each year the BGS records around 200, but only about 10% are big enough to be felt by local residents. Fortunately, most of the quakes have epicentres which are offshore. The largest earthquake recorded in the British Isles took place in 1931. This quake had a local magnitude of 6.1, but fortunately it was centred on the Dogger Bank area of the North Sea. Even so, the quake and the aftershocks were powerful enough to cause structural damage to many buildings on the east coast of England.
Finding the Epicentre
The precise epicentre of the Market Rasen quake has been calculated to be latitude 53.419 degrees north and longitude 0.354 degrees west. It is understood to have taken place approximately 5,000 metres underground.
Earthquakes are monitored by the British Geological Survey, part of the Natural Environment Research Council (NERC). There is a network of 146 seismometer stations across the UK sending data to the head office based in Edinburgh four times per day. However, during times of earthquake activity data can be sent on demand and staff at the BGS can access data and analyse results from home. They are on call 24-hours a day, as scientists don’t know when a quake will strike.
Earthquakes of this magnitude occur approximately ever 30 years or so, in the world there are about 1,300 quakes of this magnitude or bigger each year. This latest quake is the biggest since 1984, when on the 19th July North Wales was struck by an earthquake that had a magnitude of 5.4. It too caused structural damage to many buildings with cities such as Liverpool 120 kilometres from the epicentre being affected.
106 Tremors
None of the team members at Everything Dinosaur felt the quake (all sound asleep in our beds). However, one member of staff recalled the Manchester earthquakes that struck in the Autumn of 2002. A series of tremors were recorded with an epicentre in and around Manchester over a period of five weeks. The magnitude ranged from 1.1 to 3.9 ML (local magnitude). In total 106 tremors were recorded, the biggest of which (3.9 ML) hit on October 21st. Our colleague remembers particular incident very well, as he was travelling in a lift in an office block in the centre of Manchester at the time – very scary.
To read more about the work of the BGS and the latest on this mornings quake you can visit the BGS website.
Visit Everything Dinosaur’s award-winning website: Everything Dinosaur.
Rises in Oxygen Levels may Explain “Cambrian Explosion”
A new study from a multi-national team of scientists provides evidence of the link between the explosion of early life forms and the oxidation of the deep oceans. The rise of oxygen levels within the ocean between 635 and 551 million years ago may have helped trigger the increase and rapid diversification of early lifeforms, leading ultimately to the “Cambrian Explosion”.
The “Cambrian Explosion” is a term used by scientists to describe the huge increase in life that occurred around 545 million years ago, at this stage of the history of life on Earth, all life was associated with marine environments. It was during the Cambrian that most of the major groups of animals that exist today evolved.
Speedy Evolution
Soft bodied animals and the stromatolites (colonies of bacteria) were partly replaced and superseded by the evolution of organisms with hard parts such as exoskeletons and shells. The first forms of life that could be biomineralised evolved, this meant that the hard parts of their bodies could be preserved as fossils and thus this period of ancient history not only marks the increasing abundance and diversity of organisms but also marks the start of an enriched fossil record, providing palaeonotologists with more evolutionary evidence.
Complex organisms had been in existence prior to the beginning of the Palaeozoic, but the fossil record is extremely poor. Multi-cellular life forms have been recorded in rocks of approximately 600 million years of age, but these creatures seemed to have lacked any hard parts and as soft-bodied creatures, palaeontologists have only a few tantalising fossils to work with.
A Rise in Oxygen Levels
The rise in oxygen levels and the oxidation of deep oceans in the late Precambrian has been accepted for a number of years. However, it had been thought that the increase in photosynthetic bacteria such as cyanobacteria (formerly known as blue-green algae), assisted by other non-biological means such as the breakdown of water into hydrogen and oxygen by ultraviolet rays penetrating to the surface of the Earth through the ozone devoid atmosphere had led to the increase.
Now, new research from scientists studying the geochemical structure of the Duoshantuo Formation in southern China reveals that life on Earth may have been influenced by two distinct pulses of oxygen. The first increase in oxygen predates the “Cambrian Explosion” by a significant amount of time but may have led to an increase in microscopic life forms. The second burst of oxygen aerating the oceans seemed to have occurred around 550 million years ago and in geological terms immediately pre-dates the increase in life during the Early Cambrian.
Trilobites Thrived During the Cambrian
Picture credit: Everything Dinosaur
An international team of scientists from Virginia Tech, the University of Maryland, University of Nevada (Las Vegas) and the Chinese Academy of Sciences set out to test the relationship between the evolution of more complex and diverse life forms and environmental change. To do this the team needed to find sedimentary strata that pre-dates the Cambrian and a sequence of strata (stratigraphic column) that would show deposition and formation as a timeline, one that had not been altered or changed by other chemical or geological processes.
Finding pristine Precambrian strata is a challenge in itself but such locations are known, one being the Doushantuo Formation in the Yangtze Gorges area, Guizhou Province, southern China. The strata consists of phosphate and dolomite sequences, laid down at the bottom of a sea. China at this time was made up of two separate and submerged continental sheets, that lay in shallow, warm tropical waters off the coast of the super-continent Gondwana. The first part of what was to become China, closest to Gondwana, straddled the Equator, the second part lay across the Tropic of Cancer.
Mapping Oxygen Levels
By mapping the levels of oxygen at various levels in the stratigraphic column, the team could measure the amount of oxygen in the marine environment and then associate this with the biostratigraphic column (fossils used to date and correlate strata), this would provide evidence to support the increase in oxygen leading to a diversity and increase in lifeforms.
To calculate when there was enough oxygen to support animal life in the ocean, the researchers asked, “What kind of geochemical evidence would there be in the rock record?” said Shuhai Xiao, associate professor of geosciences at Virginia Tech.
Scientists hypothesized that there was a lot of dissolved organic carbon in the ocean when oxygen levels were low. If oxygen levels rose, some of this organic carbon would be oxidized into inorganic forms, some of which can be preserved as calcium carbonate (CaCO3 ) in the rock record. “We measured the carbon isotope signatures of organic and inorganic carbon in the ancient rocks to infer oxidation events,” said co-author Ganqing Jiang, assistant professor of geology at the University of Nevada at Las Vegas.
The stratigraphic column exposed during the construction of the dams in the Yangtze Gorges area represents a large slice of ancient geological history. The researchers carefully took samples from each strata of rock, the deeper the strata, then, unless the strata has been overturned, as can sometimes happen during mountain building processes for example, the older the rocks will be. This is an important geological principle it is called the “Law of Superposition”. Many hundreds of different samples were taken, representing marine deposits laid down during the Precambrian and Early Cambrian.
The researchers cleaned and crushed the small samples to powder, which they reacted with acid to release carbon dioxide from carbonate minerals, and then burned the residue to get carbon dioxide from organic matter. “The carbon dioxide that is released was measured with mass spectrometers to gives us the isotopic signature of the carbonate and organic carbon that was present in the rock,” a researcher commented.
“The relative abundances of the carbon-12 and carbon-13 isotopes, which are stable and do not decay with time, provide a snapshot of the environmental processes taking place in the ocean at the different times recorded in the layers of rock”.
The stratigraphic pattern of carbon isotope abundances suggested to these researchers that the ocean, which largely lacked oxygen before animals arrived on the scene, was aerated by two discrete pulses of oxygen.
The first pulse that occured in the Precambrian seemed to have little impact on a large organic carbon reservoir in the deep ocean, but did spark changes in microscopic life. The second event, which occurred around 550 million years ago, immediately prior to the palaeontological event known as the “Cambrian Explosion”, resulted in the reduction of the organic carbon reservoir. This indicates that the ocean became fully oxidized just before the evolution and diversification of many of Earth’s earliest animals. Perhaps this dramatic increase in the level of available oxygen provided the fuel for the rapid burst of evolution.
Certainly, scientists have speculated why all of our sudden around 545 million years ago evolution seems to have pressed the accelerator when for much of the Precambrian (Cryptozoic), evolution seemed to be progressing at a very slow pace. You could say that evolution, prior to the second pulse of oxygen had progressed at a snail’s pace but to be fair to the Gastropods (snails) these animals did not really get going until the Early Cambrian.
Photographs show a field of view 0.15 millimetres in diameter of a beautifully preserved eukaryotes fossil from the Doushantuo formation (635-550 million years old). Eukaryotes are cells with their genetic material enclosed in a cell nucleus. Eukaryotes are believed to have first appeared in the fossil from strata dated to 2,100 million years ago, but evidence from molecular biology indicates that they may have been present earlier than this but left little or no fossil evidence.
“The Doushantuo Formation has a wonderful fossil record. It allows us to look at major fossil groups, when they appear and when they disappear, and to see a relationship between oxidation events and biological groups”, a researcher commented.
“This study supports the growing view that life and environment co-evolved through this tumultuous period of Earth history,” said geochemist Alan J. Kaufman, a co-author of the study from the University of Maryland.
The triggers for the oxidation events remain elusive, scientists are still not sure what set off these oxidizing events. Members of the research team have suggested that these two events recorded in marine sediments were probably related to oxygen in the atmosphere reacting with sediments on land as rocks are eroded away. The lack of biological activity on the land would have resulted in weathered rocks and soils on the continents releasing certain dissolved ions, such as sulphate, into rivers. These would then be transported to the sea where they might be used by bacteria to oxidize the organic carbon pool in the deep oceans.
This article has been adapted from materials published by Virginia Tech, USA. The full article entitled “Pulsed oxidation and biological evolution in the Ediacaran Doushantuo Formation,” was written by Kathleen A. McFadden; Jing Huang and Xuelei Chu of the Institute of Geology and Geophysics, Chinese Academy of Sciences; Ganqing Jiang; Alan J. Kaufman; Chuanming Zhou and Xunlai Yuan of the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences; and Shuhai Xiao.
It is due to be published in March.
CollectA have recently introduced a range of models of invertebrates reflecting iconic animals from the fossil record including trilobites and members of the Mollusca: Replicas of Iconic Fossil Animals, Models, Toys and Games.
Yorkshire Lad goes “Walking with Dinosaurs” (Dinosaur Footprints Discovered)
For many palaeontologists discovering a perfectly preserved set of dinosaur footprints may mark a high point in their careers but for young Rhys Nichols of Scarborough finding dinosaur trackways is as easy as taking a walk along the beach.
Whilst walking with his father, on the beach at Burniston Rocks, north of the seaside resort of Scarborough on the North Yorkshire coast, Rhys noticed that part of cliff face had fallen away and it was here that he found the footprints. Rhys’ very proud father, Richard stated that “Rhys loves dinosaurs so for him to find something like that was wonderful. He was over the moon – I couldn’t get him away from it!
Dinosaur Footprints
“We are always coming down here beach-combing and hunting for fossils.”
Experts have agreed that these fossil prints are a “great find”. The footprints are what is known as a trace fossil. Trace fossils preserve evidence of the activity of animals, such as their tracks; unlike many dinosaur fossil bones that may have been transported after death a long way from where the dinosaur originally lived, most trace fossils such as these prints are direct “in situ” evidence of the environment at the time and the place where the dinosaurs roamed.
Picture credit: Everything Dinosaur
Pictures show two beautifully preserved fossil footprints with the three toes of dinosaur seen clearly. The raised appearance of the fossil is typical of this sort of trace fossil. Footprint fossils can either be a depression-type fossil made by the weight of the animal or a cast of the “hole” made by the animal’s foot as it walked along. Sediments can fill the print up and it is these that are fossilised and the cast preserved giving the raised appearance. Mr Nichols measured the footprints estimating that they were around 21 cm in diameter, other footprints have been found but they had been heavily eroded.
Fossilised Trackways
A number of fossilised dinosaur trackways have been found in the North Yorkshire area , much of the coast of the North East from Scarborough to near Redcar is comprised of exposed areas of delta mudstone and sandstone and thin coals that were laid down in the Middle Jurassic approximately 160 million years ago. The rock strata where the prints have been found is well known for producing dinosaur trackways and isolated footprints, in fact geologists term this strata as the Burniston Footprint Bed. As blocks of silty sandstone fall onto the beach, split apart from the cliff face by erosion, these blocks frequently come to rest at the base of the cliff upside down revealing the finely detailed tracks of dinosaurs from the Jurassic period.
It is not known what actual species of dinosaur made the prints, as with most fossil trackways, unless the culprit is found fossilised at the end of the trackway, Ichnologists (scientists who specialise in studying tracks), can only speculate what sort of creature it was. Local museum staff have stated that this dinosaur may have been an Iguanodon. The three-toed prints are indicative of an Iguanodontidae, however, the mid Jurassic date is very early for such an animal, more normally associated with the Early Cretaceous. Perhaps it could be a trackway made by a dryosaur, these animals grew to lengths in excess of 3 metres, were relatively light and had a bi-pedal stance. Dryosaurs seem to have been relatively ubiquitous, with fossil being found in both the Northern and Southern Hemispheres. Like many trackways the scientists may just have to resort to referring to these wonderfully, well preserved prints as belonging to an “in-determinant ornithopod”.
Three Toes Clearly Observed
Photographs show a close up of the footprint shown in the foreground in the picture with Rhys. The animal would have been walking from right to left as the page is viewed. The three-toes can clearly be seen, but there is little evidence of a claw mark, adding weight to the thought that this is the footprint of an ornithopod. Based on comparisons with the fossils of ornithopods such as the large amount of Iguanodontidae material available, it has been estimated that the dinosaur walking across the delta 160 million years ago would have been roughly the same size as young Rhys. It is not known whether the animal was a fully grown adult or juvenile. The pictures of the footprints indicate a bipedal stance, but as to what actual animal made these tracks, this will probably remain a mystery, unless of course Rhys happens to find another set of prints whilst beach-combing but this time with the fossils of the dinosaur which made them at the end of the track.
Here’s hoping… in the meantime well done to Rhys, palaeontology remains the only science where by going for a walk you can change the way the world views itself.
Everything Dinosaur stocks a wide range of ornithopod models and figures: Everything Dinosaur – Dinosaur Models and Figures.
Giant Frog challenges Scientists over movement of Continents
Frogs are the most common type of amphibian alive today, with an estimated 5,500 separate species, making them the most diverse and successful clade of the Lissamphibians. They are known from all the continents except Antarctica but their fossil record is quite poor. Although very much extant, scientists still debate how many actual families make up the order containing frogs and toads – Anura. With discussion ongoing as to how to classify frogs and toads around today, it is no wonder that difficulties arise when trying to piece together the development and relationships between elements of Anura when you consider how sparse the fossil evidence is.
Now the discovery of a giant, Late Cretaceous frog from Madagascar that may be related to the horned frogs of South America, has opened up the debate once again over frog family ancestry and the break up of the super-continent Gondwanaland.
Frog and Toad Evolution
Frogs and toads are very specialised Lissamphibians with a body shape (morphology) unlike their living relatives and their ancient amphibian ancestors from the Palaeozoic. In comparison with other amphibian groups, they have dramatically reduced skeletons, lacking ribs, a tail, with a simple pelvic girdle and relatively few vertebrae. One of the earliest known frogs was also found in Madagascar, called Triadobatrachus; this animal dates back to the Triassic.
Frogs and toads were probably relatively abundant during the Mesozoic but the lack of fossil evidence inhibits palaeontologists when it comes to working out Anura evolution. Fossil bones have been recorded from a number of Mesozoic sites but they are usually isolated fragments, ilia, humeri (limb bones) and the more robust skull elements such as the frontoparietals and squamosals – elements from the top and towards the rear of the skull respectively. Some upper jaws bones (maxillae) have also been located and it is the jaws and the partial skull elements that provide the greatest assistance to palaeontologists when they attempt to work out the relationships between extinct genera and species.
Researchers Study Fossil Bones
Researchers from New York’s Stony Brook University aided by a team from University College, London headed up by vertebrate morphologist and palaeontologist Susan Evans; have published their findings on this new species of Madagascan giant frog in the journal Proceedings of the National Academy of Sciences.
This discovery, led by David Krause of Stony Brook University may undermine current scientific thinking over the isolation of Madagascar that was believed to have taken place in the Cretaceous. Conventional theory states that by approximately 95 million years ago, the land mass that was to eventually form India and Madagascar had split away from Africa, part of the break up of Gondwanaland (Australia, New Zealand, Africa, Antarctica and South America). Over the next 30 million years or so a rising plume of hot magma forced its way through a fault rifting apart India and Madagascar. Madagascar was left an isolated island with its own distinct indigenous fauna and flora and India went northwards to collide with Asia.
Beelzebufo ampinga
The fossilised bones of Beelzebufo ampinga, a frog the size of a partially deflated beach ball and tipping the scales at around 4 kilogrammes, making it the largest frog found to date, resemble the bones of the extant frog group – the Ceratophyrinae. The Ceratophyrinae, termed the “horned frogs” as many members of this family have soft extensions of skin growing out from the upper eyelid, which resemble little horns; are associated with South America.
Study of the fossils have indicated that this ancient animal is not related to any of the frog species living on Madagascar today. If Madagascar was very much an isolated island when Beelzebufo was hopping around, then how do scientists explain a member of the Ceratophyrinae group on an island thousands of miles away from their ancestral home of the South Americas.
Beelzebufo – the Frog from Hell
Picture credit: Associated Press
The diagram above shows an artist’s impression of Beelzebufo, with a modern frog and a pencil for scale. Although only partial elements of the skeleton have been recovered Krause and his team estimate that this animal was 40 cm long and would have weighed as much as a large domestic cat.
The most characteristic feature of the Ceratophyrinae is not their horned eyelids (some members of the group do not possess this feature), but their large heads, huge mouths and blunt snouts. They are voracious and unfussy hunters, lying half submerged in mud waiting for any unsuspecting small animal to wander by. Basically, anything that can fit into their mouths is on the menu, mice, frogs, snakes, fish and such like.
Wide Mouth and Powerful Jaws
Beelzebufo had a very wide mouth and powerful jaws, plus teeth. The skull material recovered has ridges and groves on it; perhaps indicating that this animal had bony armour or a protective head shield.
David Krause commented: “This frog, if it has the same habits as its living relatives in South America, was quite voracious. It’s even conceivable that it could have taken down some hatchling dinosaurs.”
The name Beelzebufo is a derivative of the Greek word for Devil and bufo is the Latin for toad. The “Devil Toad” would be an apt title for a frog capable of swallowing whole baby dinosaurs.
Krause and his team began finding fragments of abnormally large frog bones whilst studying the late Cretaceous sediments of the Mahajganga basin in north-western Madagascar in 1993. Amongst the various dinosaur and crocodilian fossils a total of 60 fossil frog bone fragments were located during a number of expeditions to the area by the New York team.
A Relative of Extant Horned Frogs
The unusually large frog bones were sent to the University College, London for specialist Susan Evans to examine. The London researchers were not able to piece together a complete skeleton but they had enough of the skull elements to make a diagnosis and interpret Beelzebufo as a relative of the horned frogs group.
The giveaway clinching evidence was the skull material indicating a “short, fat skull with a huge mouth”, says Evans.
Scale Drawing of Beelzebufo Skeleton compared to Living Frogs
Picture credit: Journal of Proceedings of the National Academy of Sciences
The drawing depicts the skeleton of Beelzebufo ampinga (A) compared to the largest extant member of the South American Ceratophyrs (B) and the largest frog species found on Madagascar today (C). The skeletal material in white represents bones found, those parts of Beelzebufo skeleton in grey are a scientific impression as to what the remainder of the skeleton would have looked like.
A Palaeontology Puzzle
The link to South America raises a palaeontology puzzle. Standard theory for how the continents drifted apart show what is now Madagascar would have been long separated by ocean from the Americas during Beelzebufo’s time. Frogs with their soft permeable skins cannot survive long in salt water, so reaching Madagascar by swimming can be ruled out.
Krause contends that the giant frog provides evidence for competing theories that some bridge still connected the land masses that late in time, perhaps via Antarctica that was much warmer than today. Perhaps Gondwanaland stayed together for longer than scientists currently think, could India/Madagascar have been linked to South America by an Antarctica land bridge as recently as the late Cretaceous.
Evans says that when she first began to suspect the Madagascar fragments came from a frog related to South American Ceratophryinae, she was very cautious about the claim. “We knew it would be controversial,” she says. “There are people who believe everything on Madagascar today must have been there when it broke with Gondwanaland 160 million years ago.”
Blair Hedges, a biologist at Pennsylvania State University in University Park, agrees that Beelzebufo is an important find. “The new fossil frog, besides being large and odd-shaped, is quite unexpected because of its apparent relationship with South American species,” he says.
But he says he isn’t yet convinced that the new find is related to the South American frogs. Molecular clock data suggests that these frogs split from a common ancestor more recently than 66 million years ago, he says. “Based on molecular evidence of frog relationships, the specific resemblance to some living wide-mouthed frogs is more likely from [evolutionary] convergence than actual relationship.” Convergent evolution, where unrelated species occupying similar niches tend to look the same, is common in frogs, he says.
Even if they are related, he adds, this doesn’t mean that the frogs necessarily had to walk on land from one location to another before Gondwana split. “Any organism, including a frog, can raft on dead vegetation,” he says.
Flood events and tropical storms can wash relatively large pieces of vegetation out to sea, some of these “rafts” get washed up on foreign shores. A number of animals migrate between islands today under these circumstances.
Susan Evans and her team remain convinced that Beelzebufo is a relative of the horned frogs of the Americas and refutes the convergence evolution theory. “It is the same family. I have no doubt of that,” she says.
It seems that this large, voracious frog – a trouble maker back in the Mesozoic, eating anything that could fit in its mouth, is going to be causing just as much trouble in scientific circles here in the Holocene.
Visti Everything Dinosaur’s website: Everything Dinosaur.
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The Basal Nodosaurid Ankylosaur Europelta carbonensis n. gen., n. sp. from the Lower Cretaceous (Lower Albian) Escucha Formation of Northeastern Spain
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[
"Vertebrae",
"Skull",
"Teeth",
"Dinosaurs",
"Cretaceous period",
"Ribs",
"Spine",
"Ischium"
] | null |
[
"Mark A. Loewen",
"Eduardo Espílez",
"Luis Mampel",
"Jelle P. Wiersma",
"James I. Kirkland",
"Luis Alcalá"
] | null |
Nodosaurids are poorly known from the Lower Cretaceous of Europe. Two associated ankylosaur skeletons excavated from the lower Albian carbonaceous member of the Escucha Formation near Ariño in northeastern Teruel, Spain reveal nearly all the diagnostic recognized character that define nodosaurid ankylosaurs. These new specimens comprise a new genus and species of nodosaurid ankylosaur and represent the single most complete taxon of ankylosaur from the Cretaceous of Europe. These two specimens were examined and compared to all other known ankylosaurs. Comparisons of these specimens document that Europelta carbonensis n. gen., n. sp. is a nodosaur and is the sister taxon to the Late Cretaceous nodosaurids Anoplosaurus, Hungarosaurus, and Struthiosaurus, defining a monophyletic clade of European nodosaurids– the Struthiosaurinae.
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https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0080405
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Description and Comparisons
Skull
The skull (AR-1-544/10) was lying on its dorsal surface and is moderately well preserved although distorted through compaction (Fig. 7). The palate is crushed in toward the skull roof, resulting in the medial rotation of both maxillae with the posterior teeth displaced into the posterior palate. The sheet-like palatal bones are highly fragmented. The braincase is crushed along the plane of the cranial nerve openings and the fenestra ovalis completely obscures them. Unexpectedly, the right quadrate (Fig. 8 H–J) and associated portion of the palate was dislodged from the skull and subsequently crushed across the ventral side of the basicranium. This gives the impression that these bones had been expelled from inside the skull prior to compaction. Both the left and right nasals were separated from the skull and the premaxillae (whereas possibly present upon discovery) have not been identified.
The skull has a minimum length of 370.3 mm from the anterior end of the maxillae to the rear margin of the squamosals. The skull has a maximum width of 299.1 mm at the orbits and narrows to 203.7 mm at the posterior end of the skull at the squamosals, giving the skull the “pear-shaped” dorsal profile characteristic of derived nodosaurids [70], [71]. Although tapering posteriorly, there is no distinct post-temporal notch as in polacanthids and other nodosaurids [63].
The maxillae (Fig. 7 D–F) are irregularly sculptured externally with a flattened, horizontally oriented buccal recesses that are inset approximately 2 cm. The anterior margin of the maxilla appears to form the posterior margin of a relatively simple naris relative to derived nodosaurids and ankylosaurids. Medially, there is no evidence that the maxilla formed a portion of a secondary palate. The tooth row was arched ventrally with an estimated 22–25 alveoli increasing in size posteriorly as in Edmontonia [72]. In ventral orientation, the tooth rows are only moderately deflected medially, such that the palate would not have had a pronounced hourglass appearance typical of derived nodosaurs such as Pawpawsaurus, Edmontonia, and Panoplosaurus [73]–[75]. However, it is not dissimilar from that of the primitive nodosaurid Silvisaurus [76], [77].
The nasals (AR-1-133/10, AR-1-639/10) are relatively large and subrectangular, tapering somewhat anteriorly (Fig. 8 A–D). Both nasals extend laterally from their relatively straight, unfused midline suture before flexing down to a sutural contact with the maxillae that extends for most of their length. When rearticulated onto the skull, they appear to fit well, despite the skull's distortion. Most ankylosaurs have fused nasals except the nodosaurids Silvisaurus [76], [77] and Niobrarasaurus [78], although the nasals are unknown in European nodosaurids [24], [32], [33]. A distinct tongue-like process projects from the nasal's posterior margin and would have overlapped the frontals. The external surface is lightly textured and the internal surface is relatively smooth, suggesting the narial passage was large and simple, rather than convolute as in derived nodosaurids and ankylosaurids [79], [80].
The orbits are somewhat crushed and the sutures of the bones surrounding them are obscured by fusion. The orbits are subrectangular in shape, are slightly more elongate anteoposteriorly and are directed anterolaterally. The prominent and evenly rounded suborbital horn is formed mostly from the quadratojugal posterior to the ventral margin of the orbit, as in most derived ankylosaurs [81], [82] and unlike that in polacanthids such as Mymoorapelta, Gargoyleosaurus, and Gastonia where the suborbital horn is below the orbit and is formed exclusively by the jugal [83]–[85]. The suborbital horn appears to be unornamented and hides the head of the quadrate in lateral view.
The lateral wall of the skull extends posteriorly behind orbit with a dorsoventally wide posterior notch, such that the lower temporal opening is just visible in lateral view. There is no lateral wall of skull behind the orbits in polacanthids [70], [81] and most nodosaurids other than Peloroplites [86], Silvisaurus [76], Struthiosaurus transylvanicus [22], [23] and one specimen from the Dinosaur Park Formation assigned to Edmontonia (ROM 1215) [88], although in these taxa the lower temporal opening is still visible in lateral view as in Europelta. The lower temporal opening is completely obscured in lateral view in Cedarpelta [84], [86], Shamosaurus [89]–[91], Gobisaurus, [92] Zhongyuansaurus [93] and all derived ankylosaurids.
Although the palate is fragmented and crushed along the internal surface of the skull roof, the fragments of the vomer suggest it did not extend ventrally to the level of the tooth row. Additionally, the broad sheet-like pterygoids appear to have been flexed nearly dorsally against the anterior portion of the basicranium as in nodosaurids and not like the open transversely oriented pterygoids characteristic of ankylosaurids or polacanthids [94].
The posterolateral margin of the pterygoid is fully fused to the quadrate. There is a sutural contact between the straight, nearly vertical quadrates and the quadratojugal laterally. The quadrates are wide transversely and thin rostrocaudally as compared to the mediolaterally narrower quadrates of other ankylosaurs [82]. The contact with the squamosal is also transversely wide, unlike the narrow, rounded contact seen in many ankylosaurs such as Mymoorapelta (Kirkland, pers. obs.) and Cedarpelta [63], [86]. The mandibular articulation is proportionally wider than in any other ankylosaur examined as a part of this study and the medial condyle larger than the lateral condyle. The ratio of mediolateral quadrate width to dorsoventral quadrate length is 0.77 (94 mm/122 mm). The anteropostior length of the quadrate condyle is 31 mm. There is no fusion between the quadrates and the paroccipital processes.
Vertical compaction has obscured the posterior view of the skull, in particular the foramen magnum and the supraoccipital. However, even with compaction it is apparent that in occipital view the skull was subrectangular and wider than tall as in Gargoyleosaurus, Gastonia, and most other derived anklylosaurs, and unlike the narrow, highly arched occipital region of Struthiosaurus [22]. The paroccipital processes extend horizontally lateral to the foramen magnum and then flare dorsoventrally by approximately 100% of their minimum widths. They angle posteriorly at about 30 degrees when viewed ventrally (Fig. 7 F). In morphology and orientation, they are most similar to those in Gargoyleosaurus [95] although ventral twisting is not present. In most other ankylosaurs, the paroccipital processes extend straight laterally [81], [96] or may be flexed ventrally as in Gastonia [83]. A triangular wedge of bone of unknown identity is fused to the anterior ventrolateral margin of the paroccipital, separating it from the quadrate.
The subspherical occipital condyle (Fig. 7 B, F) has a width of 59.4 mm and height of 46.5 mm and lacks a distinct neck to separate it from the rest of the basicranium. Although no cranial sutures are visible, the occipital condyle does appear to be composed exclusively of the basioccipital. It is similar in overall morphology to that of the basal ankylosaurid Cedarpelta [88] except that the occipital condyle angles somewhat ventrally, but not as much as in more derived nodosaurids [71], [82]. The ventral surface of the relatively elongate basioccipital is broadly convex. Again, as in Cedarpelta [88], there are no distinct, separate basal tubera between the basioccipital and the short basisphenoid, but instead there is a prominent transverse flange extending across the ventral surface of the basicranium along the line of this suture. The pterygoid processes appear to be short, but are completely obscured by crushed pterygoids bone fragments that wall off the anterior part of the braincase as in most nodosaurids.
The skull roof (figs. 7 C, 9 A) is roughened texturally by remodeling of the bone surface as in Cedarpelta, the nodosaurids Sauropelta and Peloroplites, and the shamosaurine-grade ankylosaurids Shamosaurus and Gobisaurus [81], [86], [88]. Europelta differs from these specimens in that some of the margins of the scale impressions on the skull roof are visible, as seen in Edmontonia, Panoplosaurus and Struthiosaurus [22], [77]. These scale margins are represented by shallow grooves that are difficult to see relative to the textured surface of the skull and the cracks in the bone due to compaction. These grooves are particularly evident along the lateral margins of the skull roof above the orbit. An extensive median scale appears to have covered much of the central portion of the skull between and posterior to the orbits on the frontals and parietals as other nodosaurids [63], [82]. There does not appear to be any distinct nuchal ornamentation. The skull is thickened above the orbit, but there is not a distinct supraorbital boss, a condition similar to Peloroplites, Cedarpelta, Shamosaurus, and Gobisaurus [86], [88]–[90], [92]. Narrow grooves along the margin of the skull in this area above the orbits suggest that a particularly robust pair of scales were present in this area as indicated by a deep groove bisecting this ornamented area directly above the orbit. Weak grooves delineate a small scale without underlying ornamentation separating the posterior supraorbital scale from the squamosal horn forming the posteriolateral margin of the skull roof. The squamosal horn is ornamented by narrow grooves radiating from its apex onto the skull roof. Grooves on the anterolateral sides of the fronto-parietal scale appear to delineate two scales between the anterior supraorbital scales. Unfortunately, no distinctive scale boundaries are recognizable on the nasals, although the dorsal surfaces of the nasals are textured. Several elongate scales rimmed the lateral raised margin around the orbit.
In dorsal view, the posterior margin of the skull is concave, whereas it is nearly straight or convex in all other nodosaurids. This reflects the posterior angulation of the paraoccipital processes and the squamosal horns. Interestingly, the occipital condyle is barely visible, though not completely obscured in dorsal view. There is no evidence of any distinct nuchal sculpturing. The skull roof is relatively flat but a slight dome may have been present prior to crushing. However, it is clear that the skull roof is not as highly domed as in many other nodosaurids, such as Struthiosaurus [22], [26].
Attempts were made to image the skull using X-ray photography and CT scanning. The abundance of pyrite present in the skull (Fig. 4E) presents a strong limitation in the use of these techniques as pyrite is opaque to X-rays.
Mandible
A small dentary fragment extending for only four complete alveolae (AR-1-133/10) was preserved from the holotype skeleton (Fig. 8 E–G). However, a robust left dentary and splenial are preserved together (AR-1-3698/31) from the paratype specimen (Fig. 10 A–E). The splenial is not in its posteriomedial position relative to the dentary, but is fused across the posterior portion of the tooth row transversely. Additionally, an isolated left angular with a distinct highly sculptured scale along its ventral margin (AR-1-2945/31), was recovered (Fig. 10 F, G).
The dentary is 184.7 mm long with a minimum of 21 tooth positions, with no possibility of more than two unpreserved alveoli as determined by the position of the suture with the angular and surangular. As with the maxillary teeth, the alveoli are more than twice as large posteriorly. There is only 1.5 cm between the anteriormost alveoli and the symphysis, suggesting that there may have been premaxillary teeth as at least nine anterior teeth would have been positioned to oppose the premaxilla. The primitive ankylosaurs Sarcolestes [34], [98], Gargoyleosaurus, [85], Silvisaurus [76], Animantarx [97], Sauropelta [99], Anoplosaurus [17], Hungarosaurus [33] and Struthiosaurus [22] have a short anterior diastema, and thus a narrow predentary, whereas this diastema is longer in ankylosaurs with wide predentaries. However, the symphysis in Europelta is robust and dorsoventrally deeper (45.0 mm deep and 29.00 mm across) than in ankylosaurs [82], and is most similar to the deep symphysis of Hungarosaurus [32], further suggesting a reduced predentary with a rudimentary ventral process. The symphysis is marked by two deep anteroposteriorly directed grooves. A row of foramina extends posteriorly on the lateral surface of the dentary from just dorsal to the buccal recess to the notch for the surangular, whereas nutritive foraminae are not clearly visible ventral to the alveolae on the medial side of the dentary as in other ankylosaurs. The recessed tooth row is deflected medially and forms a convex arch in lateral view. The dentary of Hungarosaurus is deeper dorsoventrally than that of Europelta [33].
The splenial (Fig. 10 A-D) is a thin bone with a convex ventral margin 156.6 mm long that contacts the angular. It has the appearance of an obtuse triangle in medial view. There is large, well-developed intermandibular foramen (7 mm long and 5.3 mm wide) 50 mm from its anterior end.
The angular (Fig. 10 F, G) has a maximum length of 175 mm. The lateral margin is highly rugose, because the bone is textured and remodeled to support a large scale, extending about 10–12 mm ventral to the ventral margin of the angular for most of its length. A distinct ridge marks the dorsal limit of the mandibular ornament medially, where it is in contact with the ventral margin of the splenial. Dorsal to this contact the bone is smooth. The ventral extent of the textured bone supporting the mandibular scale is similar to that observed in ankylosaurids such as Euoplocephalus [95] and Minataurasaurus [100], rather than the more lateral orientation found in Gargoyleosaurus [93] and in nodosaurids like Sauropelta [99] and Panoplosaurus [101].
Teeth
A large number of teeth are preserved from both the holotype AR-1/10 (20+) and the paratype AR-1/31 (15+) although many have drifted away from the alvaeolae. We assume that the teeth associated with the holotype pertain to the maxilla (several are preserved in the palate and in the maxilla) and those of the paratype pertain to the dentary (several are preserved in the dentary). In general, the cutting surfaces of the teeth are not well preserved, but a few exceptions exist. Wear facets were not observed on any of the teeth. The roots for both dentary and maxillary teeth are swollen lingually, are three to four times the length of the crowns, and are subquadrate in cross-section. One small tooth (AR-1-343/10) is more highly asymmetrical mesiodistally and may represent a premaxillary tooth (Fig. 11 L, M).
The isolated maxillary teeth (Fig. 11 A–K, N–FF) have a weakly developed labial cingulum and a strongly developed lingual cingulum. The best preserved right tooth AR-1-324/10 is 11.50 mm wide, 9.99 mm tall with seven to eight mesial denticles and five to six distal denticles (Fig. 11 A–E). A large right tooth AR-1-564/10 is 17.23 mm wide and 12.95 mm tall with eight to nine mesial denticles and ∼six to seven distal denticles (Fig. 11 V–Z).
The isolated dentary teeth (Fig. 11 GG–FFF) are identical to the maxillary teeth and have a weak lingual cingulum and a strongly developed labial cingulum. The best preserved tooth AR-1-3700/31 is 14.03 mm wide and 12.69 mm tall with eight to nine mesial denticles and six to seven distal denticles (Fig. 11 LL–PP). The largest dentary tooth AR-1-3650/31 is 16.58 mm wide and 13.50 mm tall (Fig. 11 GG–KK).
With their relatively large size and well-developed cingula, the teeth of Europelta are most comparable to those of other nodosaurids [72]. They similar to the teeth of Cedarpelta, Sauropelta [34], [97], [102], Edmontonia and Panoplosaurus [72], but are not as high crowned as in the Jurassic ankylosaurs Sarcolestes and Priodontognathus [103], the Jurassic polacanthids Gargoyleosaurus [93] and Mymoorapelta (Kirkland, pers. obs.), the nodosaurids Peloroplites [84] or Hungarosaurus [33]. Additionally, the large teeth of Gobisaurus are more inflated labiolingually than in Europelta and other ankylosaurs. The teeth of Gastonia and putative Polacanthus teeth are also inflated, but are smaller proportionally [83], [103]. The teeth of Europelta differ from an isolated tooth from the Cenomanian of France which is about half the size, and proportionally is longer mesiodistally with more deeply divided denticles forming ridges on the labiolingual surfaces of the tooth [104]. Likewise, lower Cenomanian teeth assigned to “Acanthopholis” have more deeply divided denticles in what is a proportionally taller tooth [17]. The teeth of Struthiosaurus languedocensis [31] from the lower Campanian of France also differ in size and in having longer, lower tooth crowns.
Axial skeleton
There are numerous ribs and vertebrae preserved from the holotype (AR-1/10) and the paratype specimen (AR-1/31). Vertebral measurements are presented in Table 1.
The complete atlas (AR-1-649/10) from the holotype has a total width of 195.6 mm (Fig. 12 A–F). The neural arch is divided dorsally with the left side fused to the centrum and the right side unattached. The anterior face of the atlantal intercentrum is 73.7 mm wide by 71.7 mm tall and its posterior face is 99.9 mm wide by 61.2 mm tall with a length of 62.0 mm. The axis is not present in either associated skeleton.
There are five post-axis cervical vertebrae (AR-1-431/10, 449, 533, 637, 650) preserved from the holotype skeleton (Fig. 12 I– II) and five from the paratype skeleton; of which four are illustrated (AR-1-3586/31, 3632, 3671, and 3676) (Fig. 13). Overall, they are typical of most other described ankylosaur cervical vertebrae. The centra are amphicoelus, wider than tall, anterorposteriorly short, and medially constricted. Anterior and mid-cervical vertebrae have the anterior faces of the centra dorsally elevated relative to the posterior faces. This is in contrast to the posterior cervical centra which have horizontally aligned faces. The ventral sides of the anterior centra are characterized by two anteroposteriorly-oriented paired fossae separated by a low keel (Figs.12, N, T, Y, EE, II, 13, F), as observed in the primitive nodosaurid Animantarx [97]. The dorsal ends of the neural spines are expanded transversely. AR-1-638/10 may either be the last cervical vertebra or the first dorsal vertebra based on the position of the parapophyses.
There are two complete cervical ribs preserved for the holotype. AR-1-450/10 is a relatively anterior cervical rib (Fig. 12 G, H) and AR-1-4452/10 is a posterior cervical rib. There is no evidence of fusion of cervical ribs to the cervical vertebrae as in the ankylosaurid Saichania [105], [106] or Ankylosaurus [107]. The cervical ribs are Y-shaped overall and much like the cervical ribs of other ankylosaurs such as Silvisaurus [76], [78], [82].
Several amphiplatan to amphicoelus dorsal vertebra are preserved: eight for the holotype AR-1/10 and nine for the paratype AR-1/31. The diapophyses originate at the level of the post-zygopophyses at the dorsal extent of the neural canal. The more anterior vertebrae have large cylindrical amphiplatan centra which lack a constricted ventral keel with circular neural canals and fused ribs (AR-1-448/10, 478, and 535). The broad transverse processes are T-shaped in cross-section and angled dorsally, unlike the laterally directed transverse processes in Polacanthus [10], [38]. Two dorsal vertebrae from the holotype appear to be pathological with the centra overgrown by about 0.5 cm of lumpy reactive bone (Figs. 14, G–K, W–BB). One of these pathologic vertebrae (AR-1-535/10) has fused ribs (Fig. 14 G–K) although the other (AR-1-430/10) does not (Fig. 14 W–BB). Two additional dorsal vertebrae (AR-1-478/10, 448) with fused ribs are not pathologic (Fig. 14 L–V). More posterior dorsal vertebrae have shorter, taller, more medially constricted centra, laterally compressed neural canals, more dorsally directed transverse processes, and lack fused ribs (AR-1-155/10, 322, and 556). The neural spines are thin and rectangular with narrowly expanded dorsal ends as in Sauropelta [99]. The neural spines are oriented dorsally as opposed to the posteriorly inclined neural spines of some other ankylosaurs such as Sauropelta [97]. None of the paratype vertebrae (AR- 1-3489/31, 3633, 3662, 3672, 3673, 3674, 3675, 3677 and 3704) have fused ribs (Fig. 15), suggesting that this character is ontogenetic because the paratype AR-1/31 represents a somewhat smaller (and presumably younger) individual than the holotype AR-1/10. More expanded neural spines are present in Shamosaurus [91].
There are a number of rib fragments preserved with AR-1/10, but there are only three (AR-1-331/10, 333, 476) relatively complete ribs (Fig. 16). As with most other ankylosaurs, the ribs are sharply arched and L-shaped in cross-section proximally in anterior ribs and broadly arched and T-shaped in cross-section proximally in more posterior ribs.
The sacrum is not preserved in AR-1/10 other than an anteriormost centrum (AR-1-154/10) of the synscacrum (Fig. 17 W, X). However, for the paratype, AR-1-3466/31, there is a largely complete but fragmented synsacrum (Fig. 17 A–V) that includes an interpreted anteriormost synsacral centrum (AR-1-3451/31), more of the anterior synsacrum composed of two dorsal centra (AR-1-3450/31), four sacral vertebrae with the sacral ribs from the left side (AR-1-3446/31), two sacral ribs from the right side (AR-1-3452/31, 3460), and one caudosacral vertebra (AR-1-3512/31). Given that at least one intermediate and one anterior fused synsacral dorsal vertebra are missing, the vertebral formula for the synsacrum would be five or more dorsosacral vertebrae, four sacral vertebrae, and one sacrocaudal vertebra. The entire synsacrum would have been over 50 cm long and measures about 44 cm across the sacral ribs. The middle section of the preserved dorsal synsacrum thins anteriorly from about 7 cm wide to about 5.5 cm wide. It then expands again anteriorly as indicated by the anteriormost centrum of the synsacrum. This differs from the sacrum of Euoplocephalus [108] and Saichania [106] in which each centrum making up the synsacrum is constricted medially. The sacrum is distinctive in being more strongly arched anteroposteriorly than other described ankylosaur sacra. The neural spines are dorsoventrally shorter than the height of the centra and are fused into a vertical sheet of bone along the length of the sacrum. The caudosacral neural spine is longer and unexpanded, transitional in form between the sacral neural spines and those of the proximal caudal vertebrae. The neural spines are broken off the anterior end of the synsacrum. The ventral side of the sacrum and anterior synsacrum is longitudinally depressed. The distal ends of the sacral ribs are expanded and the most robust medial sacral rib is about 50% taller (9.4 cm) than wide (6 cm) at its attachment with the ilium. There is no sign of expansion of the dorsal termination of the neural spine on the sacrocaudal vertebra. Additionally, the caudal rib is reduced compared to the sacral ribs.
The sacrum of Struthiosaurus languedocensis [31] is similar overall, but based on the description is not so strongly anteroposteriorly arched as in Europelta. Similarly, the sacrum of Hungarosaurus, as exhibited at the Hungarian Natural History Museum, appears to be moderately arched. The moderate angulation of the faces of the sacral centra (somewhat wedge-shaped in lateral view) in Anoplosaurus [17] indicates that a moderately arched sacram may have been present in this taxon as well. Among North American nodosaurids, we have observed only a moderate anteroposteriorly arching of the synsacrum of Silvisaurus, which appears to be restricted to the posterior part of the sacrum and two sacrocaudals. In other ankylosaurs, the downward flexure of the tail from the hips is taken up in the proximal caudal vertebrae as in Mymoorapelta [84], [109] and Euoplocephalus [70], [82].
Only three proximal caudal vertebrae (AR-1-562/10, 635, 636) are present (Fig. 18 A–F, J–O, V–AA). The proximal-most caudal vertebrae are not preserved for the holotype. The preserved vertebrae probably represent caudal vertebrae positions in the interval of about 3–7. The centra are anteroposteriorly shorter than dorsoventrally tall and somewhat wedge-shaped in anterior and posterior views. The posterior chevron facets are well developed. The neural spines are inclined posteriorly and the dorsal ends of the neural spines are only slightly expanded transversely as in Gargoyleosaurus [95] and some other ankylosaurs such as Cedarpelta [86], Edmontonia [110], Hungarosaurus [32] and Euoplocephalus [70], [82]. The neural spines are strongly expanded in most polacanthids such as Mymoorapelta [84], [109], Gastonia [83], and Polacanthus [10], and some North American nodosaurids such as Sauropelta [99], and Silvisaurus [76]. The neural spine of AR-1-562/10 is broken, erroneously giving it the appearance of being strongly inclined posteriorly. The caudal ribs (transverse processes) in Europelta originate high on the sides of the centrum and angle ventrally proximal to flexing laterally, giving them a dorsally concave profile in anterior view like Hungarosaurus, Struthiosaurus, and Peloroplites, and unlike the ventrally flexed caudal ribs of many polacanthids [10], [84], [109] and the caudal vertebra assigned to “Acanthopholis” [17] or straight caudal ribs of Gargoyleosaurus [95], Cedarpelta, Peloroplites [86], and Edmontonia [87]. The proximal caudal ribs of Hylaeosaurus differ in being swept back posteriorly [111]. The lateral terminations of the caudal ribs do not expand dorsoventrally as they do in Peloroplites [86] and Struthiosaurus, which actually appear to bifurcate [25], [26].
Additionally, there are four chevrons preserved from about the same region of the tail (AR-1-560/10, 561, 569, and 4451) of which three are illustrated (Fig. 18 G–I, P–U). The proximal chevrons are approximately as long as the neural spines as in most other ankylosaurs. They are relatively straight and expanded into teardrop shapes distally in lateral view. Unlike in many ankylosaurs, there is no fusion of proximal chevrons to their respective caudal vertebrae as in Pinacosaurus and Saichania [105], [106], Ankylosaurus [107], [112], and Edmontonia (ROM 1215) [87].
Several more distal caudal vertebrae are preserved in the paratype. The two most proximal of these (AR-1-3348/31, AR-1-3717/31) have centra of nearly equal height, width, and length, with a ventral groove, and caudal ribs shorter than the diameter of the centrum that extend laterally and angle posteriorly (Fig. 19 A–J). The chevron facets are well developed with the posterior facets more strongly developed than the anterior facets. The neural spines are not developed and the zygapophyeses only extend a short distance beyond the anterior and posterior margins of the centra. These vertebrae are interpreted to represent mid-caudal vertebrae. Two more distal mid-caudal vertebrae (AR-1-3616/31, AR-1-3716/31) are similar in morphology except that the caudal ribs are reduced to anteroposteriorly directed ridges on the lateral margins of the centra (Fig. 19 K–N). Their neural spines incline posteriorly, merging with the postzygapophyses as posterior processes extending laterally past the faces of the centra to overlie and articulate between the paired prezygapophyses of the immediatly distal vertebra. This morphology is retained in the distal caudal vertebra. More distally, as in AR-1- 2950/31, 3206, 3243, 3265, 3478, and 3615, the caudal ribs are lost and the centra become more elongate (Fig. 19 O–FF). Unlike many ankylosaurs, the faces of the centra maintain a well-rounded to heart-shaped surface distally down the caudal series [82]. For many of these vertebrae, ventrally anteroposteriorly elongated skid-shaped (inverted T) chevrons are fused to the posterior chevron facets. Fusion of distal chevrons to their respective vertebrae is widespread among ankylosaurs [84], [106], [110] although it is not present in some, such as Nodosaurus [113]. One pair of distal caudal vertebrae is fused by their mutually shared chevron (Fig. 19 GG–II) such as has been documented in Mymoorapelta [84]. The most distal four caudal vertebrae (Fig. 19 JJ–LL) and their chevrons are fused together in AR-1-3204/31 to form a tapering, terminal rod of bone at the end of the tail somewhat similar to that of Sauropelta [71].
Pectoral Girdle
Parts of the right scapulocoracoid are preserved. A portion of the distal scapular blade (AR-1-429/10) is preserved with a portion of the distal ventral margin missing with a curved section broken away. There is no evidence of any distal expansion of the scapular blade as in many nodosaurids [94].
The coracoid (AR-1-657/10) is preserved with only the most proximal portion of the scapula fused on (Fig. 20 D–H). It appears to have been sheared off just dorsal to the suture between the coracoid and the scapula, perhaps in the process of removing the overlying coal seam. The coracoid is relatively equidimensional (201.3 mm long by 186.5 mm tall) relative to the elongate coracoids characteristic of many other nodosaurids [114] such as Peleroplites [86], Texasites [77], [115], and Animantarx [97]. The medial surface is concave and the lateral surface is convex giving it a bowl-shaped appearance. The ventral margin is evenly convex as in many polacanthids and nodosaurids and there is no anteroventral process as in all ankylosaurids, including Shamosaurus [91], [94]. The articular surface of the ventrally directed glenoid is wide, bounded by a flange that extends beyond the medial surface of the coracoid.
Both xiphisternal plates are preserved (Figure 20I–L). The best preserved xiphisternal is approximately 350 mm long. They appear to be arcuate flat bones. Xiphisternal plates are only known in a few nodosaurids, but those of Europelta, whereas similar in overall shape to other nodosaurid xiphisterna, are not fenestrate or scalloped along their margins as in North American nodosaurids for which they are known [82], [87], [116].
Forelimb
Parts of both humeri are preserved. The right humerus (AR-1-655/10) is represented by the proximal end (Fig. 21 A–D). It is 249.2 mm wide with a well-developed proximal head 91.9 mm wide that extends onto the posterior side of the humerus. Distinct notches separate both the laterally directed deltopectoral crest as in nodosaurids such as Sauropelta [70], [71], [99] and the internal tuberosity from the humeral head. The deltopectoral crest extends lateraly from the humerus and is not flexed anteriorly as in polacanthids and ankylosaurids [94].
The left humerus (AR-1-327/10) is represented by a midshaft for which both the proximal and distal ends appear to have rotted off and the core of the shaft has rotted away (Fig. 21 E–H). The shaft is deeply waisted relative to the proximal and distal ends. Although relatively uninformative, enough of this humerus is preserved to indicate that the deltopectoral crest would have made up less than 50% of the length of the humerus as in nodosaurids [71], [117] and in the basal ankylosaur Mymoorapelta (Kirkland, pers. obs.) compared to the longer deltopectoral crests of ankylosaurids [70], [71]. Overall, the humerus of Europelta is similar in proportions to Niobrarasaurus [118], [119]. The wide proximal end of the humerus figured by Ősi and Prondvai [120] as cf. Struthiosaurus is similar to that of Europelta, whereas the humerus of co-occuring Hungarosaurusis is more slender proportionally.
Among the nine unguals preserved for AR-1/31, one specimen (AR-1-3711/31) may represent a manual ungual. It is more equidimensuional than the other eight more elongate unguals.
Pelvic Girdle
The right ilium of AR-1/10 is fused with its ischium and pubis (AR-1-479/10) which are flexed medially due to compaction (Fig. 22 A–D). The acetabulum is completely enclosed as in all derived ankylosaurs [70], [71], [82], [94], [108]. Only Mymoorapelta is known to retain an open acetabulum [84], [109]. The acetabulum is directed verntrally and is situated medially near the contact of the ilium with the sacrum so that the ilium extends far out beyond the acetabulum laterally for a distance nearly equal to its width. The lateral and anterior margins of the laterally oriented ilium are broken away. The prepubic portion of the ilium diverges from the midline of the sacrum at about 30 degrees and is thickened ventrally along its midline. Large, fairly equi-dimensional, closely appressed osteoderms (7-10 cm in diameter) cover the dorsal surface of the ilium posterior to and medial to the acetabulum. As discussed below, this morphology of sacral armor compares well with “Category 3” pelvic armor of Arbour and others [121]. Anteriorly, the smooth dorsal surface of the ilium is exposed. The pubis is fully fused to the anterior margin of the ischium with no visible sutures; its presence is indicated by a slot-shaped foramen along the anterior side of the ischium. This foramen represents the obturator notch between the postpubic process and the main body of the pubis as in Scelidosaurus and stegosaurs [122]. The distal end of the ischium is broken away. Additionally, AR-1-129/10 is a poorly preserved, proximal left ischium with the pubis fully fused to its anterior margin (Fig. 22 E, F).
Beyond some relatively uninformative fragments of the ilium (Fig. 23 A–C), AR-1/31 includes both the right (AR-1-3648/31) and the left (AR-1-3649/31) ischia with fully fused pubes (Fig. 23 D–M). Both exhibit the slot-shaped foramen along the anterior side of the ischium formed by the obturator notch. The proximal ends appear enrolled such that the anterior and posterior margins are nearly parallel due to compaction. Both display an anterior kink at their distal end as in Cedarpelta [86], [88], but overall are straight-shafted as in the Ankylosauridae [70], [82], [123] and the other European nodosaurids Struthiosaurus [31] and Hungarosaurus [32]. The distal end of the left ischium is the best preserved and measures 299.9 mm long along its anterior margin, including the fully fused pubis forming an ischiopubis. Given the asymmetry of the proximal end of the fused ischium and pubis and the position of the obturator foramen, it appears that the pubis still makes up some of the acetabular margin. The contact between the ilium and the fused ischiopubis is straight with about one-fourth to one-third of the acetabulum formed by the fused ischiopubis.
A straight ischium has been considered to be the primitive character state for ankylosaurs, with the bent ischium of Polacanthus and nodosaurids, a derived character [63], [82], [83], [94], [114], [123]. It is possible that as opposed to being primitive, a straight ischium may be secondarily acquired in the ankylosaurids and European nodosaurids. The only known ischium from the Jurassic ankylosaur (Mymoorapelta) is bent, a trait that is also observed in some stegosaurs such as Kentrosaurus [124]. Stegosaur ischia, even when straight, have an angular thickening near the mid-point of the posterior margin [124] that is shared by the polacanthids Mymoorapelta (Kirkland pers. obs.) and Gastonia [83]. Europelta is the oldest known ankylosaur preserving a straight ischium. The slight kink in the distal end of the ischium of Europelta suggests the straight ischium in European nodosaurids and ankylosaurids is achieved by shortening the ischium distal to the bend.
Hindlimb
The right femur, tibia, and fibula were closely associated (Fig. 24 A–F). The robust right femur (AR-1-3244/31) is 502.9 mm long and 178.9 mm wide at the proximal end and has been flattened anteroposteriorly, with the most distortion to the mid-shaft region. The femoral head is distinct with much of its articular surface directed dorsally and only somewhat medially. It forms an angle of about 115° with the long axis of the femur. The femoral head is directed more dorsally under the ilium in polacanthids [7], [12], [82], [95], [125], and several nodosaururids. In addition, the femoral head of Europelta is expanded such that it overhangs the femoral shaft both anteriorly and posteriorly. The greater trochanter is well demarcated from the femoral head by a constriction across the proximal end of the femur, and the anterior trochanter forms a ridge ventral to the greater trochanter that is fully fused to the femur. The robust fourth trochanter overlaps the midpoint of the femoral shaft and its midpoint is located proximal at the midpoint of the femur. Polacanthids and nodosaurid ankylosaurs have this configuration, whereas in ankylosaurids the fourth trochanter is distal to the middle of the shaft [63], [82], [95], [120], [125]. The distal end of the femur is flattened and forms a planar articular surface relative to the straight femoral shaft. The intercondylar notch is not expressed ventrally, and is better developed posteriorly than anteriorly
The right tibia (AR-1-3237/31) and fibula (AR-1-3238/31) were closely associated (Fig. 6) and post-depositionally compressed. Compression has distorted the distal end of the tibia such that the wide posterior surface is twisted counterclockwise in line with the wide lateral side of the anterior end relative to the orientation of the proximal and distal ends of the tibia in most other ankylosaurs, such as Mymoorapelta [84] (Kirkland, pers. obs.). The fibula was taphonomically displaced ventrally and with the ventral end rotated posteriorly relative to its position in life with the tibia.
The tibia is 458.8 mm long and robust for its entire length (Fig. 24 G–K, Q) as in Cedarpelta [86]. The proximal end is 169.2 mm wide by 93.1 mm wide and its distal end is 146.8 mm wide by 70.2 mm. It is significantly more narrowly waisted in Mymoorapelta [84], Gastonia [83], Polacanthus [7], [12], [18], Sauropelta [69], [71], [99], [108], Peloroplites [86], and in Zhejiangosaurus [126] and ankylosaurids like Saichania [106]. The cnemial crest is broadly rounded. The even curvature of the distal end of the tibia suggests that the astragalus was fully fused to it with no evident sutural contact as in most ankylosaurs [63], [82], [121]. The astragalus is not fused to the distal end of the tibia in Mymoorapelta [84], Gastonia [83], Hylaeosaurus [11], and Peloroplites [86].
Generally, ankylosaurids have tibiae that are less than two-thirds the length of their femora, as opposed to nodosaurids which have proportionally longer lower leg elements [127]. With a tibia to femur ratio of 0.91, Europelta has the proportionally longest tibia of any ankylosaur for which this ratio is known. Both Cedarpelta and Peloroplites have relatively longer tibiae than other ankylosaurs [86], with a tibia to femur ratio of 0.82 in both. Peloroplites differs in its proportionally more narrowly waisted tibial shaft.
The fibula is 395.5 mm long (Fig. 24 L–P, R) and laterally flattened. The proximal end is not expanded anteroposteriorly, such that the slender fibula changes little in size and shape from the proximal to distal end. In lateral view, the proximal end is rounded and the distal end is concave. In cross-section, it is flattened medially and convex laterally. It is longer relative to the tibia than in most other ankylosaurs [108].
A calcaneum (AR-1-3289/31) was identified in association with the lower right leg of AR-1/31. It is laterally compressed, convex laterally and concave medially (Fig. 24 S, T). Its dorsal margin is flattened where it would articulate with the fibula. Calcanea are practically unknown in ankylosaurs, but one has been identified in the juvenile specimen of the derived ankylosaur Anodontosaurus [128]. The type of Niobrarasaurus coleii preserves an articulated lower hind limb, with an astragalus fully fused with the tibia and possessing an articulation with the distal end of the fibula and an unfused calcaneum of similar morphology to that of Europelta [118]. The calcaneum is fully fused to the distal end of the fibula in Saichania [106].
A number of metatarsals and phalanges are associated with AR-1/31. The metatarsals have subrectangular proximal ends, indicating that they were closely articulated in a well-integrated pes in life (Fig. 25 A–W). The pedal phalanges (Fig. 25 X–JJJ) are short, as in other ankylosaurs. There are eight relatively large, elongate, spade-like unguals (Fig. 25 KKK–WWWW) of a morphology similar to pedal unguals in other ankylosaurs in which the unguals are nearly as long as the digits[82], which indicates that portions of both feet are present in AR-1/31. We interpret that the pes of Europelta possesses four pedal phalanges as in most other nodosaurids [80]. Liaoningosaurus has three digits on the pes. The eight similar unguals are interpreted as pedal unguals and the smallest ungual (Fig. 25 XXXX–BBBBB) is interpreted as an isolated manual ungual. The overall proportions of the preserved pedal elements are similar to those of Niobrarasaurus [119], which also has pedal unguals nearly as large as its metatarsals.
Armor
There was an abundance of dermal armor recovered with both AR-1/10 and AR-1/31. On comparison with the quarry maps, none of the osteoderms appears to be preserved in situ with any of the skeletal elements or with each other, and there is no fusion between any of the osteoderms recovered. Therefore, the armor has been divided into several broad morphotypes for the purpose of description and comparison to armor described for other ankylosaurs. Although morphotypes and terminologies have been proposed [129], [130], no system fits for all armor types in all ankylosaurs. A number of researchers have divided armor into types as in Type 1, 2, etc. [131]; for this discussion the armor types are alphabetized to ensure minimal confusion with previous descriptions. The term osteoderm is used to describe relatively larger dorsal and lateral armor elements with the presence of an external keel or tubercle, whereas the term ossicle describes relatively smaller dermal armor lacking a keel, in the sense of Blows [130]. It is recognized that a consistent methodology for describing armor is achievable, but must be done within a phylogenetic framework to be of maximum utility.
Osteoderm surface texture may be broadly useful in differentiating ankylosaurids from nodosaurids [132], [133]. The vast majority of the osteoderms examined in Europelta has a moderately rugose texture with sparse pitting more in keeping with nodosaurids and basal ankylosaurids rather than more derived ankylosaurids. Whereas histological studies have proven useful in the study of thyreophorans [132], [134], [135], that is beyond the scope of this study.
It is noteworthy that no portions of distinct cervical rings were recovered, although cervical vertebrae are known for both skeletons of Europelta. Additionally, only one spine from the cervical or pectoral region was tentatively identified. We postulate that these elements were lost through the process of coal removal or may have been taphonomically removed from the skeletal associations. Only the discovery of additional specimens of Europelta can further reveal the presence of cervical half-rings.
Type A armor.
An isolated fragmentary spine (AR-1-128/10), possibly from the cervical or pectoral region, is recognized from the holotype (Fig. 26 A–D). It appears to represent only the anterior half and may have been cut in two as the overlying coal was removed. This sharp, broken margin reveals an asymmetric, Y-shaped cross-section. The base flares more and is is less excavated than in a Type 2 caudal plate, suggesting that it was positioned on a broad flank of the body. From the possible anterior margin, the spine slopes posteriorly 15 cm to the broken margin in a gradual arc. There is no indication that the spine could not have been longer. The spine is compressed as in the cervical spines of Sauropelta [77], [99] and Edmontonia [110], [136], and the pectoral spines of Gastonia [83] and Polacanthus [7], [10]. The base is asymmetrical in a manner similar to the elongate osteoderms in Mymoorapelta [84], with one side of the base extending lower anteriorly and the other posteriorly. There is no evidence of a basal plate incorporated into fusion of the cervical half-ring as in mature ankylosaurs like Mymoorapelta [84] Gargoyleosaurus [85], [95], Gastonia [83], Polacanthus [10], [130], and Sauropelta [77], [99]. This may relate to the anchoring of larger elements into the dermis in Gastonia and Polacanthus [130]. We tentatively interpret AR-1-128/10 as a pectoral spine. However, if the complete element extends beyond the break for more than twice the length of the preserved portion, it would fall into the category of Type B armor, although that is unlikely because it is more massive form than the Type B elements.
Type B armor.
Dorsoventrally compressed, hollow, asymmetric-based plate-like osteoderms with sharp anterior and posterior edges and lateroposteriorly directed apices are identified for AR-1/10 (Fig 26 E–J) and AR-1/31 (Fig. 27 A–L). Similar large osteoderms have been described as caudal plate ostederms in Mymoorapelta [84], [109], Gargoyleosaurus [85], [95], Gastonia [83], and Polacanthus [8]-[10], [38], [130]. Similar, more anterorposteriorly symmetrical caudal plate osteoderms are also known in Minmi [137], [138] and several Asian ankylosaurids [131]. The few plate-like osteoderms of this morphology that are identified in Europelta are mediolaterally shorter and anteroposteriorly longer with a more posteriorly swept apices. Two pairs of similar plates are known for the holotype of Sauropelta (AMNH 3032), with one of the larger plates being illustrated [99]. One plate from the Yale collections of Sauropelta has a unique double apex (YPM 5490). Given the rarity of Type B armor in Sauropelta and Europelta we hypothesize that caudal plates in these nodosaurids ran down the sides of the tail but decreased in size more rapidly, such that long-keeled osteoderms of Type E morphology made up the lateral armor down most of the length of the tail. It is also possible that these large plate-like osteoderms were on the lateral margin of the sacrum as has been documented by Carpenter and others [106] in Saichania. Struthiosaurus preserves several osteoderms of this morphology that have been reconstructed as in Polacanthus as being medial, dorsally-projecting caudal osteoderms [25], [26]. The relative rarity of these plate-like osteoderms suggests that they were restricted to the base of the tail as well.
Type C armor.
Both AR-1/10 (Fig. 28 A–H, O, P) and AR-1/31 (Fig. 29 A– F) preserve fairly large (∼15–25 cm long) subrectangular to subtrapezoidal, solid osteoderms with low, evenly developed keels running down the long axis of the osteoderm either medially or to one side of the mid-line. Their distal and medial surfaces are subparallel and the entire plate may be slightly flexed across the short axis perpendicular to the crest. The straight, longer margins of these plates appear to have been tightly affixed but not fused to adjoining osteoderms. Armor of Type C morphology is not common but is most similar to medial cervical osteoderms of half-rings, and most distinctively, across the mid-line of the pectoral region in some nodosaurids such as Stegopelta [138], Niobrarasaurus [140], [141], Panoplosaurus [74], [101], and Edmontonia [74], [110].
Type D armor.
Both AR-1/10 and AR-1/31 preserve large (∼10-20 cm long) asymmetric, diamond (Fig. 28 I–N, Q–T; Fig. 29 M–P) to tear-drop shaped (Fig. 29 G–L, Q, R, U,V) osteoderms with a long keel rising to an apex medially to posteriorly and in some specimens extending past the posterior margin of the base. They are distinguished from Type E osteoderms because they are wider than 50% of their length. The wider osteoderms are thinner and more solid than the narrower osteoderms with small pockets under the apices. The more diamond-shaped forms may be more closely appressed to each other in anterior bands similar to Type C armor.
Type D Armor is widely known in the nodosaurids such as Sauropelta [99], Panoplosaurus [101], and Edmontonia. Gastonia is documented to have similar armor [142], although more solid in cross section with less basal excavation, which occurs in oblique rows anterior to the sacrum with each osteoderm separated by a single row of small Type H ossicles. This pattern is similar to the dorsal dermal ornamentation documented for the ankylosaur Tarchia by Arbour and others [130], except that in Tarchia most of the intermediate scales lacked ossified cores. Similar armor is known from the lateral sides of the legs in some ankylosaurs such as Saichania [106].
Type E armor.
Both AR-1/10 and AR-1/31 preserve large (10-15 cm long) moderately asymmetric osteoderms more than twice as long as wide with a long keel higher on the assumed posterior end (Fig. 28 Q, R, II–NN; Fig. 30 G–FF). These osteoderms have proportionally more deeply excavated bases than Type D armor, have chevron-shaped cross-sections, and are distinguished from Type D armor by their width being less than 50% of the length. Type E armor is gradational with Type D armor (Fig. 28 S–T; Fig 29 A–F) and may represent lateral or distal armor from the trunk of the body and along the sides of the tail. This armor type is present in Sauropelta [99] and Texasetes [115]. Similar armor is present on the sides of the limbs in Scelidosurus and Saichania [106].
Type F armor.
Medium to large (∼5-15 cm long) oval to circular osteoderms of low profile with a median keel extending into an apex near or overhanging the posterior margin of the osteoderm are represented in both AR-1/10 (Fig. 28 U–Z) and AR-1/31(Fig. 29 W–VVV). The basal surface of the osteoderm is generally solid except for a small pocket under the apex, reminiscent of Type D armor. Less commonly, the base may be more extensively excavated. Armor of this morphology is abundant in many nodosaurids and makes up the major elements of the armor of Sauropelta anterior to the sacrum in AMNH 3036 [142] and is present in Panoplosaurus [101]. These osteoderms may reside within more expansive spaces among the larger dorsal armor as in Edmontonia (AMNH, 5665) and the polacanthids [81], [82], [93], [107], or may be major armor elements on the posterior portion of the sacrum as in Sauropelta (AMNH 3036). They may also lie on the tail between the Type B caudal plate-like osteoderms, or could be arranged along the lateral side of the limbs as in Saichania [106].
Type G armor.
One piece (AR-1-192/10) of flat, oval to subtriangular armor (AR-1-192/10) from AR-1/10 is about 12 cm long and 7 cm wide and is about 0.5 cm thick throughout (Fig. 28 AA, BB). A pair of similar, osteoderms from the Sauropelta specimen AMNH 3032 was curated with a note from the collector, Barnum Brown, stating that these distinct osteoderms were associated with the forelimbs. Therefore, we suggest a similar position for Type G armor in Europelta.
Type H armor.
Small (∼1-4 cm long) solid ossicles are abundant, with 71 examples from both AR-1/10 (Fig. 31) and AR-1/31 (Fig. 32) illustrated. These ossicles range in shape from round, to oval and even irregularly shaped, and are probably filling in the spaces between larger osteoderms. Small interstitial ossicles are not known for every ankylosaur taxon, but appear to be present in many nodosaurid taxa such as Sauropelta [99], [143] and Edmontonia [74], [136], in polacanthid ankylosaurs such as Gastonia [83] and in some ankylosaurids such as Tarchia [131], in which epidermal scales interstitial to osteoderms do not preserve deeper, interstitial ossicles. Their absence may be real, in that they never form deep to the epidermal scales, taphonomic, in that they are selectively transported away because of their small size and low density, or ontogenetic; in that they only ossify late in ontogeny. The surface texture of Gastonia ossicles is smoother than those of Europelta.
Sacral armor.
Armor is present on the posterior margin of the ilium AR-1-479/10. It is composed of large, subequal-sized (7-10 cm) osteoderms that are tightly sutured together (Fig. 22 A) as in the poorly known Stegopelta [139], Nodosaurus [113], Aletopelta [127], and Glyptodontopelta [132], [144]. These low-relief ossicles lack a central apex or keel. The boundary between the margins of the osteoderms and the area devoid of osteoderms on the ilium is sharply demarcated along the margins of unbroken osteoderms, suggesting the armor was not coossified as in Aletopelta [127] and unlike the fully fused sacral armor in the polacanthids Polacanthus and Gastonia [63], [83]. This form of pelvic armor fits that of Arbour and others' Category 3 pelvic armor [121].
Additionally, there is a unique osteoderm AR-1-653/10 that has a large, posteriorly-curved, plate-like keel extending out from the surface that, considered in isolation, is comparable in size and morphology to Type B armor (Fig. 26 K–N). The base is smooth and gently convex, suggesting it may have been closely appressed to the more anterior portion of the ilium. In overall morphology, this large osteoderm is comparable to the spine-bearing armor plate-like osteoderm identified in Hungarosaurus and interpreted to be present in Struthiosaurus [33].
Unique armor pieces.
Some irregularly shaped armor specimens are not represented by more than one element among this material or in the armor from other taxa. At this time, we can offer no positional interpretation of this armor. AR-1-447/10 is an irregular mass of what we interpret as an osteoderm, although it could be sacral armor (Figure 28 CC–FF). AR-1-438/10 is a small, cap-shaped shaped with a small excavation in the center of the external surface (Fig. 28 OO, PP). Two small, deeply basally excavated, oval osteoderms (Fig. 30 GG–JJ) were collected from AR-1/31(AR-1-3239/31, 3721). These osteoderms lack the external excavation.
Discussion
Europelta (Fig. 33) can be distinguished from any of the ankylosaurs assigned to the Polacanthidae (sensu Kirkland's Polacanthinae [83] and Carpenter's Polacanthidae [63] from the Upper Jurassic and Lower Cretaceous as defined by Yang and others [64]; see Terminology) by its rounded, tear-drop shaped skull and a suborbital horn developed on the posterior portion of the jugal and the quadratojugal posterior to the orbit, as opposed to a triangular-shaped skull that is widest at the posterior margin and a suborbital horn developed exclusively on the jugal (as seen in polacanthids). Post-cranially, it can also be distinguished from polacanthids, by its elongate lower hind limbs, the apparent rarity of cervical, pectoral, and thoracic spines, and reduction in the number of caudal plate-like osteoderms. Likewise, it has an abundance of Type D, asymmetric, tear-drop shaped osteoderms like those observed in many nodosaurids and absent in all polacanthids.
Europelta is also distinguished from derived ankylosaurids by its weakly ornamented teardrop-shaped skull in which the lower temporal opening is visible in lateral view. The absence of a tail club also distinguishes the taxon from these ankylosaurids. More basal “shamosaurine grade” ankylosaurids [63], [86] are more similar to Europelta, but also have the lower temporal openings completely obscured laterally by expanding the lateral margin of their skulls. “Shamosaurine grade” ankylosaurids also possess skulls that are approximately as wide mediolaterally between the orbits as they are across the posterior margin.
Europelta shares a number of derived characters with nodosaurids [71], [72], [83], [94], [114]. It has a tear-drop shaped skull that is longer than wide with its greatest width dorsal to the orbits, whereas the short, boxy skulls of Minmi and all anklosaurids are essentially as wide at the posterior edge of the skull, as are the elongate skulls of “shamosaurine-grade” ankylosaurids. Grooves in the remodeled textured skull roof define epidermal scale impressions, with the largest covering the frontoparietal area. Although poorly preserved, the laterally extensive pterygoids are pressed up against the anterior face of the braincase. All known nodosaurid scapulae have a prominent acromion process extending on to the blade of the scapula that terminates in an expanded knob. Unfortunately, this portion of the scapula is as yet unknown in Europelta.
Some character states considered typical of nodosaurids are absent in Europelta. Instead of having a distinct hourglass-shaped palate typical of nodosaurids [70], [71], [82], [83], [114], the upper tooth rows show less lateral emargination and diverge posteriorly. This is also true of Silvisaurus, which also shares an expanded lateral wall of the skull [76], [77]. The coracoid of Europelta is nearly as long as it is tall, whereas in other nodosaurids, for which the corocoid is known, it is expanded anteriorly and longer than tall [71], [72], [83], [94], [114].
The only other Early Cretaceous nodosaurid to have large cranial scales as in Europelta is Propanoplosaurus, known only from an embryonic to hatchling specimen from the base of the Potomac Group of Maryland [145]. However, only the anterior cranial scales are well defined in Propanoplosaurus, whereas only the posterior scale pattern in Europelta. The unusual preservation and extremely small size of Propanoplosaurus lead us to suspect that the fossil preserves the actual scales overlying the skull and not the remodeled skull roof, because this is such a young specimen and remodeling of the cranial bones is not expected to have occurred so early in ontogeny [129], [146].
Additionally, a number of important characters traditionally used to define nodosaurids are not known in Europelta, as yet, because of the missing anteroventral half of the scapula and the absence of premaxilla and surangulars. Thus, the presence absence of premaxillary teeth, if the tooth row joined the margin of premaxillary beak, the morphology of the naris, the height of the coronoid process, and the morphology of the acromion process are unknown for Europelta.
Europelta is distinguishable from European nodosaurids from the Albian through the Cenomanian. The juvenile Anoplosaurus from the Albian Gault Clays of southern England differs in a number of characters, such as possessing a proportionally longer coracoid, a narrower proximal end of the humerus, and a femur with a separate anterior trochanter [17] although the latter two characters are consistent with the juvenile nature of Anoplosaurus. No pectoral spines of the morphology described for “Acanthopholis” from the Cenomanian Lower Chalk in southern England by Huxley [13] are known in Europelta. Additionally, the tall teeth assigned to “Acanthopholis” are distinct in the long apicobasal ridges extending from the denticles to the root on medial and lateral faces of the teeth, and in the presence of caudal ribs that extend laterally and flex ventrally, whereas the caudal ribs in Europelta extend ventrolaterally and flex laterally [16], [17]. Europelta is like other Late Cretaceous European nodosaurids in having a short symphysis for the predentary, a mediolaterally wide and anteroposteriorly thin quadrate, an anteroposteriorly arched sacrum, and a straight ischium [21], [32].
The domed skull and elongate cervical vertebrae in Struthiosaurus clearly distinguish it from Europelta. Likewise, Hungarosaurus also has more elongate cervical vertebrae [32]. Both Hungarosaurus and Struthiosaurus possess a pair of spines on the anterior portion of the pelvis [33], whereas we interpret the presence of a pair of upright plate-like armor elements in this position in Europelta (Fig. 33).
The lateral wall of the skull in most North American nodosaurids is typically narrow [82], whereas in Europelta it is relatively wider, although a broad notch along its posterior margin permits the caudal margin of the lower temporal opening to be observed in lateral view. This morphology in Europelta is similar to that in the nodosaurids Silvisaurus [76], [77] and Peloroplites [86]. Although, the skull of Struthiosaurus transylvanicus is highly reconstructed [22], it appears that the lateral wall of the skull is expanded laterally, whereas not completely obscuring the lower temporal opening. This character state is not known in other species of Struthiosaurus, but appears to be moderately developed in Hungarosaurus [32].
Comparisons of Europelta with the Asian”nodosaurids” Zhongyuansaurus [93] and Zhejiangosaurus [126] from the lower Upper Cretaceous of China hinges partially on the question of whether those taxa have been validly referred to Nodosauridae. Carpenter and others [86] noted that the skull of Zhongyuansaurus is morphologically similar to that of a “shamosaurine-grade” (like Shamosaurus and Gobisaurus) ankylosaurids and was the first shamosaurine-grade ankylosaurid documented to not have a tail club. However, its distal tail is modified into a stiffened structure of the same morphology as the “handle” of the tail club in more derived ankylosaurids [147], [148]. Zhejiangosaurus was assigned to the nodosaurids based on characteristics of the femur and sacrum, together with the lack of a tail club [126]. We hypothesize that it lacked a knob as in basal ankylosaurids, polacanthids and nodosaurids because ankylosaurids with a full tail club have distal free caudal vertebrae bearing caudal ribs at the base of the handle. Most of the distal caudal vertebrae of Zhejiangosaurus have raised ridges on the sides of the centra as in the distal vertebrae of polacanthids and nodosaurids. Additionally, whereas the position of its most proximal preserved caudal vertebrae is not known, morphologically, they do not appear to represent the most proximal caudal vertebrae. Thus, while Zhejiangosaurus' 13 preserved caudal vertebra are more than the number of free caudals preserved in most ankylosaurs with tail clubs (10 in Saichania [106] and Dyoplosaurus [148]), the total number of free caudals in its tail would appear to be more than the 14 in Tarchia [130] and 15 in Pinacosaurus [129]. Unlike nodosaurids, Zhejiangosaurus has an exceedingly low ratio of femur to tibia length of 0.46 similar to that of with ankylosaurids and polacanthids rather than nodosaurids. Dongyangopelta [149] was described as a second nodosaurid from the same area and stratum as Zhejiangosaurus, which was found to be its sister taxon in their phylogeny [149]. With few overlapping elements, we feel that the proposed differences between these taxa may be due to preservation, individual variation, or ontogeny. Additionally, given the presence of a pelvic shield and numerous caudal plate-like osteoderms in Dongyangopelta, we suggest that both specimens may pertain to the same taxon and represent the first polacanthid described from Asia. Given the recent description of the polacanthid Taohelong from the upper portion of the Lower Cretaceous of Gansus Province in western China [64], this hypothesis has added support. We also do not think that the partial ankylosaur skull reported from the lower Upper Cretaceous of Hokkaido, Japan [150] can be diagnosed as a nodosaurid with any confidence at this time, due to the incomplete nature of the specimen. Thus, we do not presently recongnize the presence of true nodosaurids in Asia.
In his seminal paper defining a bipartite division of the Ankylosauria into Ankylosauridae and Nodosauridae, Coombs [71] hypothesized that Acanthopholis (as a nomen dubium in which he would have included Anoplosaurus) and Struthiosaurus might represent a separate lineage of European nodosaurids. Unlike Hylaeosaurus (in which he included Polacanthus), these taxa had a well-developed supraspinus fossa developed anteriorly on the scapula as did all North American nodosaurids. This European lineage was hypothesized based on their small body size, presence of premaxillary teeth, and their possessing an unfused scapula and corocoid. Although, none of the characters are valid in defining such a group, our research on Europelta has resulted in supporting the taxonomic hypothesis of Coombs [71], [72] as correct, just for the wrong reasons.
Relationships to Other Taxa
We use Struthiosaurinae to define the clade of European nodosaurs. Nopcsa [25] proposed Acanthopholidae as a family of relatively lightly built thyreophorans, that included Acanthopholis ( = Anoplosaurus), Polacanthus, Stegopelta, Stegoceras, and Struthiosaurus. In 1923, he divided the Acanthopholidae into an Acanthopholinae and a Struthiosaurinae without comment [69]. Subsequently, he relegated the Acanthopholidae to a subfamily of the Nodosauridae, in which he also included Ankylosaurus and restricted the Acanthopholinae to Acanthopholis, Hylaeosaurus, Rhodanosaurus, Struthiosaurus, Troodon [26], [151]. This artificial grouping included a polacanthid ankylosaur [72], [83], a pachycephalosaur [152] and Acanthopholis, now considered a nomen dubium [17], [82]. Thus, the term Acanthopholinae is not acceptable for this newly recognized clade of nodosaurids. Thus, Struthiosaurinae is the next published term available to use for this clade and is derived from the first described and youngest member of this clade. Struthiosaurinae is defined as the most inclusive clade containing Europelta but not Cedarpelta, Peloroplites, Sauropelta or Edmontonia.
In order to determine the systematic position of Europelta, it was found that previous cladistic analyses [71], [72], [82], [83], [114], did not include many of the character states that we identify as significant in our research on Upper Jurassic and Lower Cretaceous ankylosaurs. A major weakness of these analyses is the limited recognition of postcranial skeletal and dermal characters that restricts the testing the phylogenetic relationships for taxa for which skulls are either poorly known or not known at all.
We present a character based definition of Struthiosaurinae as: nodosaurid ankylosaurs that share a combination of characters including: narrow predentary; a nearly horizontal, unfused quadrate that is oriented less than 30° from the skull roof, and mandibular condyles that are 3 times transversely wider than long; premaxillary teeth and dentary teeth that are near the predentary symphysis; dorsally arched sacrum; an acromion process dorsal to midpoint of the scapula-coracoid suture; straight ischium, with a straight dorsal margin; relatively long slender limbs; a sacral shield of armor; and erect pelvic osteoderms with flat bases. This suite of characters unites Europelta with the European nodosaurids Anoplosaurus, Hungarosaurus and all species assigned to Struthiosaurus. This clade of European nodosaurids has not been previously recognized. Europelta represents the earliest member of the European clade Struthiosaurinae.
Biogeogeographic Implications
The near simultaneous appearance of nodosaurids in both North America and Europe is worthy of consideration (Fig. 34). Europelta is the oldest nodosaurid known in Europe, it derived from strata in the lower Escucha Formation that is dated to early Albian. The oldest nodosaurid from western North America is Sauropelta, which in the lower part of its range is in the lower Albian Little Sheep Mudstone Member (B interval) of the Cloverly Formation in northern Wyoming and southern Montana [99], [153] with an ash bed 75 meters above the base near the top of the member providing an age of 108.5±0.2 Ma [154]. Nodosaurid remains from eastern North America appear to be older. Teeth of a large nodosaurid Priconodon crassus are known from the Arundel Clay of the Potomac Group [77], [155], which palynology dates as near the Albian-Aptian stage boundary [156]. The hatchling Propanoplosaurus is from the base of the underlying Patuxent Formation of the Potomac Group of Maryland, which has been dated as late Aptian [157], [158], making Propanoplosaurus the oldest known nodosaurid. Polacanthid ankylosaurs characterize pre-Aptian faunas in both Europe [11], [12], [37]-[39] and North America [70], [95], [159]. We have not been able to document a specific example of Polacanthus in the Lower Aptian Vectis Formation of the Wealden Group, although Polacanthus has been reported to occur in those strata [10]-[12], [82], [160]. However, polacanthids are present in the lower Aptian Morella Formation of northeastern Spain [40]. Blows [10] illustrated a block with ankylosaur dorsal vertebrae with the uninformative ventral portion of a pelvic shield fragment and noted it as being from Charmouth, suggesting that there were upper Albian polacanthids in England [160]. However, the specimen NMW 92.34G.2 was actually found on the beach further west at Charton Bay and may have come from either the Aptian (Lower Greensand) or Albian (Upper Greensand). Only preparation of the dorsal surface of the pelvic shield would reveal if the specimen is a polacanthid or nodosaurid. A large polacanthid (BYU R254) occurs in the Poison Strip Sandstone Member of the Cedar Mountain Formation [156]. It is not a nodosaurid close to Sauropelta as reported by Carpenter and others [97], but a polacanthid that was initially described as cf. Hoplitosaurus [161]. These rocks have been dated as lower to middle Aptian by laser ablation of detrital zircons and by U-Pb dating of early diagenetic carbonate [162]. A fragmentary large nodosaurid with massive cervical spikes that may be referred to as cf. Sauropelta (DMNS 49764) has been recovered from the overlying Ruby Ranch Member about 20 m up section in the same region [163] in strata interpreted to be of Lower Albian age [162]. Thus, the youngest polacanthids occur in the lower to possibly mid-Aptian and the oldest documented nodosaurids occur in the upper Aptian or lower Albian in both Europe and North America with no discernible stratigraphic overlap (Fig. 34). Why this faunal discontinuity occurs is unknown. There are no documented significant changes in sea level or shifts in geochemical indicators to suggest a geological or environmental change that would affect ankylosaurs on both continents at approximately the same time [164]. However, the OAE1a or “Sella” organic burial episode near the base of the Aptian was followed by a positive carbon isotope excursion that may have precipitated longer-term environmental effects that would result in the turnover of ankylosaurs in the “middle” Aptian [165]. In North America, “medial” grade iguanodonts (basal Steracosterna) are replaced by the considerably more primitive basal iguanodont Tenontosaurus at this time, while in Europe the lower Albian more derived iguanodont Proa is phylogenetically close to Iguanodon [43], [159] at the base of Hadrosauriformes [43], documenting different patterns of faunal change for iguanodonts and ankylosaurs. Therefore, a cause for this faunal turnover, which might specifically have affected ankylosaurs, should be sought. Ankylosaurs are low feeders, so perhaps the rapid ongoing radiation of flowering plants at this time [166]-[170] might have driven their diversification. It has been proposed that this floral revolution was linked to a decline in atmospheric CO2 concentrations [171] or, more likely, an increase in CO2 and global warming resulting from massive early Aptian volcanic activity forming the Ontong Java and Manihiki plateaus [172], [173]-[174]. Therefore the rapid domination of shrubby angiosperms may have caused a disruption in the availability of forage to which polacanthids were adapted. Kirkland and others have proposed that North America became isolated from Europe at the end of the Barremian [159], [175]. Certainly the timing of the appearance of nodosaurids on both continents indicates that the origins of the clade preceded the complete isolation of North America and Europe pushing up this date in to at least the “middle” Aptian. The separation of the Nodosauridae into a North American Nodosaurinae and a European Struthiosaurinae by the end of the Aptian, would thus provide a revised date for the isolation of North America from Europe with rising sealevel.
Additionally, whereas there is no definitive evidence for nodosaurids in Asia, apparently polacanthids entered Asia in the later portion of the Early Cretaceous and survived there in isolation into the early Late Cretaceous.
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https://www.academia.edu/66199999/Campanian_Maastrichtian_Ankylosaurs_of_West_Texas
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en
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Campanian-Maastrichtian Ankylosaurs of West Texas
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] |
[] |
[] |
[
""
] | null |
[
"Bryanna West",
"tcu.academia.edu"
] |
2021-12-28T00:00:00
|
Big Bend National Park is known for its unique Late Cretaceous fauna, such as Alamosaurus sanjuanensis and Quetzalcoatlus northropi. Most major groups of dinosaurs are represented in the Late Cretaceous strata, which ranges from the Early Campanian
|
https://www.academia.edu/66199999/Campanian_Maastrichtian_Ankylosaurs_of_West_Texas
|
"A new, small ankylosaurid, Ahshislepelta minor, from the upper Campanian Kirtland Formation (Hunter Wash Member), San Juan Basin, New Mexico, consists of shoulder girdle and forelimb elements, vertebral fragments, and numerous osteoderms. Ahshislepelta minor differs from other ankylosaurids on the basis of a prominent dorsolateral overhang of the acromion and its osteoderm texture. It ranks as one of the most complete ankylosaur specimens known from New Mexico and adds to our understanding of ankylosaurid paleobiogeography, stratigraphy, and taxonomy."
Isolated bones and osteoderms of ankylosaurian dinosaurs recovered from Late Cretaceous sediments of northern Coahuila, northeastern Mexico, have been identified as remains of nodosaurids. Here, we summarize these discoveries and provide a review on Mexican Ankylosauria from a taxonomic perspective. We also present a new taxon, Acantholipan gonzalezi gen. et sp. nov. from the Pen Formation and provide a phylogenetic analysis integrating the new taxon. A. gonzalezi is the first named ankylosaur from Mexico that adds to the currently rare nodosaurid diversity from southern Laramidia.
Nodosaurid ankylosaur remains from the Upper Cretaceous of Mexico are summarized. The specimens are from the El Gallo Formation of Baja California, and the Pen and Aguja Formations of northwestern Coahuila, Mexico. These specimens show significant differences from other known nodosaurids, including ulna with very well developed olecranon and prominent humeral notch, the distal end of the femur not flaring to the extent seen in other nodosaurids, and a horn-like spine with vascular grooves on one side. The specimens are important because they are the southern-most occurrences in North America, and provide an important biogeographical link between nodosaurids of the United States and Canada on the one hand, and Argentina and Antarctica on the other.
Fossil evidences of the presence of ankylosaurian dinosaurs in Gondwana are scarce but consistent, being found in Antarctica, Oceania and South America. In spite that there are no nominated species in South America, the ankylosaur fossil record has increased in the last years. Indeterminate nodosaurid specimens, some isolated osteoderms and many trackways are known from the Upper Cretaceous of South America. The aim of the present contribution is to report new ankylosaurian remains from the Allen Formation (Campanian-Maastrichtian) at the Salitral Moreno locality, Northern Patagonia, Argentina. These osteoderms are small and conical, and includes thoracic, sacral and caudal scutes. The thoracic and sacral pieces are similar to those belonging to nodosaurids. The caudal osteoderm is a new element for the record of South American ankylosaurs. It resembles the caudal plates of Kunbarrasaurus and some ankylosaurids. The scutes show a mixture of characters so it is not possible to assign these pieces to a nodosaurid-like or ankylosaurid ankylosaur. These elements are consistent with the previously known ankylosaur fossil record of the Upper Cretaceous of Argentina, being a new sample of the diversity of the latest Cretaceous from South America.
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https://cyberleninka.ru/article/n/the-braincase-of-bissektipelta-archibaldi-new-insights-into-endocranial-osteology-vasculature-and-paleoneurobiology-of
|
en
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THE BRAINCASE OF BISSEKTIPELTA ARCHIBALDI - NEW INSIGHTS INTO ENDOCRANIAL OSTEOLOGY, VASCULATURE, AND PALEONEUROBIOLOGY OF ANKYLOSAURIAN DINOSAURS Текст научной статьи по специальности « Биологические
|
https://cyberleninka.ru/article/n/the-braincase-of-bissektipelta-archibaldi-new-insights-into-endocranial-osteology-vasculature-and-paleoneurobiology-of/og
|
https://cyberleninka.ru/article/n/the-braincase-of-bissektipelta-archibaldi-new-insights-into-endocranial-osteology-vasculature-and-paleoneurobiology-of/og
|
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[
"научная статья бесплатно на тему THE BRAINCASE OF BISSEKTIPELTA ARCHIBALDI - NEW INSIGHTS INTO ENDOCRANIAL OSTEOLOGY",
"VASCULATURE",
"AND PALEONEUROBIOLOGY OF ANKYLOSAURIAN DINOSAURS текст научной работы по биологическим наукам из научного журнала Biological Communications. DINOSAURIA",
"ANKYLOSAURIA",
"ENDOCAST",
"BLOOD VESSELS",
"PALEOBIOLOGY",
"LATE CRETACEOUS",
"UZBEKISTAN"
] | null |
[
"Kuzmin Ivan",
"Petrov Ivan",
"Averianov Alexander",
"Boitsova Elizaveta",
"Skutschas Pavel",
"Sues Hans-Dieter"
] |
2020-08-25T00:00:00
|
We describe in detail three braincases of the ankylosaur Bissektipelta archibaldi from the Late Cretaceous (Turonian) of Uzbekistan with the aid of computed tomography, segmentation, and 3D modeling. Bissektipelta archibaldi is confirmed as a valid taxon and attributed to Ankylosaurinae based on the results of a phylogenetic analysis. The topographic relationships between the elements forming the braincase are determined using a newly referred specimen with preserved sutures, which is an exceedingly rare condition for ankylosaurs. The mesethmoid appears to be a separate ossification in the newly referred specimen ZIN PH 281/16. We revise and discuss features of the neurocranial osteology in Ankylosauria and propose new diagnostic characters for a number of its subclades. We present a 3D model of the braincase vasculature of Bissektipelta and comment on vascular patterns of armored dinosaurs. A complex vascular network piercing the skull roof and the wall of the braincase is reported for ankylosaurs for the first time. We imply the presence of a lepidosaur-like dorsal head vein and the venous parietal sinus in the adductor cavity of Bissektipelta. We suggest that the presence of the dorsal head vein in dinosaurs is a plesiomorphic diapsid trait, and extant archosaur groups independently lost the vessel. A study of two complete endocranial casts of Bissektipelta allowed us to compare endocranial anatomy within Ankylosauria and infer an extremely developed sense of smell, a keen sense of hearing at lower frequencies (100-3000 Hz), and the presence of physiological mechanisms for precise temperature control of neurosensory tissues at least in derived ankylosaurids.
|
/favicon.ico
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КиберЛенинка
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https://cyberleninka.ru/article/n/the-braincase-of-bissektipelta-archibaldi-new-insights-into-endocranial-osteology-vasculature-and-paleoneurobiology-of
|
FULL COMMUNICATIONS
PALAEONTOLOGY
The braincase of Bissektipelta archibaldi — new insights into endocranial osteology, vasculature, and paleoneurobiology of ankylosaurian dinosaurs
Ivan Kuzmin1, Ivan Petrov2, Alexander Averianov3, Elizaveta Boitsova1, Pavel Skutschas1, and Hans-Dieter Sues4
1Department of Vertebrate Zoology, Faculty of Biology, Saint Petersburg State University, Universitetskaya nab., 7-9, Saint Petersburg, 199034, Russian Federation; 2Saint Petersburg City Palace of Youth Creativity, Nevsky pr., 39A, Saint Petersburg, 191011, Russian Federation;
3Zoological Institute, Russian Academy of Sciences, Universitetskaya nab., 1, Saint Petersburg, 199034, Russian Federation;
4Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, MRC 121, P.O. Box 37012, Washington, DC 20013-7012, USA
Address correspondence and requests for materials to Ivan Kuzmin, kuzminit@mail.ru
Abstract
Citation: Kuzmin, I., Petrov, I., Averianov, A., Boitsova, E., Skutschas, P., and Sues, H.-D. 2020. The braincase of Bissektipelta archibaldi — new insights into endocranial osteology, vasculature, and paleoneurobiology of ankylosaurian dinosaurs. Bio. Comm. 65(2): 85-156. https://doi.org/10.21638/spbu03.2020.201
Authors' information: Ivan Kuzmin, Master of Sci. in Biology, PhD student, Junior Researcher, orcid.org/0000-0003-3086-2237; Ivan Petrov, School student, orcid.org/0000-0003-3617-2317; Alexander Averianov, Dr. of Sci. in Biology, Head of Laboratory, orcid.org/0000-0001-5948-0799; Elizaveta Boitsova, Master of Sci. in Biology, orcid.org/0000-0001-8590-9835; Pavel Skutschas, Dr. of Sci. in Biology, Associate Professor, orcid.org/0000-0001-8093-2905; Hans-Dieter Sues, PhD, Senior Scientist, orcid.org/0000-0002-9911-7254
Manuscript Editor: Nikita Zelenkov, Cabineth of Palaeoornithology, Borissiak Palaeontological Institute, Russian Academy of Sciences, Moscow, Russia
Received: November 21, 2019;
Revised: February 25, 2020;
Accepted: March 10, 2020.
Copyright: © 2020 Kuzmin et al. This is an open-access article distributed under the terms of the License Agreement with Saint Petersburg State University, which permits to the authors unrestricted distribution, and self-archiving free of charge.
Funding: The field work in 1997-2006 was funded by the National Science Foundation (EAR-9804771 and EAR-0207004 to J. D.Archibald and H.-D. Sues), the National Geographic Society (5901-97 and 628198 to J. D.Archibald and H.-D. Sues), and the Navoi Mining and Metallurgy Combinat. The laboratory research received support from the Russian Science Foundation (19-1400020). AA was supported by the Zoological Institute, Russian Academy of Sciences (project AAAA-A19-119032590102-7).
Competing interests: The authors have declared that no competing interests exist.
We describe in detail three braincases of the ankylosaur Bissektipelta archibaldi from the Late Cretaceous (Turonian) of Uzbekistan with the aid of computed tomography, segmentation, and 3D modeling. Bissektipelta archibaldi is confirmed as a valid taxon and attributed to Ankylosaurinae based on the results of a phylogenetic analysis. The topographic relationships between the elements forming the braincase are determined using a newly referred specimen with preserved sutures, which is an exceedingly rare condition for ankylosaurs. The mesethmoid appears to be a separate ossification in the newly referred specimen ZIN PH 281/16. We revise and discuss features of the neurocranial osteology in Ankylosauria and propose new diagnostic characters for a number of its subclades. We present a 3D model of the braincase vasculature of Bissektipelta and comment on vascular patterns of armored dinosaurs. A complex vascular network piercing the skull roof and the wall of the braincase is reported for ankylosaurs for the first time. We imply the presence of a lepidosaur-like dorsal head vein and the venous parietal sinus in the adductor cavity of Bissektipelta. We suggest that the presence of the dorsal head vein in dinosaurs is a ple-siomorphic diapsid trait, and extant archosaur groups independently lost the vessel. A study of two complete endocranial casts of Bissektipelta allowed us to compare endocranial anatomy within Ankylosauria and infer an extremely developed sense of smell, a keen sense of hearing at lower frequencies (1003000 Hz), and the presence of physiological mechanisms for precise temperature control of neurosensory tissues at least in derived ankylosaurids. Keywords: Dinosauria, Ankylosauria, endocast, blood vessels, paleobiology, Late Cretaceous, Uzbekistan.
Introduction
Ankylosaurs constitute a clade of quadrupedal heavily armored ornithischian dinosaurs. Their remains are known from the Jurassic to the Late Cretaceous from every continent except Africa (Tumanova, 1987; Vickaryous et al., 2004). Aspects of ankylosaurian anatomy, phylogeny, and paleobiogeography have been thoroughly studied in the last few decades (e.g., Maryanska, 1977; Tumanova, 1987, 2012; Coombs and Maryanska, 1990; Carpenter, 2001; Vickaryous et al., 2004; Thompson et al., 2012; Arbour and Currie, 2016). Despite this progress, our knowledge of the neurocranial osteology and endocranial morphology within
Table 1. Measurements of the studied braincases of Bissektipelta archibaldi. All linear measurements in millimeters
Parameter ZIN PH 1/16 ZIN PH 281/16 ZIN PH 2329/16
Length from the anterior margin of the sphenethmoidal complex to the posterior tip of occipital condyle 89.2 82.7 84
Depth from the dorsal tip of the laterosphenoid capitate process to the ventral margin of the parabasisphenoid 60.4 58.5 -
Dorsoventral depth of the cranial nerve II foramen 8 6.8 -
Paroccipital process, dorsoventral depth at the mid-section 23.5 22.5 -
Occipital condyle, dorsoventral depth 23.8 21.5 21
Occipital condyle, transversal breadth 36 31.4 42.6
Basioccipital, transversal breadth at the basioccipital-parabasisphenoid contact 52 42 46
Basioccipital, length from the posterior tip of the condyle to the basioccipital-parabasisphenoid contact, in sagittal plane 36 30 35
Foramen magnum, transversal breadth 22 18 18
Foramen magnum, dorsoventral height 19 20 19
Parabasisphenoid, transversal breadth between basipterygoid processes 33 23.8 34
the clade is comparatively poor (see the recent review by Paulina-Carabajal et al. [2018]).
A number of isolated specimens belonging to An-kylosauria are known from the Late Cretaceous of Central Asia (Averianov, 2009). Bissektipelta archibaldi is the only valid ankylosaur species from the territory of the former USSR reported to date. It was initially described as "Amtosaurus" archibaldi based upon a single braincase incorporating the skull roof from the Late Cretaceous Bissekty Formation of Uzbekistan (Averianov, 2002). Later, it was re-assigned to a new genus (Bissektipelta) by Parish and Barrett (2004) as these authors concluded the type species of "Amtosaurus" "A. magnus" is nondiagnostic and should be considered a nomen dubium. Since the initial description, the affinities and phyloge-netic position of Bissektipelta have been debated (Averianov, 2002; Parish and Barrett, 2004; Tumanova, 2012; Arbour and Currie, 2016; see "Phylogenetic analysis" below) but have never been formally assessed. Recently, Alifanov and Saveliev (2019) described a high-quality synthetic endocast made from the holotype of Bissektipelta archibaldi. However, many of their anatomical interpretations and biological inferences appear to be controversial and in need of further review.
Here, we redescribe in detail the holotype of Bissektipelta archibaldi (ZIN PH 1/16) with the aid of CT scanning. Additionally, two new ankylosaur braincases from the Bissekty Formation are described and assigned to the same species. One of these (ZIN PH 281/16) preserves clear sutures between the elements forming the brain-case, which is exceedingly rare for ankylosaurs. Endo-casts for two studied specimens have been made, which is the largest sample for a single species of ankylosaurs to date. A thorough review of the literature and com-
parison between the described taxa allowed us to propose new and revise previously known braincase characters from the most current taxon-character matrices of ankylosaurs (Thompson et al., 2012; Arbour and Currie, 2016; Arbour and Evans, 2017; Zheng et al., 2018) and subsequently test the phylogenetic relationships of Bissektipelta. Based on a solid phylogenetic framework and detailed digital endocranial casts, we discuss aspects of cranial vasculature and inferences concerning the paleobiology of ankylosaurs.
Material and methods
Institutional abbreviations. OUVC, Ohio University Vertebrate Collection, USA; ZIN PH, Paleoherpetologi-cal Collection, Zoological Institute, Russian Academy of Sciences, Saint Petersburg, Russia.
Material. The studied material comprises three braincases: the holotype of Bissektipelta archibaldi (ZIN PH 1/16) and two newly described specimens (ZIN PH 281/16 and ZIN PH 2329/16). The material came from the Late Cretaceous (Turonian) Bissekty Formation at the Dzharakuduk locality in the Central Kyzylkum Desert, Uzbekistan. The measurements for the specimens are provided in Table 1.
The holotype of Bissektipelta archibaldi ZIN PH 1/16 is a well-preserved, fully ossified braincase with a partial skull roof. This specimen was the only known cranial material of the Bissekty ankylosaur and constituted the basis of the original description of "Amtosaurus" ar-chibaldi (Averianov, 2002) and subsequent taxonomic reappraisal of this taxon as Bissektipelta archibaldi (Parish and Barrett, 2004). The newly described specimens include ZIN PH 281/16, a partial braincase of slightly
smaller size with open sutures between some bones, and ZIN PH 2329/16, which is similar in size to the holotype of Bissektipelta archibaldi (Table 1). ZIN PH 2329/16 preserves most of the braincase and part of the skull roof. The sutures cannot be traced in ZIN PH 2329/16 because it is damaged and partially covered with matrix.
Computed tomography. The holotype ZIN PH 1/16 and the referred specimen ZIN PH 281/16 were X-ray CT scanned using a Toshiba Aquilon 64 medical tomographer at 0.5 mm slice thickness, 120 kV, and 300 mA. The resulting stacks compile 334 images (512x512x334 resolution) in DICOM format for ZIN PH 1/16 and 149 images (512x512x149 resolution) for ZIN PH 281/16. Data acquired from CT scans were imported into the visualization software Amira 6.3.0 (FEI-VSG Company) and manually segmented. The resulting 3D models have the voxel size of 0.313x0.313x0.3 for ZIN PH 1/16 and 0.625x0.625x0.8 for ZIN PH 281/16. Measurements on the 3D models were performed using Amira 6.3.0 and MeshLab (Cignoni et al., 2008). The CT scan data and 3D models are available upon request from the first author.
Description of the holotype of
Bissektipelta archibaldi ZIN PH 1/16 (Figs. 1-9)
General comments. The braincase of Bissektipelta is highly ossified, and the bones of the skull roof are completely fused to it. Most sutures were obliterated. We do not support previous assumptions about incompletely ossified portions of some elements in the holotype (e.g., basal tubera, right basipterygoid process, occipital condyle, and the distal tip of the paroccipital process; Averianov, 2002) and regard those as preservational artifacts. These structures are variably preserved in the three studied braincases (notably, also in the smaller specimen ZIN PH 281/16) and are frequently broken off. The braincase is non-pneumatic. CT scans show that no internal pneumatic structures are present. Externally, there is neither the medial pharyngeal recess on the ventral surface of the basicranium nor a well-defined anterior/lateral pneumatic recess on the lateral surface of the parabasisphenoid.
Skull roof. The preserved skull roof has a relatively flat dorsal surface (Fig. 1A, B). Sutures are completely obliterated and are not evident either on the specimen's surface or in the CT images. General observations suggest that ZIN PH 1/16 preserves the posterior portion of the skull roof that corresponds to the frontoparietal region of taxa with known sutural relationships (e.g., Pinacosaurus, Maryañska, 1977; Godefroit et al., 1999; Hill et al., 2003; Kunbarrasaurus, Leahey et al., 2015; Ce-darpelta, Carpenter et al., 2001; "ZhongyuansaurusXu et al., 2007, = Gobisaurus in Arbour and Currie, 2016: Fig. 6D). A truncated Y-shaped groove that separates
three polygonal areas of remodeled bone (= caputegu-lae; Blows, 2001; Arbour and Currie, 2013a) is present. The resulting areas are identified here as the paired pos-terolateral nuchal caputegulae (nuca, Fig. 1B) and central parietal caputegulum (paca, Fig. 1B) using the terminology of Arbour and Currie (2013a). Each groove terminates in a pronounced pit; a small offshoot of the left groove is present and is directed anteromedially from the corresponding pit. The CT data for ZIN PH 1/16 shows that these grooves, paired pits, and the skull roof surface are pierced by numerous vascular foramina that connect through canals with the endocranial cavity and the lateral braincase wall. The left branch of the Y-shaped groove interrupts its course for one millimeter, and there is a short contact between the left nuchal and the central parietal caputegulae. The skull roof surface of ZIN PH 1/16 was remodeled, but it is uncertain if osteo-dermal ossifications were involved in that process. According to the hypothesis of Vickaryous et al. (2001a), "the superficial furrows that divide the cranium.. .represent the areas of coosification between adjacent cephalic osteoderms". The presence of the Y-shaped groove thus implies that the osteoderms are preserved and co-ossified with the skull roof in ZIN PH 1/16. There is no frontoparietal depression. The posterior edge of the skull roof is broken off. The broken lateral edges of the skull roof overhang the adductor cavities, and there are no traces of the supratemporal fenestrae.
The boundaries between the skull roof and brain-case are partly recognized on the preserved right par-occipital process in the occipital view (Fig. 1E), and are inferred on the lateral surface of the specimen based on the position of small vascular foramina that frequently lie near the border between the skull roof and brain-case (Galton, 1988; Galton and Knoll, 2006; Fig. 2A). The pattern of facets on the skull roof in the referred specimen ZIN PH 281/16 supports this reconstruction of the boundaries in the holotype. The parietal has two posterolateral processes that are sutured ventrolaterally to the dorsal surface of the paroccipital processes and medially to the supraoccipital (the latter contact is hard to trace; Fig. 1E). The posterolateral processes are an-teroposteriorly thin and oriented almost perpendicular to the sagittal plane of the skull. The posterior surface of the posterolateral processes is slightly posteroventrally-anterodorsally inclined. On the lateral aspect of ZIN PH 1/16, the skull roof appears to form an almost horizontal, slightly posteroventrally inclined contact with the braincase posterior to the capitate process of the lateros-phenoid and a gently anteroventrally inclined contact anteriorly (Fig. 2A). Posteriorly in lateral view, the parietal roofs a small vascular recess (nvr, Fig. 2A, B) and forms the dorsomedial wall of the adductor cavity. Here the skull roof reaches its greatest dorsoventral thickness of 21 millimeters.
oc
Fig. 1. ZIN PH 1/16, holotype of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. Photographs and corresponding CT-based models in dorsal (A, B), ventral (C, D), and occipital (E, F) views. Scale bars each equal 1 cm. Abbreviations: bo, basioccipital; bofe, basioccipital fenestra; bpt, basipterygoid process; bt, basal tuber; CN II — XII, cranial nerve foramina; fm, foramen magnum; fr, frontal; fvOC, foramen for orbitocerebral vein; fvSo, foramen for supraoccipital vein; MF, metotic foramen; nuca, nuchal caputegulum; nvf, neurovascular foramen; nvp, neurovascular pit; oc, occipital condyle; olff (CN I), olfactory fenestra; oto, otoccipital; p, parietal; paca, parietal caputegulum; pbsro-ios, fused parabasisphenoid rostrum and interorbital septum; pop, paroccipital process; proaf, proatlas facet; ptf, posttemporal fenestra; so, supraoccipital; tencr, tentorial crest.
Ventral surface of the basicranium. The basioccipi-tal and the parabasisphenoid meet at an angle of approximately 90o in ZIN PH 1/16; the suture between these bones is evident in lateral and ventral views (Figs. 1D, 2). Overall the basioccipital is massive and robust. The neck of the occipital condyle is barely defined. The ventral surface of the basioccipital is posteroventrally oriented, concave, and broad; it is slightly wider lateromedially than the corresponding surface of the parabasisphenoid. The basal tubera (= sphenoccipital tubera in Kurza-nov and Tumanova [1978] and Tumanova [1987]) are rounded, anteroposteriorly thin, and project laterally (bt, Fig. 1D, F). The basioccipital fenestra (bofe, Fig. 1D) is present as a distinct blind fissure on the ventral surface between the basal tubera. CT data show that two small, presumably vascular canals extend from it anteriorly and posteriorly inside the bone and gradually disappear in the trabeculae. The basioccipital fenestra is present in the same location ventral to the occipital condyle in present-day crocodylians; a small vein traverses this foramen (Owen, 1850).
The parabasisphenoid has a triangular, anteroven-trally oriented ventral surface (Fig. 1C, D). The surface between the basipterygoid processes is mediolaterally wider and gradually tapers anteriorly. The left basiptery-goid process is slightly incomplete (bpt, Fig. 1D). The basipterygoid process is a knob that projects ventro-laterally. It is oval in cross-section, with the longer axis directed anteriorly. Its anteroposterior length is nearly twice the mediolateral width at its base. The surface between the basipterygoid processes is relatively flat; there is a shallow depression on each side close to the base of the process. Only the base of the fused parabasisphenoid rostrum (= cultriform process) and the ossified/calcified interorbital septum is preserved. It is situated anterior to the basipterygoid processes (pbsro-ios, Fig. 1D). The base of the fused parabasisphenoid rostrum-interorbital septum extends obliquely anteriorly to the spheneth-moidal complex, where it merges with the septum that separated the olfactory bulbs (= mesethmoid in Miyas-hita et al. [2011]; Figs. 1D, 2E). Regarding the preserved part, the base of these elements is slightly transversally constricted at its mid-length and then expands anteriorly. On each side of the fused parabasisphenoid rostrum-interorbital septum are longitudinal depressions (possibly for the sphenopalatine artery; gaSP, Fig. 2D). A pronounced ridge ventral to the foramen for the optic cranial nerve (CN II) delimits the course of the longitudinal depression dorsally. No sutures in the region of the sphenethmoidal complex are discernable.
Occipital surface. The occipital surface is inclined at the angle of about 125o to the dorsal surface of the skull (Fig. 2A). When the specimen is held such that its skull roof surface is oriented horizontally, the occipital condyle is directed posteroventrally and barely projects
beyond the occipital plane. The articular surface of the condyle is crescent-shaped and transversely elongated (lateromedial length nearly 1.5 times larger its dorsoven-tral depth; Fig. 1E, F). The articular surface of the con-dyle is slightly eroded. The suture with the otoccipital is visible on the right lateral and posterior surfaces of the condyle (Figs. 1E; 2A); it indicates that the otoccipitals formed the dorsolateral corners of the occipital condyle. The posterior surface of the basioccipital ventral to the condyle is notably arched dorsally and overall faces pos-teroventrally (Fig. 2).
The foramen magnum is nearly circular. Its lateral and dorsal margins are formed by the otoccipitals; the supraoccipital appears to be excluded from the dorsal margin. Paired triangular surfaces (proatlas facets) project from dorsolateral corners of the foramen magnum (proaf, Fig. 1F). They merge medially and form a dorsal shelf over the foramen magnum. The proatlas facets overhang rounded notches that are sometimes interpreted as the path of the first spinal nerve (Kurzanov and Tumanova, 1978; Parish and Barrett, 2004). In addition, or as an alternative hypothesis, these sulci can correspond to the route of a venous vessel that branches off from the longitudinal venous sinus or its posterior expansion (occipital sinus) at the foramen magnum and courses ventrolaterally (Porter, 2015). Just dorsal to the proatlas facets, there are paired small foramina with associated grooves. These foramina pierce the occipital surface of the braincase directly to the endocranial cavity and likely transmitted small supraoccipital veins (fvSo, Fig. 1F). A venous foramen in a similar position above the foramen magnum was noted for "Amtosaurus magnus" (Kurzanov and Tumanova, 1978). Medial to these vascular foramina, on the assumed posterior surface of the supraoccipital, there is the base of the sagittal nuchal crest; dorsally, this surface is obscured by damage.
Paired rounded posttemporal fenestrae are present lateral to the sagittal nuchal crest (ptf, Fig. 1E, F). In general, the posttemporal fenestrae appear to lie near the contact of the parietal, supraoccipital, and otoccipital, but the precise sutural pattern is entirely obscured on the left side and is not clear on the right. The presumed parietal-otoccipital suture is situated at the ventrolateral margin of the posttemporal fenestra; thus, the ventral margin of the posttemporal fenestra is likely formed by the paroccipital process of the otoccipital, and its dorso-lateral margin by the parietal. It is likely that the supra-occipital contributed to the margin of the fenestra medially; alternatively, the medial margin of the fenestra may have been formed by the otoccipital and the parietal. The posttemporal fenestra pierces anteriorly into a small recess on either side of ZIN PH 1/16. This recess is evident in lateral view (nvr+g, Fig. 2A, B); it lies dorsal to the paroccipital process and medial to the adductor cavity. A notable groove is present at the anterior margin of the
Fig. 2. ZIN PH 1/16, holotype of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. Photographs and corresponding CT-based models in right lateral (A, B), left lateral (C, D), and oblique left lateral (E, F) views. Scale bars each equal 1 cm. Abbreviations: bo, basioccipital; bpt, basipterygoid process; bt, basal tuber; ca+vSO, canal for supraorbital artery and vein; ci, crista interfenestralis; CN II — XII, cranial nerve foramina; CN III / aOr, foramen for oculomotor nerve or orbital artery; CR+FO, columellar recess and fenestra ovalis; crp, crista prootica; fa+vSO, foramen for supraorbital artery and vein; faCC, foramen for cerebral carotid artery; faSP, foramen for sphenopalatine artery; fr, frontal; fvOC, foramen for orbitocerebral vein; gaSP, groove for sphenopalatine artery; ls, laterosphenoid; meth, mesethmoid; MF, metotic foramen; nvg, neurovascular groove; nvr+g, neurovascular recess and groove; olff (CN I), olfactory fenestra; ors, orbitosphenoid; oto, otoc-cipital; p, parietal; pbs, parabasisphenoid; pbsro-ios, fused parabasisphenoid rostrum and interorbital septum; pop, paroccipital process; pro, prootic; r, ridge; speth, sphenethmoid.
recess, suggesting the course of a blood vessel along the dorsomedial wall of the adductor cavity. Both the walls of the recess and the anterior groove are pierced by numerous small vascular foramina.
The preserved right paroccipital process extends laterally and slightly posteriorly and is incomplete distally (pop, Fig. 1). It is anteroposteriorly thin at its distal end and thick and robust at its base. The process is relatively narrow dorsoventrally; its depth equals the height of the foramen magnum. Two blunt ridges curve dorso-laterally and converge to form the ventral margin of the paroccipital process. The ventral margin of the paroc-cipital process is slightly arched dorsally and is nearly at the same level as the ventral border of the foramen magnum. Dorsally, the process is sutured to the skull roof. There is a small but pronounced depression at the posterior surface of the paroccipital process.
Lateral braincase wall. The elements forming the lateral wall of the braincase are fused (e.g., the sphen-ethmoidal complex, the orbitosphenoid, the lateros-phenoid, the parabasisphenoid, the prootic, and the otoccipital). No clear sutures can be observed, with the exception of the basioccipital-otoccipital suture on the condyle on the right side and the suture between the basioccipital and parabasisphenoid. All structures are paired, and the right and left sides of ZIN PH 1/16 have the same general structure and proportions. The lateral wall is penetrated by numerous neurovascular foramina (Fig. 2). These are clustered into two major groups and are relatively closely spaced within the cluster. The anterior group includes the foramina for CN II-VII and two primarily vascular foramina (for the cerebral carotid artery and the sphenopalatine artery and vein). The posterior group is situated ventral to the base of the par-occipital process and comprises the columellar recess/ fenestra ovalis, the metotic foramen, and the foramina for CN XII.
The two clusters of foramina are separated by a flattened strip of bone that extends ventrally between the basioccipital and the parabasisphenoid portion of the basal tuber. Dorsally, its posterior margin arches over the fenestra ovalis onto the ventral edge of the paroc-cipital process (crp, Fig. 2D). This structure corresponds to a poorly developed crista prootica (= otosphenoidal crest in Sampson and Witmer, 2007) that in diapsids separates the more anterior cranial nerve foramina from the posterior depression containing ear-related structures (fenestra ovalis plus metotic foramen). Generally in diapsids, the crista prootica arches posterodorsally from the parabasisphenoid lateral surface, just above the basipterygoid process. The crista prootica in Bissek-tipelta contacts ventrally the basal tubera instead of the basipterygoid proces. This is likely due to the highly divergent braincase structure of Bissektipelta (and other ankylosaurs) from the basic diapsid pattern, specifically
the posterior position of the basipterygoid processes close to the basal tubera.
The olfactory fenestrae are the only neurovascular foramina directed anteriorly instead of laterally (olff, Fig. 2D, F). They are paired and separated by a thick bony septum (= mesethmoid in Miyashita et al. [2011]). They are the largest neurovascular foramina and approach the foramen magnum in size. The olfactory fe-nestrae housed short paired olfactory bulbs and the ethmoid vessels, and they communicated directly with the olfactory region of the nasal cavity (Miyashita et al., 2011). The internal walls of the olfactory fenestrae are covered by numerous anteroposterior grooves, indicating that a large number of neurovascular bundles passed through them (nvg, Fig. 2E, F). The two separate cavities for the olfactory bulbs converge posteriorly into a single chamber that is separated from the rest of the endocra-nial cavity by a rounded constriction.
Only the base of the broken preorbital septum is preserved. The preorbital septum is a thin transversal bony lamina that separates the nasal and orbital cavities in ankylosaurs; it was first named by Maryanska (1977) (= ectethmoid in Miyashita et al. [2011]; see the description of ZIN PH 2329/16 below). Between the base of the preorbital septum and the anterior cluster of neurovas-cular foramina, the surface of the braincase wall bears no foramina and has dorsoventral striations. The largest foramen among the anterior cluster is that for CN V; the opening for CN II is the second largest. The foramina for the cerebral carotid artery and for the sphenopala-tine vessels are prominent and nearly equal in size (faCC and faSP, Fig. 2F). The large recess of the ganglion of CN V has a triangular projection from its dorsal margin that separates the anteriorly directed groove for CN V1 (ophthalmic branch of the trigeminal nerve) from postero-ventrally directed grooves for CN V2+3 (maxillary and mandibular branches of the trigeminal nerve; see Holli-day and Witmer [2007]). The small foramen for CN VII lies in the same large recess with that for CN V and is separated by a small ridge from the latter. The foramen for CN II is separated from the more posterior foramina by a thick vertical strut of bone. A small groove on the ventral margin of the foramen for CN II possibly indicates the course of a small vessel (Fig. 2D, E). There is a prominent depression on the lateral braincase wall dorsal to the foramen for CN IV and anterior to the adductor cavity (the postocular shelf is not preserved here in ZIN PH 1/16; see the description of ZIN PH 2329/16 below). The depression is pierced by two foramina for the orbitocerebral vein and a series of smaller vascular openings (fvOC, Fig. 2F).
The columellar recess/fenestra ovalis (CR/FO), the metotic foramen (MF), and three external foramina for CN XII are closely spaced and situated in a single depression ventral to the paroccipital process. This de-
pression is bordered by the crista prootica anteriorly, the basal tuber ventrally, and the prominent blunt ridge posteriorly (r, Fig. 2D). The latter connects with the ventral margin of the paroccipital process so that the foramina for CN XII are not evident in posterior view (Fig. 1E, F). The external openings of the CR/FO and MF are almost equal in size and large. The crista interfenestralis (= ventral ramus of opisthotic in more basal archosaurs; e.g., Gower, 2002; Sobral et al., 2016) separates FO and MF (ci, Fig. 2B). It is a slightly anteroventrally inclined lamina of bone that is barely visible in posterior view. The three foramina for CN XII are almost vertically arranged posterior to MF. The posteriormost foramen is the largest of the three. The anteriormost foramen for CN XII is the smallest and lies below MF.
Endocranial surface. The complex endocranial surface can be anteroposteriorly subdivided into four main concave regions (olfactory and cerebral fossae, and two fossae anterior and posterior to the otic capsule) separated by convex crests (Fig. 3). The anterior part of the endocranial cavity in ZIN PH 1/16 corresponds the pos-teriormost portion of the olfactory region of the nasal cavity (distinguished by rugose walls with numerous neurovascular grooves) and paired cavities of the olfactory bulbs that merge posteriorly into the cavity for the olfactory tract (olfbc and olftc, Fig. 3). The olfactory tract cavity is constricted laterally by paired blunt crests, which emphasize the division between the olfactory region anteriorly and the cerebral fossa posteriorly.
The cerebral cavity is circumscribed by the blunt olfactory crest anteriorly and by the tentorial crest (sensu Sedlmayr [2002]) posteriorly on each side (olfcr and tencr, Fig. 3A). Several neurovascular structures pierce the surface of the cerebral fossa, including the foramen for CN IV and two conspicuous foramina for the orbito-cerebral vein (Fig. 3A). The surface of the cerebral cavity has a gently corrugated texture but lacks prominent vascular valleculae, indicating that the brain was not in close relationship to the endocranial wall and loosely fitted the cerebral cavity (Evans, 2005). The large transverse groove for CN II is offset anteroventrally and opens posteriorly into the cerebral cavity (Fig. 3B). Its dorsal margin forms a blunt crest that arches posterodorsally onto the lateral endocranial surface on each side and merges with the tentorial crest. This oblique crest marks the subdivision of the cerebral fossa into two smaller fossae, roughly corresponding to the cerebral hemispheres anteriorly and the optic lobes posteriorly. The ventral margin of the CN II groove forms a sharp crest that denotes the anterior dorsal limit of the hypophyseal cavity.
The cerebral cavity merges into the hypophyseal cavity ventrally (hypc, Fig. 3). The hypophyseal cavity is comparatively shallow, being half the depth of the cerebral cavity. Foramina for the cerebral carotid and sphenopalatine arteries and for CN III pierce its surface
(Fig. 3). The internal foramen for CN III is unexpectedly situated ventrally, well in the limits of the hypophyseal cavity. A groove connects the internal openings of the sphenopalatine artery and CN III, raising the possibility that the latter may actually be a vascular foramen, perhaps for a branch of the cerebral carotid/sphenopalatine artery (e.g., the orbital artery of extant birds) or for the orbital/hypophyseal vein that drains into the cavernous sinus (Bruner, 1907; Porter and Witmer, 2015; Porter and Witmer, 2016a). In that case, the actual CN III would leave the braincase through the dorsally situated foramen for CN IV, as in Euoplocephalus (Miyashita et al., 2011). We tentatively follow the initial description by Averianov (2002) and maintain a conservative interpretation of the foramen in question as for CN III.
The dorsum sellae bulges over the hypophyseal cavity dorsally. It has a short anterior triangular projection surrounded by two grooves medially. This projection is also evident in the referred specimen of Bissektipelta ZIN PH 281/16, in "Amtosaurus magnus" (Kurzanov and Tumanova, 1978), and is possibly present, though less developed, in several other Mongolian taxa (Averianov, 2002; Parish and Barrett, 2004). We regard these grooves as vascular impressions that indicate the course of posterior venous branches of the cavernous sinus (caudo-ventral cerebral veins) or, as an alternative hypothesis, the course of the caudal encephalic arteries (Sedlmayr, 2002; Porter, 2015; Porter et al., 2016). Posteriorly to the dorsum sellae, the floor of the endocranial surface is essentially flat.
The tentorial crest is prominent; ventrally, it is confluent with the lateral aspect of the dorsum sellae, arches anterodorsally over the anterior margin of the foramen for CN V, and then curves posterodorsally and extends to the roof of the endocranial cavity (tencr, Fig. 3A). The internal opening for CN VI pierces the base of the tento-rial crest; the foramen for CN VII lies dorsolateral to it (Fig. 3B). The tentorial crest circumscribes a fossa dorsal to the foramen for CN V that was likely occupied by the cerebellum and a large venous vessel (middle cerebral vein; vMC, Fig. 3A). The latter opened externally through a series of foramina at the posterodorsal aspect of the fossa. The floccular (= auricular) fossa is very shallow.
The medial wall of the otic capsule is incompletely ossified (sc+ves in Fig. 3A). The amount of the exposure may have been exaggerated by postmortem fracture; however, both the referred specimens have largely medially open vestibules. The recesses for the vestibule, common crus, and lagena are medially open and confluent with the endocranial cavity. Paired unossified fossae with unfinished margins at the floor of the endocranial cavity mark the position of the lagenae (lagf, Fig. 3B). These fossae are comparatively large and probably contained additional structures such as supportive vascular plexus. A bifurcating groove extends posterodorsally
Fig. 3. ZIN PH 1/16, holotype of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. Parasagitally sectioned CT-based models showing left endocranial surface, in medial (A) and posteromedial (B) views. Scale bar equals 1 cm. Abbreviations: cerc, cerebral cavity; CN II — XII, cranial nerve foramina; CN III / aOr, foramen for oculomotor nerve or orbital artery; faCC, foramen for cerebral carotid artery; faSP, foramen for sphenopalatine artery; fvOC, foramen for orbitocerebral vein; hypc, hypophyseal cavity; lagf, lagenar fossa; MF, metotic foramen; olfbc, olfactory bulb cavity; olfcr, olfactory crest; olftc, olfactory tract cavity; sc+ves, cavities of semicircular canals and vestibule; tencr, tentorial crest; vMC, groove for middle cerebral vein.
from the internal foramen for CN VII and indicates the course of CN VIII (Fig. 3A); a similar reconstruction of this region was made for some ornithopods (Hopson, 1979; Sobral et al., 2012). The internal opening of the metotic foramen (MF) is just posterior to the otic capsule. The metotic foramen is undivided (see discussion in Rieppel [1985]; Gower and Weber, 1998; Sobral et al., 2012) and likely transmitted the perilymphatic sac, CN IX-XI, and the posterior cerebral vein (vagal vein in Sedlmayr [2002]). The wall between the vestibular recess and the MF is incised. This notch indicates the position of the incompletely ossified perilymphatic foramen that transmitted the perilymphatic sac from the otic capsule to the MF (Baird, 1960; Gower, 2002; Gower and Walker, 2002). Two larger internal foramina for CN XII pierce the endocranial wall posterior to the MF; a single small opening is just ventral to it. An extensive shallow depression with a deeper pit above these structures indicates the position of the occipital venous sinus (Fig. 3B; Sedlmayr, 2002; Witmer et al., 2008; Porter, 2015).
Endocast. The endocast of ZIN PH 1/16 generated from a CT scan data is complete, undistorted, and relatively detailed (Fig. 4; Table 2). It comprises casts of the endocranial cavity, cranial nerves, both endos-seous labyrinths, and vascular canals. The morphology of the inner ear and braincase vasculature of ZIN PH 1/16 are described in separate sections below. The brain of Bissektipelta loosely fitted the endocranial cavity as is common for many non-avian dinosaurs and for reptiles in general (Hopson, 1979; Witmer et al., 2008). Thus, the produced endocast is more a cast of the meninges (including endocranial venous sinuses) rather than the brain itself. Nevertheless, it appears to be a faithful inference of gross morphology of the brain as is suggested by recent research on extant archosaurs (Watanabe et al., 2019). Additionally, the endocranial vessels of various extant diapsids have a rather conservative pattern (Porter, 2015; see Vasculature and Fig. 9 below), and their disposition revealed on the endocast of Bissektipelta is a reliable proxy for recognition of major brain divisions.
olft ch
Fig. 4. ZIN PH 1/16, holotype of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. Cranial endocast with endosseous labyrinth of the inner ear in dorsal (A), ventral (B), and left lateral (C) views. Scale bar equals 1 cm. Abbreviations: aCC, cerebral carotid artery and vein; aSP, sphenopalatine artery and vein; cbl, cerebellum; ch, cerebral hemisphere; lab, endosseous labyrinth; MF, metotic foramen passage; olfb, olfactory bulb; olfc, olfactory cavity cast; olft, olfactory tract; optt, optic tectum of the midbrain; vn?, vomero-nasal bulbs?; II — XII, cranial nerves.
The endocranial cast is elongate and relatively straight, with low angles of cerebral and pontine flexures of about 30o; the cranial nerves and the fenestra ovalis of the labyrinth are correspondingly linearly arranged (Fig. 4C). The olfactory bulbs diverge anteriorly at an angle of approximately 80o from a short and broad olfactory tract (CN I; olfb and olft in Fig. 4A). The bulbs and the tract are nearly circular in cross-section. The olfactory ratio (the ratio of the greatest diameter of the olfactory bulb to the greatest diameter of the cerebral hemisphere regardless of their orientation; Zelenitsky et al., 2009) equals nearly 63-69 %. It is comparable to that of other ankylosaurs (see Discussion below) and suggests proportionally large olfactory bulbs in Bissektipelta and other ankylosaurs. Anteriorly, the endocast of each olfactory bulb terminates in a rounded expansion with wrinkled walls that corresponds to the olfactory region of the nasal cavity (olfc, Fig. 4A). The wrinkles most likely represent neurovascular bundles passing to and from the endocranial cavity, and part of them was visualized as the vascular olfactory plexus (olfP, Fig. 6; see "Vasculature" section below). Paired cavities within the ossified ethmoidal region, ventral to the olfactory tract, were segmented (vn?, Fig. 4A, B). A similar cavity was also found ventral to the broken mesethmoid in the referred specimen ZIN PH 281/16 (see Figs. 11-14). These struc-
tures lie below the olfactory complex of Bissektipelta and could correspond to the part of the vomeronasal bulb or vascular sinuses. These structures are reported for the first time in dinosaurs and require additional study to elucidate their nature.
The short olfactory tract posteriorly merges into a rounded cast of the cerebral cavity. The anterior part of the cavity's endocast reaches maximum dorsoventral depth and lateromedial breadth (Table 2) and corresponds to the cerebral hemispheres (ch, Fig. 4A). The cerebral hemispheres are relatively discrete on the endo-cast. The absence of a dorsal groove between the hemispheres indicates that the dorsal longitudinal venous sinus occupied the space above the latter.
The endocast of Bissektipelta lacks the dorsal dural peak present on endocasts of stegosaurs (Galton, 1988; Galton, 2001; Leahey et al., 2015: Fig. 10), most sauro-pods (Witmer et al., 2008), and some theropods (e.g., Sampson and Witmer, 2007). Dorsal expansions of dinosaur endocasts have been interpreted as an unossified gap plugged with cartilage in life (Hopson, 1979) or as corresponding to extensive dural sinuses that may have surrounded the pineal complex (Sampson and Witmer, 2007; Witmer et al., 2008). In Bissektipelta, the only structure of the endocast that may correspond to the pineal complex is a conspicuous median vessel (medVs,
Table 2. Endocast measurements of Bissektipelta archibaldi. All linear measurements in millimeters; volume in cm3
Parameter ZIN PH 1/16 ZIN PH 281/16
Whole endocast length 103 70
Endocast length without the cast of the olfactory region of the nasal cavity 85 -
Endocast volume (without vessels and nerves) 53
Endocast width across cerebral hemispheres 33 29
Olfactory bulb maximum cross-sectional diameter 15 -
Olfactory tract width 19.5 12
Pituiatary depth 19 18
Pituitary diameter 16.5 12.5
Fig. 7). Its canal pierces the skull roof all the way through to the endocast and connects to the anterior branching plexuses laterally (see Vasculature below). The median vessel emerges at the surface of the endocast just an-terodorsal to the inferred division between the cerebral hemispheres and the optic lobes (assessed by the position of the cerebrotectal venous sinus and the disposition of crests on the endocranial surface), at the level of the optic chiasm and the pituitary. This position broadly corresponds to that of the pineal complex in other di-apsids (e.g., Sphenodon; Dendy, 1911). Notably, the pineal complex, optic chiasm, and the neurohypophysis are all diencephalic derivates. However, the external pineal (parietal) foramen was lost early in archosauriform evolution (Hopson, 1979; character 63 in Nesbitt, 2011); extant birds have a pineal that lies internally within the braincase, adjacent to the skull roof (Ralph, 1970). Thus, we doubt that the median canal of Bissektipelta contained a pineal/parapineal organ that was exposed on the dorsal surface of the skull roof and consider the structure a vascular canal. However, noting its remarkable position, we hypothesize this canal enclosed vessels that may have been connected to pineal vasculature. The point of emergence of the median vessel from the endocast thus marks a possible position of the pineal complex in Bissektipelta.
The optic chiasm is located on the ventral surface of the endocast of Bissektipelta, below the cerebral hemispheres and just anterior to the hypophysis (Fig. 4). Each CN II leaves the braincase by a separate lateral foramen. The endocasts of the canals for CN II and of the optic chiasm form a single straight trunk that is oriented strictly perpendicular to the longitudinal axis of the braincase (Fig. 4B).
The complete cast of the hypophyseal (pituitary) fossa is present just posterior to the optic chiasm and ventral to the hemispheres (Fig. 4B, C). The pituitary projects vertically from the ventral surface of the en-docast. Overall, the pituitary cast is a tubular structure with an even diameter throughout; the stalk itself is not
expressed. It is relatively short dorsoventrally (its dorso-ventral depth equals nearly 19 mm and is half the depth of the cerebral cavity above it), broad, and nearly circular in cross-section (Table 2). The hypophyseal fossa apparently contained the infundibulum (hypophyseal stalk) and the hypophysis, which were likely surrounded by the cavernous venous sinus, as it in extant archosaurs (Neumeier and Lametschwandtner, 1994; Sedlmayr, 2002; Porter et al., 2016; Porter and Witmer, 2016a). The hypophyseal fossa of Bissektipelta was well vascularized. Large cerebral carotid arteries entered the hypophyseal cavity transversely as in most other ankylosaurs (e.g., Paulina-Carabajal et al., 2018); the sphenopalatine arteries branched off of them and left the hypophyseal cavity slightly anteriorly (a+vCC and a+vSP in Fig. 4B, C). The cerebral carotid and sphenopalatine veins that drained the cavernous sinus and the orbit/palate apparently shared canals with similarly named arteries. The stalk of CN III appears roughly at the mid-height of the endo-cast of the hypophyseal cavity, which occupies an unusually ventral position compared to those on most other dinosaur endocasts (Fig. 4C). A swelling on the lateral surface of the pituitary endocast connects the CN III trunk with the sphenopalatine artery endocast, which, combined with its low position, possibly indicates that the former represented a vessel (e.g., orbital artery and vein) rather than a nerve.
The optic lobes of the midbrain (optt, Fig. 4C) are not directly discernable on the endocast of ZIN PH 1/16 as they were likely overlain by sizable dural venous sinuses. The approximate position of the midbrain could be determined through the disposition of major encephalic vessels and general topographic cues of the diapsid brain (reviewed by Hopson [1979], Witmer et al. [2008], and others). The optic tectum of the midbrain in Bissektipelta apparently laid between the cerebral hemispheres anteriorly and the cerebellum posteriorly (Fig. 4C); thus, the brain had a linear arrangement that is similar to that in extant crocodiles, plesiomorphic for dinosaurs in general, and characteristic of many ornithischians in par-
ticular (Hopson, 1979; Balanoff and Bever, 2017). The endocranial cast of Bissektipelta is slightly constricted mediolaterally at the level of the optic tectum; its dorsal outline smoothly arches posteroventrally in lateral view. The short trunk of CN IV projects anterolaterally and slightly ventrally from the endocast above the pituitary (Fig. 4C). If we assume that the canal for CN III housed vascular structures rather than the actual cranial nerve, CN III must have left the braincase through the canal for CN IV.
The cerebellum is not distinctly expressed, and there is no prominent flocculus on the endocast of ZIN PH 1/16 (cbl, Fig. 4A, C). The region of the endocast that corresponds to the cerebellum is posterior to a groove reflecting the position of the tentorial crest. The cerebellum was circumscribed by extensive dural vessels, e.g., the middle cerebral vein anterodorsally and the longitudinal sinus (torcular Herophili part) dorsally (Figs. 6A, 7A, 9A).
Part of the endocast corresponding to the medulla oblongata is nearly as broad mediolaterally as it is anteroposterior^ long. The structure of the medulla ob-longata is obscured by extensive occipital venous sinus (sOc, Fig. 7A). The ventral surface of the brainstem is essentially flat and straight in lateral view; it is only slightly notched behind the endocast of the hypophyseal fossa anteriorly (Fig. 4B, C).
The single large trunk of CN V expands shortly after its emergence from the lateroventral surface of the endocast and superficially subdivides into three lobes (Fig. 4C). This expansion of CN V endocast likely corresponds to the Gasserian ganglion, and the three lobes reflect its main branches — the ophthalmic (CN VI), maxillary (CN VII), and mandibular (CN VIII) nerves (see Holliday and Witmer [2007] for a survey of the diapsid condition). The middle cerebral vein exited the brain-case together with CN V (vMC, Fig. 9). The endocast of CN VI extends anteroventrally and slightly laterally from the ventral surface of the brainstem. It comes off at the level of CN V, passes by the hypophyseal cavity, and exits the braincase through a separate foramen anterior to CN V (Figs. 2, 3, 4). The trunk of CN VII emerges between the endocasts of CN V and the inner ear and parallels the course of CN V. CN VIII was not digitally rendered; however, a groove at the endocranial surface of ZIN PH 1/16 that extends posterodorsally from the internal foramen of CN VII into the inner ear recess, just below the ampullary spaces, corresponds to the course of CN VIII (Fig. 3A). CN IX and CN X share the same exit via the metotic passage. The endocast of the MF is directly posterior to that of the inner ear and is relatively large (comparable to the endocast of CN II and slightly smaller than that of CN V). Three trunks of CN XII are evenly spaced posterior to the MF endocast; the anteri-ormost trunk is the smallest and lies adjacent to the MF.
Inner ear. The endosseous labyrinth of the inner ear was digitally reconstructed for both sides of ZIN PH 1/16 (Figs. 4, 5). The endosseous labyrinth is the endocast of inner skull cavities that carried the endolymphat-ic (otic or membranous) labyrinth surrounded by the perilymphatic (periotic) labyrinth (Baird, 1960; Witmer et al., 2008). Part of the perilymphatic labyrinth associated with semicircular canals is uniform among reptiles and closely matches the semicircular ducts of the endolymphatic labyrinth in shape. The lower part of the perilymphatic labyrinth that surrounds the saccule and the cochlear duct (lagena) of the endolymphatic system has a more complex structure that obscures the form of the endolymphatic labyrinth (Baird, 1960). The endosseous labyrinths of Bissektipelta, as well as those of other dinosaurs, reflect the structure of both the endolymphatic and perilymphatic systems as a whole. The perilymphatic labyrinth of Bissektipelta has an extracapsular portion (perilymphatic sac) that extends posteromedi-ally into the undivided metotic passage (MF) to participate in a compensatory secondary tympanic membrane (pls, Figs. 4B, 5B). This is a common condition for many diapsids including Sphenodon, basal archosaurs, and dinosaurs (Baird, 1960; Gower, 2002; Gower and Walker, 2002; Witmer et al., 2008). The position of extracapsular portion of the perilymphatic sac is marked by a notch between the vestibular recess of the inner ear cavity and the MF and was digitally visualized as part of the endosseous labyrinth (pls, Fig. 5B, C).
The endosseous labyrinths from both sides of ZIN PH 1/16 are undistorted and symmetrical. The medial aspects of both labyrinths are incomplete due to incomplete endocranial ossification of the otic capsules (Fig. 5C). The semicircular canals are robust. The am-pullar regions are not discrete and are present as expansions at proximal ends of the canals. Each semicircular canal lies in a single plane and does not curve beyond its limits. The anterior canal is the tallest and the largest of the three; it is roughly circular in shape (asc, Fig. 5B). The angle between the anterior and posterior canals equals approximately 90o. The posterior semicircular canal has a marked elliptical shape and is relatively low (psc, Fig. 5B); the anterior canal is one and a half times taller than the posterior one. The common crus is low (nearly equals the depth of the posterior canal) and broad (nearly twice the average canal breadth) (crc, Fig. 5). The lateral semicircular canal is ovoid in shape and appears to be equal to or only slightly smaller than the posterior canal (lsc, Fig. 5D). The utricular and sac-cular compartments of the endolymphatic labyrinth are not apparent as they were laterally covered by the peri-otic cistern of the perilymphatic labyrinth (as in other diapsids; Baird, 1960).
Below the level of semicircular canals, the endosse-ous labyrinth is markedly constricted anteroposteriorly.
Fig. 5. ZIN PH 1/16, holotype of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. Digital reconstruction of the endos-seous labyrinth of the left inner ear in anterior (A), lateral (B), medial (C), and oblique dorsolateral (D) views. Scale bar equals 1 cm. Abbreviations: asc, anterior semicircular canal; cochd, endosseous cochlear duct; CR, columellar recess; crc, crus communis; CR+FO, external openning of columellar recess leading to fenestra ovalis; lsc, lateral semicircular canal; pls, perilymphatic sac; psc, posterior semicircular canal.
This point is level with the columellar recess and the fenestra ovalis and marks the approximate line of division of the endosseous cochlear duct (lagena). The cochlear duct curves ventromedially below the vestibular portion of the labyrinth (cochd, Fig. 5A). Large unossified spaces below the endocranial cavity on both sides of ZIN PH 1/16 (lagf, Fig. 3B) were segmented as parts of the inner ear labyrinth. Although these structures are natural, symmetric, and observed in all studied specimens, we doubt that the actual cochlea was that elongate in Bissek-tipelta and curved below the endocast as in birds (e.g., Witmer et al., 2008). We suggest that these spaces could have contained either enlarged outgrowths of the peri-lymphatic sac and/or supportive neurovascular tissues of the inner ear. Overall, they just might have been plugged
with cartilage. Accounting for the uncertainty regarding the actual extent of the cochlear duct in ZIN PH 1/16, we perform two sets of measurements of its length (Table 3): a conservative assessment that does not include the medioventral portion of the cochlear endocast and an extended assessment that accounts for a complete model length. The presence of the extracapsular portion of the perilymphatic sac in Bissektipelta (which is the continuation of the scala tympani that encircles the la-gena medially in archosaurs; see Baird, 1960: Fig. 2) suggests that small portions of the perilymphatic labyrinth bulged out intracranially and, thus, were not visualized in the model. According to our approximate estimates, the length of the endosseous cochlear duct equals nearly 10-11 mm under a conservative measurement and 11-
Table 3. Endosseous labyrinth measurements and hearing properties of Bissektipelta archibaldi. The best frequency of hearing and the high-frequency hearing limit are calculated based on the equations from Gleich et al. (2005). We assume that the basilar papilla represents only two thirds of the cochlear duct length as in Gleich et al. (2005). For ZIN PH 1/16, two types of measurements were conducted — a more conservative approach (when a straighter line through the cochlea was measured) and an extended approach (when a strongly ventromedially curved line through the cochlea was measured). A single set of cochlear dimensions was taken from ZIN PH 281/16. All linear measurements in millimeters, hearing frequencies in hertz
Parameter ZIN PH 1/16 ZIN PH 281/16
Left labyrinth cochlear duct length, conservative 10.8
Left labyrinth cochlear duct length, extended 13.9 13.1
Right labyrinth cochlear duct length, conservative 10.1
Right labyrinth cochlear duct length, extended 11.6 14.4
Mean cochlear duct length for both labyrinths, conservative 10.45
Mean cochlear duct length for both labyrinths, extended 12.75 13.75
Best frequency of hearing, conservative 1002
Best frequency of hearing, extended 682 576
High-frequency hearing limit, conservative 2889
High-frequency hearing limit, extended 2299 2105
14 mm under an extended assessment, which amounts to 38-41 % and 41-53 % of the overall height of the en-dosseous labyrinth (the height of the vestibular part is around 15-16 mm). Thus, the lagena of Bissektipelta was moderately elongate.
A long canal extends laterally from the cochlear duct of ZIN PH 1/16 (CR, Fig. 5A). It has two parallel oblique sharp margins along its sides. We hypothesize that this structure represents the stapedial recess partly enclosed in bone due to extensive ossification of the lateral wall of the braincase in ZIN PH 1/16. In dorsal and anterior views, the distal part of the recess delimited by the aforementioned margins resembles the shape of the oblique stapedial footplate (Fig. 5A, D). Thus, the actual fenestra ovalis was likely displaced internally from the lateral surface of the braincase.
Vasculature. The CT data allowed digital reconstruction of a complex pattern of blood vessels in the ho-lotype of Bissektipelta archibaldi (Figs. 6-9). Endocranial vasculature and the system of vessels piercing the skull roof and lateral braincase wall has been reconstructed for Bissektipelta based on relative osteological correlates such as grooves and canals within bone (Witmer, 1995; Porter, 2015). Major vessels that are external to the braincase and did not leave direct bony features are only briefly mentioned here and are discussed later (see Discussion). As it is often hard to discriminate which component (arterial/venous) is prevalent in a given os-teological structure (save for well-known features such as the cerebral carotid canal predominated by the arterial component or grooves for the dural venous sinuses at the endocranial surface; see Porter [2015]), we have not distinguished between the types of blood vessels that pierced the skull roof and the braincase wall in our model. However, many of them are considered mainly or exclusively venous as encephalic arteries form a closed network around the brain under the dura matter and do not communicate with the orbital and temporal vessels (Sedlmayr, 2002; Almeida and Campos, 2010, 2011), with the exception of the ethmoid artery that communicates anteriorly with the supraorbital artery to form the nasal artery (Porter et al., 2016; Porter and Witmer, 2016a).
In extant diapsids, the main artery that supplies the braincase is the internal carotid artery and two of its branches — the cerebral carotid and the stapedial arteries (Porter and Witmer, 2015, 2016; Porter et al., 2016) (aIC, aCC, and aST in Fig. 8). In Bissektipelta, each cerebral carotid artery enters almost at the floor of the hypophyseal cavity (a+vCC, Fig. 6A). Small vessels branching off of the cerebral carotid curve anteroven-trally along the floor of the hypophyseal cavity (Fig. 4B). These small lobose vessels, though visualized as parts of the cerebral carotid artery endocast, possibly represent ventral parts of the cavernous sinus that drains into the
cerebral carotid vein (compare Fig. 4B with Neumeier and Lametschwandtner, 1994: Fig. 15). The latter vein shares the canal with the cerebral carotid artery; thus, both vessels are represented by a single trunk in the endocast of ZIN PH 1/16. The cerebral carotid arteries were likely connected medially because extant birds and crocodylians show anastomizing vessels/plexuses in the posteroventral region of the hypophyseal cavity (Sedlmayr, 2002; Porter et al., 2016; Porter and Witmer, 2016a). A horizontal swelling at the posterior surface of the pituitary endocast of ZIN PH 1/16, between cerebral carotid arteries, possibly corresponds to the intercarotid anastomosis (Fig. 4B). Just anterior to its entrance into the hypophyseal cavity, the cerebral carotid artery gives off the sphenopalatine artery (a+vSP, Fig. 6A). It is a distinct but smaller-caliber vessel compared to the cerebral carotid. A possible anterior course of the sphenopalatine artery is marked by a notch and depression on each side of the parabasisphenoid (gaSP in Fig. 2D, aSP in Fig. 8). The artery courses anterodorsally into the nasal region. A similar route of the sphenopalatine artery is present in extant birds (Porter and Witmer, 2016a). The dorsal courses of the common encephalic artery (= cerebral carotid after branching off sphenopalatine artery) and its branches that ramify around the brain inside the endocranial cavity are hard to trace. Possible osteologi-cal correlates are paired grooves on the dorsum sellae that could correspond to the caudal encephalic artery (Fig. 3A).
Another branch of the internal carotid is the sta-pedial artery, which continues anteriorly through the temporal region as the temporoorbital artery and then divides into three main orbital vessels (supra-, infraorbital, and ophthalmotemporal arteries; Sedlmayr, 2002; Porter, 2015) (aST, aTO, aSO, aOpt + aIO in Fig. 8). Anterior to the supratemporal fossa, a large curved canal within the lateral wall of the braincase, dorsal to the foramina for CN II-IV, is interpreted as the passage for the supraorbital artery and vein (preserved on the left side of ZIN PH 1/16 and opened by fracture on the right) (ca+v and fa+vSO in Fig. 2B, F; a+vSO in Fig. 6A; aSo in Fig. 8; vSO in Fig. 9). These vessels accompany each other through their course over the anterior surface of the laterosphenoid and ventral surface of the frontal in extant archosaurs (Porter et al., 2016; Porter and Witmer, 2016a). The supraorbital vessels course external to the bone surface in extant taxa; however, some dinosaurs with heavily ossified skulls (e.g., pachycephalo-saurids) show evidence for the bony enclosure of their branches into canals (Porter, 2015). The same is likely true for ZIN PH 1/16. The supraorbital artery/vein canal communicates via small-caliber vascular canals with the anterior branching plexus of the cranial roof dorsally and with endocranial vessels medially and posteromedi-ally (Figs. 6, 7, 9). The latter corresponds to numerous
Fig. 6. ZIN PH 1/16, holotype of Bissektipelta archibaldifrom the Bissekty Formation (Turonian), Uzbekistan. CT-based models showing braincase vasculature in left lateral view; endocast with surrounding vessels (A), semitransparent view of the braincase showing vessels within the skull roof and lateral braincase wall (B), solid braincase (C). Scale bar equals 1 cm. Abbreviations: a+vCC, cerebral carotid artery and vein; a+vSO, supraorbital artery and vein; a+vSP, sphenopalatine artery and vein; ABP, anterior branching plexus; olfABP, olfactory part of the anterior branching plexus; olfP, olfactory plexus; PBP, posterior branching plexus; PBP-vCD, anastomotic vessel between the posterior branching plexus and the dorsal head vein; PbsVs, parabasisphenoid vasculature; sOc, occipital venous sinus; sP, parietal venous sinus; vCD, dorsal head vein; vg, venous groove; vOC, orbitocerebral vein; vSo, supraoccipital vein; vTOc, transverso-occipital vein. Black arrow heads mark small anastomotic connections between main vascular elements.
Fig. 7. ZIN PH 1/16, holotype of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. CT-based models showing braincase vasculature in dorsal view; endocast with surrounding vessels (A), semitransparent view of the braincase showing vessels within the skull roof (B), solid braincase and skull roof (C). Scale bar equals 1 cm. Abbreviations: ABP, anterior branching plexus; anastABP, anastomotic connection between left and right anterior branching plexuses; medVs, medial vessel; olfABP, olfactory part of the anterior branching plexus; PBP, posterior branching plexus; sOc, occipital venous sinus; sP, parietal venous sinus; vCD, dorsal head vein; vOC, orbitocerebral vein; vSo, supraoccipital vein; vTOc, transverso-occipital vein. Black arrow heads mark small anastomotic connections between main vascular elements. White arrow heads mark connections between the dorsal head vein and the middle cerebral vein.
Fig. 8. ZIN PH 1/16, holotype of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. CT-based model of the braincase in right lateral view showing reconstructed pattern of arteries. Red vessels are reconstructed upon corresponding osteological correlates. Black vessels inferred from broader studies of diapsid vascular patterns. Not to scale. Abbreviations: aCC, cerebral carotid artery; aET, ethmoidal artery; aIC, internal carotid artery; aMan, mandibular artery; aNas, common nasal artery; aOc, occipital artery; aOpt+aIO, ophthalmotemporal and infraorbital arteries; aSO, supraorbital artery; aSP, sphenopalatine artery; aST, stapedial artery; aTO, temporoorbital artery; braSO, dorsal branches of supraorbital artery.
venous communications with endocranial dural veins (mainly, the dorsal longitudinal sinus with its tributaries and the cerebrotectal sinus, see Fig. 9). The supraorbital artery/vein canal also receives a number of small canals from the lateral surface of the braincase. The anterior course of the supraorbital vessels is marked by an antero-ventrally directed groove and the orbitonasal foramen in the preorbital septum (Fig. 8). The latter structure is broken in the holotype but preserved in the referred specimen ZIN PH 2329/16 (Fig. 15). The communication of the supraorbital and ethmoid vessels (situated dorsal to the olfactory tract and bulbs in extant taxa; Almeida and Campos, 2010, 2011; Porter et al., 2016; Porter and Wit-mer, 2016a) occurred through some of the small canals piercing the lateral wall of the braincase (Figs. 6A and 7A) and further anterior to the orbitonasal foramen (see Figs. 8-9). The latter communication of the supraorbital and ethmoid vessels gave rise to the nasal vessels that supplied and drained the nasal cavity.
The encephalic vessels seldom leave direct traces on the endocranial surface (with some notable exceptions; see Evans [2005]); however, their basic pattern appears to be rather conservative among known diapsids (Bruner, 1907; Dendy, 1909; Sedlmayr, 2002; Witmer et
al., 2008; Porter, 2015; Porter and Witmer, 2015, 2016; Porter et al., 2016). Major endocranial veins are recognized as swellings on the endocast surface that communicate with external vasculature via vascular or nervous canals (Hopson, 1979; Sampson and Witmer, 2007; Witmer et al., 2008; Porter, 2015). The latter are important landmarks that trace the course of the vessel. The digital endocast of the holotype of Bissektipelta allows recognition of several dural venous vessels/sinuses and their communications with external vasculature (Fig. 9).
The dorsal longitudinal sinus appears as a shallow but broad prominence on the top of the endocast that extends from the olfactory tract anteriorly to the level of the otic capsules posteriorly (Figs. 7, 9). Anterior to the olfactory tract, the dorsal longitudinal sinus apparently splits into a pair of vessels (olfactory veins in Dendy [1909]; ethmoid vein in Porter and Witmer [2016]; Porter et al., 2016) that overlaid the olfactory bulbs and continued forward to drain the olfactory cavity as nasal veins (vET + vNas in Fig. 9). A large number of neurovascular grooves are preserved around the olfactory bulbs/posterior portion of the olfactory region of the nasal cavity in ZIN PH 1/16 (Fig. 2F). These grooves indicate the presence of a vascular plexus around the ol-
olfP sLon
Fig. 9. ZIN PH 1/16, holotype of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. CT-based models of the cranial endocast and braincase in left lateral view showing reconstructed pattern of veins. A, encephalic veins that drain endocranial cavity, labeled on the endocast; B, semitransparent view of the braincase showing veins within the skull roof and lateral braincase wall (solid), encephalic veins (transparent), and their intercommunications; C, non-transparent view of the braincase showing external braincase veins (solid) and their parts within canals (transparent). Note controversial interpretation of veins at the supratemporal fossa; see Discussion for the preferred reconstruction. Blue vessels are reconstructed upon corresponding osteological correlates. Black vessels inferred from broader studies of diapsid vascular patterns. Not to scale. Abbreviations: ABP, anterior branching plexus; anastv, circum-occipital anastomotic loop between dorsal head vein, transverso-occipital vein, and parietal venous sinus; olfABP, olfactory part of the anterior branching plexus; olfP, olfactory plexus; PBP, posterior branching plexus; sCT, cerebrotectal sinus; sLon, dorsal longitudinal sinus; sOc, occipital venous sinus; sOc/vCP, occipital venous sinus/ posterior head vein; sP, parietal venous sinus; vCD, dorsal head vein; vET, ethmoidal vein; vMC/sT, middle cerebral vein and transverse venous sinus; vNas, common nasal vein; vOC, orbitocerebral vein; vOpt+vIO, ophthalmotemporal and infraorbital veins; vSC, superior cerebral veins; vSo, supraoccipital vein; vSO, supraorbital vein; vST+vTO, stapedial and temporoorbital veins; vTOc, transverso-occipital vein.
factory bulbs that drained into the ethmoid vein and the anterior branching plexus (olfP in Figs. 6-7, 9). In extant birds, the olfactory bulbs are surrounded by a dense venous plexus that eventually drains into the longitudinal sinus via the ethmoidal veins and/or the olfactory sinus and their anastomoses (Sedlmayr, 2002; Porter and Wit-mer, 2016a).
A conspicuous median vessel arises sagittally at the top of the endocast of Bissektipelta, dorsal to the cerebral hemispheres (medVs, Fig. 7). The vessel extends dor-sally within a canal inside the bone and opens through a foramen on the dorsal surface of the skull roof. Along the way, it issues left and right major branches as well as smaller branches in a slightly asymmetrical manner; each major branch communicates with a corresponding anterior branching plexus (Fig. 7A). The median vessel was apparently connected to the dorsal longitudinal sinus ventrally. Undoubtedly this structure was vascular (most likely, venous), as it commences from the endo-cast at the inferred position of the longitudinal sinus and connects to vascular branching plexuses. We are unaware of any similar dorsal extensions of the encephalic vessels in extant diapsids. As noted earlier in the description of the endocast, the position of the median vessel in Bissektipelta generally corresponds to the position of the pineal complex in extant lepidosaurs and birds. In extant diapsids, the pineal complex is well vascularized, supplied by branches of the posterior cerebral artery and drained by the dorsal longitudinal sinus (Dendy, 1909; Ralph, 1970). We doubt that the canal housed the pineal/ parapineal organ itself but hypothesize that the median vessel and its branches may represent dorsal continuations of the pineal complex vasculature.
The dorsal longitudinal sinus is most prominent posteriorly where it received the middle cerebral veins and appears as a broad triangle in the dorsal view (tor-cular Herophili; Dendy, 1909; Sedlmayr, 2002; sOc, Fig. 7A). Posterior to this, the dorsal longitudinal sinus likely bifurcated into two sinus-like posterior cerebral veins (vena cerebralis posterior, vena cephalica posterior, vagal vein, occipital venous sinus of different authors) (sOc, Fig. 7A; sOc/vCP, Fig. 9A). The posterior cerebral veins likely left the endocranial cavity through the foramen magnum, as in most extant diapsids (Bruner, 1907; Sedlmayr, 2002; Witmer et al., 2008; Porter and Witmer, 2015, 2016; Porter et al., 2016). However, Sphenodon shows an important variation of the course of the posterior cerebral vein, which leaves the endocra-nial cavity through the metotic foramen (Dendy, 1909). The same route for the posterior cerebral vein through the metotic foramen was reconstructed for basal croco-dylomorphs (Walker, 1990) and other pseudosuchians (Gower, 2002; Gower and Nesbitt, 2006; Sulej, 2010). Both pathways for the venous drainage (via the metotic foramen/foramen magnum) are likely traced in extant
crocodylians through their development (Dendy, 1909; Sedlmayr, 2002). For Bissektipelta, we imply that most, if not all, of the venous blood left the endocranial cavity posteriorly through the foramen magnum, with possible additional venous drainage through the metotic foramen via the posterior cerebral vein (sOc and sOc/vCP, Fig. 9A). Sobral et al. (2012) arrived at a similar conclusion regarding the pathway of the posterior cerebral vein in the ornithopod Dysalotosaurus.
A pair of small foramina (fvSo in Fig. 1) directly above the foramen magnum of Bissektipelta apparently transmitted small veins and accompanying arteries (su-praoccipital veins; vSo in Figs. 6A, 7A and 9A). In Sphen-odon, these vessels drain from the dura matter and the dorsal part of the occipital sinus extracranially through similarly distributed foramina (Dendy, 1909).
Along its course, the dorsal longitudinal sinus receives several transverse veins that drained lateral aspects of the endocranial cavity. We assume the presence of a number of superior cerebral veins that extended along the lateral aspects of the olfactory tract and the anterior cerebrum, dorsal to CN II, as was described for Sphenodon (venae cerebrales superiores; Dendy, 1909) (vSC, Fig. 9A). Additionally, the presence of corresponding but unidentified vessels dorsal to CN II was discussed for Caiman and reported for fossil endocasts (Hopson, 1979). The presence of superior cerebral veins in Bissektipelta is established by numerous small vascular canals that connected the anterior branching plexus and the canal for supraorbital vessels with endocranial dural veins (Fig. 7A). These veins joined the corresponding ethmoid vein/dorsal longitudinal sinus dorsally.
Posteriorly, at the level of CN IV, conspicuous swellings on the endocast and a pair of the orbitocerebral veins on each side indicate the course of a transverse venous sinus (sCT, Fig. 9A, B). The latter received confusing terminology in the literature: vena cerebri posterior in Hopson (1979); sphenotemporal sinus in Sedlmayr (2002); sphenoparietal sinus in Witmer et al. (2008); and cerebrotectal sinus in Porter et al. (2016) and Porter and Witmer (2016). In extant archosaurs, this venous sinus extends dorsally along the tentorial crest to join the dorsal longitudinal sinus and wedges in between the posterior region of the cerebrum and the optic tectum (Hopson, 1979; Sedlmayr, 2002). A series of veins on each side of the brain in the same region (venae begime-nales superiores) was described for Sphenodon (Dendy, 1909). We use the term "cerebrotectal sinus", as it clearly reflects the anatomical position of the vessel. In Bissek-tipelta, the cerebrotectal sinus extends transversally as a swelling on the posterior aspect of the cerebral endocast (compare Figs. 4C, 6A, and 9A). The orbitocerebral veins drain into the cerebrotectal sinus from the orbital cavity (vOC in Figs. 6A, 9B). The cerebrotectal sinus directly communicates via small vascular canals with the
middle cerebral vein and dorsal head vein/parietal sinus, the posterior branching plexus, and the canal for supraorbital vessels (Figs. 6, 9).
The succeeding large transverse tributary of the dorsal longitudinal sinus with complicated nomenclature is the middle cerebral vein (vena cerebralis media in Bruner [1907]; transverse sinus in Dendy [1909] and Porter and Witmer [2015]; recessus lateralis of longitudinal sinus in Hopson [1979]; rostral petrosal sinus in Sedlmayr [2002]; cerebellotectal sinus in Porter and Witmer [2016] and Porter et al. [2016]) (vMC, Fig. 9). In extant diapsids, it is a large vessel that extends between the optic tectum and cerebellum, just in front of the otic capsule. At the point of its divergence from the longitudinal sinus, the middle cerebral vein is sinus-like and broad, and thus its dorsal portion was designated the transverse, rostral petrosal, or cerebellotectal sinus (Dendy, 1909; Sedlmayr, 2002; Sampson and Witmer, 2007; Porter and Witmer, 2016a; Porter et al., 2016). Ventrally, the sinus drains into one or several smaller and more defined veins (= middle cerebral vein sensu stricto, e.g., Sampson and Witmer, 2007; trans-versotrigeminal vein of Porter and Witmer [2015]; rostral middle cerebral vein in Paulina-Carabajal et al. [2016]) that frequently pass through the trigeminal foramen ex-tracranially. We use the simpler term "middle cerebral vein" for both portions of the vessel (the transverse sinus and its continuations) in an effort to keep the terminology as concise as possible and to ensure compatibility with previous accounts on dinosaurian cranial vasculature (Sampson and Witmer, 2007; Witmer et al., 2008; Miyas-hita et al., 2011; Leahey et al., 2015; Paulina-Carabajal et al., 2016, and others).
In Bissektipelta, the middle cerebral vein/transverse sinus extends dorsally from the foramen for CN V as a bulge on the endocast surface, then arches posterodor-sally, parallel to the anterior semicircular canal, and finally joins the dorsal longitudinal sinus (Figs. 6A and 9). In Bissektipelta, there is no separate branch of the middle cerebral vein that passes extracranially in the lateral direction through its own canal (= rostral middle cerebral vein of some authors). Thus, the middle cerebral vein likely exited the braincase via the large foramen of CN V. Dorsally, a conspicuous posterodorsally curved swelling on the endocast marks the course of the middle cerebral vein. Here, the middle cerebral vein is laterally confluent with the dorsal longitudinal/occipital sinus (Fig. 9A). Three short vascular branches extend posterodorsally and laterally from the middle cerebral vein on each side of ZIN PH 1/16 (Fig. 7A). These vascular branches connect the middle cerebral vein with the external veins of the temporal and occipital regions of the skull (dorsal head vein, transversooccipital vein, parietal sinus; Figs. 6A, 7A, 9B). Additionally, a separate vessel extends from the anterior-most of the three described vascular branches and connects with the cerebrotectal sinus anteriorly (precisely,
with the dorsal orbitocerebral vein, Figs. 6A, 9B). As described, the course of the middle cerebral vein in Bissek-tipelta is consistent with observations on extant diapsids (Bruner, 1907; Dendy, 1909; Porter and Witmer, 2015; Porter and Witmer, 2016a; Porter et al., 2016) and various dinosaurs (see Discussion).
The dorsal head vein and the transverso-occipital vein (caudal middle cerebral vein of some authors) exit the braincase of Bissektipelta via separate foramina on the lateral (nvr+g, Fig. 2B; vCD, Fig. 6B, C) and occipital (ptf, Fig. 1; vTOc, Fig. 7B) surfaces of the skull, correspondingly. However, their endocast suggests that they either represent a single vessel or a continuous anasto-motic loop that extends from the temporal to the occipital region of the skull and maintains the connection with the middle cerebral vein/transverse sinus (Fig. 7A). In Bissektipelta, the groove passes anterior from the foramen of the dorsal head vein (nvr+g in Fig. 2B). This groove corresponds to the continuation of the dorsal head vein; we term this continuation as the parietal sinus (sP, Figs. 6B, 7A, 9B-C) following the terminology of extant squamates (Bruner, 1907; Porter and Witmer, 2015; see also Discussion).
Numerous small openings at the dorsal surface of the skull roof of Bissektipelta lead into the canals within bones that eventually converge ventrally (Fig. 7). This pattern of vascular canals is herein referred to as branching plexus. There are paired anterior and posterior branching plexuses that supplied and drained the skull roof and overlying dermis in Bissektipelta (ABP, Figs. 6-7 and 9). The anterior branching plexus can be subdivided into two parts: one part that lies above the olfactory bulb and the olfactory cavity and is connected ventrally to the ethmoid vessels (olfABP in Fig. 9B) and the other part that lies posteriorly and communicates with the supraorbital vessels ventrally and dural veins medially (ABP in Fig. 9B). Some parts of these canals likely transmitted small branches of the supraorbital artery that pierce the frontal and emerge onto the outer surface of the skull in extant birds (Porter and Witmer, 2016a) and some dinosaurs (Porter, 2015) in a similar way (braSO in Fig. 8). The posterior plexus is situated above the dorsal head vein (PBP in Figs. 6, 9); it is less distinct compared to the anterior plexus and was not visualized on the right side of ZIN PH 1/16. As previously described, small vascular canals integrate the anterior and posterior branching plexuses as well as various endo- and extracranial vessels into a single vascular network around the brain (see Discussion for physiological implications).
Description of ZIN PH 281/16 (Figs. 10-14)
General comments. ZIN PH 281/16 is exquisitely preserved, with fine features of the external and endocra-nial surfaces and clear sutures and facets. It appears to
be only slightly smaller compared to the holotype; most of its measurements are only 5-15 % smaller than those for ZIN PH 1/16 (Table 1). The braincase is externally and internally non-pneumatic, as is evident from the CT data.
Skull roof. ZIN PH 281/16 does not preserve bones of the skull roof and has slightly rugose fine facets on its dorsal surface (Figs. 10A, 11A). This indicates that the skull roof was not completely co-ossified with the braincase in this particular specimen. A lack of fusion between the skull roof and braincase is present in adults of Pinacosaurus (Maryanska, 1977; Tumanova, 1987) and Minotaurasaurus (Miles and Miles, 2009; Penkalski and Tumanova, 2017), and the two cranial components are strongly sutured in adult individuals of other anky-losaurs. The unfused skull roof in ZIN PH 281/16, along with open sutures between individual neurocranial elements, indicates that the specimen probably represents a somatically subadult individual.
Ventral surface of the basicranium. The ventral aspect of the specimen is formed by the basioccipital posteriorly and the parabasisphenoid anteriorly; the suture between these bones is clearly visible on both the lateral and ventral surfaces (Fig. 10C). Unlike in the holotype, the two bones join at an obtuse angle of approximately 120o. The ventral surface of the basioc-cipital is smoothly arched and bears a vascular foramen (basioccipital fenestra). The triangular ventral surface of the parabasisphenoid bears the base of the fused para-basisphenoid rostrum-interorbital septum anteriorly and small, bump-like basipterygoid processes (left one is broken off), which are offset posteriorly, close to the suture between the basioccipital and parabasisphenoid (Fig. 10D).
Occipital surface. The occipital surface of the specimen is formed by the supraoccipital, basioccipital, and paired otoccipitals; the sutures between these bones are easily recognized, unlike in the holotypic cranium and most other known ankylosaurs (Fig. 11C). The occipital surface forms the same angle of about 125o with the skull roof (inferred from the plane of corresponding facets) as in the holotype. The occipital condyle barely projects beyond the occipital plane. It has a more rounded shape compared to those of the holotype and ZIN PH 2329/16. The otoccipitals form the dorsolateral portions of the condyle. The otoccipital-basioccipital suture is evident on both sides of the specimen (Figs. 11C, 12). The suture extends onto the lateral and endocranial surfaces, where it gradually disappears toward the external and internal openings of the metotic foramen, respectively. The foramen magnum is bounded by the basioccipital ventrally and by the otoccipitals laterally and dorsally. The supraoccipital was probably excluded from the dorsal margin of the foramen magnum by a short dorsal contact between the otoccipitals; however, the latter is
not preserved. The supraoccipital-otoccipital suture is apparent dorsal to the foramen magnum and further anterolaterally where the supraoccipital reaches the prootic and probably the laterosphenoid on either side (Figs. 12A, 13A).
The supraoccipital bears a clear sagittal crest with two depressions on its sides (scr, Fig. 11D). These depressions could correspond to the ventral border of the posttemporal fenestra (ptf? in Fig. 11D) and the course of the transverso-occipital vein; they are, however, too close to the sagittal plane compared to the position of the posttemporal fenestra in the holotype (Fig. 1F). Paired canals within the paroccipital processes, just ventral to the contact with the parietal and lateral to the suture with the supraoccipital, almost certainly transmitted vascular elements. These canals could have transmitted some tributaries of the dorsal head/transverso-occipital veins or the occipital artery (vf in Figs. 11D, 12B, 13B). The latter canals are absent in the holotype ZIN PH 1/16; thus, the arrangement of vascular foramina on the occipital surface is variable among the specimens assigned to Bissektipelta. Small paired foramina for the supraoc-cipital vein are present at the suture between the supra-occipital and otoccipital (fvSo, Fig. 11D).
Lateral braincase wall. In general, the structure of the lateral wall of the braincase of ZIN PH 281/16, including the distribution of the neurovascular foramina, closely matches that of the holotype. The neurovascular foramina are grouped into anterior and posterior clusters divided by a flattened crista prootica.
The otoccipital forms most of the posterior aspect of the lateral wall of the braincase and encloses much of the posterior cluster of foramina: the undivided metotic foramen (MF), two or three external foramina of the CN XII (varying between the two sides of the specimen), and partly the fenestra ovalis (FO) and the columellar recess (Fig. 12). These openings are incised ventral to the broad base of the paroccipital process. A pair of grooves begins from the FO and MF and extends distally on the ventral surface of the paroccipital process. The openings for CN XII and the MF are completely enclosed by the otoccipital; the basioccipital is apparently excluded from the external ventral border of the MF (Fig. 12E). The MF is separated from the foramina for CN XII by a laminalike process of the otoccipital that descends anteroven-trally toward the basal tuber. The metotic foramen is separated from the anteriorly situated FO by an oblique crista interfenestralis (= ventral ramus of opisthotic in more basal archosauriforms). The crista interfenestralis is a minor process that is not visible in occipital view. The columellar recess and the FO are bounded by the prootic anteriorly and by the otoccipital posteriorly, which is a common condition for diapsids in general (e.g., Sobral and Müller, 2016). The columellar recess is more open laterally that in the holotype ZIN PH 1/16. It leads into
Fig. 10. ZIN PH 281/16, referred specimen of Bissektipelta archibaldifrom the Bissekty Formation (Turonian), Uzbekistan. Photographs and corresponding CT-based models in dorsal (A, B) and ventral (C, D) views. Scale bars each equal 1 cm. Sutures are represented by solid lines; possible sutures are represented by dashed lines. Abbreviations: bo, basioccipital; bpt, basipterygoid process; cap, capitate process; CN II — XII, cranial nerve foramina; fvOC, foramen for orbitocerebral vein; hypc, hypophyseal cavity; ls, laterosphenoid; MF, metotic foramen; ors, orbitosphenoid; oto, otoccipital; pbs, parabasisphenoid; pop, paroccipital process; pro, prootic; so, supraoccipital; speth, sphenethmoid; vf, vascular foramen.
the FO; the latter communicates medially with the vestibular recess of the inner ear via a foramen (Fig. 13C).
The prootic forms the posterior border of the foramen for CN V, encloses the foramen for CN VII, and partially bounds the columellar recesss/FO (Fig. 12A, C, E). It broadly adheres to the anterior surface of the par-occipital process, as in various archosauriforms except crocodylomorphs (character 105 in Nesbitt [2011]). The anterior contact of the prootic with the laterosphenoid is evident on both sides of the specimen. The prootic-supraoccipital contact is not clearly observable, and the ventral contacts of the prootic with the parabasisphenoid
are obliterated. The prootic forms a triangular projection that descends from the dorsal margin of the foramen for
CN V and partially subdivides it.
The suture between the laterosphenoid and prootic and a prominent capitate process (cap in Figs. 11B, 12B) mark the posterior extent of the laterosphenoid. The latter participates in the anterior margin of the foramen for CN V. It is likely that the laterosphenoid also encloses the orbitocerebral vein openings and the foramen for CN IV. A pair of foramina for the orbitocerebral veins is present on both laterosphenoids; additionally, a groove at the presumed laterosphenoid-frontal contact on both
Fig. 11. ZIN PH 281/16, referred specimen of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. Photographs and corresponding CT-based models in anterior (A, B) and posterior/occipital (C, D) views. Scale bars each equal 1 cm. Sutures are represented by solid lines; possible sutures are represented by dashed lines. Abbreviations: ameth, articular surface for mesethmoid; bo, basioccipital; bpt, basipterygoid process; bt, basal tuber; cap, capitate process; CN V, trigeminal cranial nerve foramen; oc, occipital condyle; cvn?, cavity for vomero-nasal bulb?; fCC, cerebral carotid artery and vein foramen; fm, foramen magnum; fvSo, supraoccipital vein foramen; ls, laterosphe-noid; oto, otoccipital; pbs, parabasisphenoid; pbsro-ios, fused parabasisphenoid rostrum and interorbital septum; pop, paroccipital process; proaf, proatlas facet; ptf?, posttemporal fenestra?; scr, sagittal crest; so, supraoccipital; speth, sphenethmoid; vf, vascular foramen.
sides of the specimen marks the course of a similar vascular element (fvOC in Fig. 12D). An internal vascular canal for the supraorbital vessels was reconstructed for the holotype at this region; the canal is absent in ZIN PH 281/16, indicating a lesser degree of ossification of the braincase wall. The capitate process is stout and bears a rounded head with an unfinished articular surface (Fig. 12A). The facet on its dorsal surface and round head indicate a synovial joint between the laterosphe-noid and the postorbital (Holliday and Witmer, 2008) that was nevertheless akinetic, as in extant crocodylians. The blunt crista antotica (Sampson and Witmer, 2007, and references therein; laterosphenoid buttress in Hol-liday and Witmer [2009]) descends from the capitate process and subdivides the orbital and adductor aspects of the external surface of the laterosphenoid (Fig. 12B,
D). On the left side of ZIN PH 281/16, a groove passes through the crista antotica. It likely indicates the course of the temporoorbital artery/vein (gTO in Fig. 12D).
Sutures cannot be distinguished between the preserved elements of the sphenethmoidal complex and between them and the parabasisphenoid. The medial septum that separated the olfactory bulbs is broken off in ZIN PH 281/16, but the preserved surface is symmetrical on both sides and most likely represents a facet (ameth, Fig. 11A). Thus, this medial septum was a separate element (mesethmoid in Miyashita et al. [2011]). It contacted the elements of the lateral wall of the braincase laterally, the parabasisphenoid ventrally, and the skull roof dorsally (based on the holotype that preserves both the medial septum and the skull roof) and was probably continuous anteriorly with the ossified nasal septum as
CR+FO CN VII bP*
bt
faCC
Fig. 12. ZIN PH 281/16, referred specimen of Bissektipelta archibaldifrom the Bissekty Formation (Turonian), Uzbekistan. Photographs and corresponding CT-based models with cranial endocast in right lateral (A, B) and oblique left lateral (C, D) views, with a close-up of the posterior cranial nerve foramina (E). Scale bars each equal 1 cm; E not to scale. Sutures are represented by solid lines; possible sutures are represented by dashed lines. Abbreviations: bo, basioccipital; bpt, basipterygoid process; bt, basal tuber; cap, capitate process; CN II — XII, cranial nerve foramina; cvn?, cavity for vomero-nasal bulb?; faCC, cerebral carotid artery and vein foramen; faSP?, sphenopalatine artery and vein foramen?; CR+FO, columellar recess and fenestra ovalis; fvOC, foramen for orbitocerebral vein; gTO, temporoorbital artery and vein groove; ls, laterosphenoid; MF, metotic foramen; ors, orbitosphenoid; oto, otoccipital; pbs, parabasisphenoid; pro, prootic; so, supraoccipital; speth, sphenethmoid; vf, vascular foramen.
Fig. 13. ZIN PH 281/16, referred specimen of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. Photograph and corresponding CT-based model in oblique anterodorsal view (A, B) and parasagitally sectioned CT-based model showing left endocranial surface in medial view (C). Scale bars each equal 1 cm. Sutures are represented by solid lines; possible sutures are represented by dashed lines. Abbreviations: bo, basioccipital; cap, capitate process; cerc, cerebral cavity; CN II — XII, cranial nerve foramina; cvn?, cavity for vomero-nasal bulb?; dsOc, occipital venous sinus depression; fCC, cerebral carotid artery and vein foramen; floc, floccular fossa; fvMC, middle cerebral vein foramen; fvOC, foramen for orbitocerebral vein; lagf, lagenar fossa; Is, laterosphenoid; olfc, olfactory cavity; oto, otoccipital; pbs, parabasi-sphenoid; pro, prootic; so, supraoccipital; speth, sphenethmoid; ves, vestibular cavity; vf, vascular foramen.
in other ankylosaurs (Miyashita et al., 2011). Features of the complex external surface of the sphenethmoidal complex include prominent vertical striations dorsal to the foramen for CN II (possible site of attachment of the preorbital septum; ectethmoid in Miyashita et al. [2011]) and multiple grooves, ridges, and bumps around the foramen for CN II (indicating courses of neurovas-cular elements and possible attachment sites of ocular musculature).
The parabasisphenoid constitutes most of the an-teroventral aspect of the lateral surface of the braincase (Fig. 12A, C). Paired grooves mark the course of CN VI from the dorsum sellae toward the external surface of the braincase, bypassing the hypophyseal cavity (Figs. 12D, 13). The participation of the prootic in the canal for CN VI is not clear. A large foramen for the cerebral carotid artery is present on either side of the specimen (faCC in Fig. 12B, D). The breakage of the specimen anterior to the cerebral carotid foramen on both sides makes the
interpretation of certain foramina challenging. On the left side, the anterior rounded margin of a foramen is preserved (faSP?, Fig. 12D). This opening is comparable in size to the foramen for the cerebral carotid artery. Dorsal to it, the bone is broken, and no additional foramina could be identified. On the right side, the margin of a small-sized foramen is preserved anterior to the foramen for CN VI (faSP?, Fig. 12B). It is half as large as the abovementioned foramen on the left side but is located at the same level as the latter. Considering the position of these foramina, both of them could equally likely represent foramina for the sphenopalatine artery or foramina for CN III. In the latter case, the sphenopal-atine artery would have branched off from the cerebral carotid artery before the latter entered the hypophyseal cavity in ZIN PH 281/16. However, as the nature of foramen for CN III is controversial in the holotype (see above), we assume that these foramina represent exits of the sphenopalatine artery (faSP?, Fig. 12B, D). Thus, the
Fig. 14. ZIN PH 281/16, referred specimen of Bissektipelta archibaldi from the Bissekty Formation (Turonian), Uzbekistan. Cranial endocast with endosseous labyrinth of the inner ear in right lateral (A), ventral (B), dorsal (C), and oblique ventrolateral (D) views. Scale bars each equal 1 cm. Abbreviations: aCC, cerebral carotid artery and vein; aCE/vCC?, caudal encephalic artery/caudoventral cerebral vein; aSP?, sphenopalatine artery and vein; cbl, cerebellum; ch, cerebral hemisphere; CN II — XII, cranial nerves; cochd, endosseous cochlear duct; hyp, hypophysis (pituitary); lab, endosseous labyrinth; MF, metotic passage; mo, medulla oblongata; olfb, olfactory bulb; olft, olfactory tract; pls, perilymphatic sac; vCD, dorsal heard vein; vn?, vomero-nasal bulbs?; vOC, orbitocerebral veins.
presence and position of separate foramina for CN III in ZIN PH 281/16 are uncertain.
Endocranial surface. The endocranial surface of ZIN PH 281/16 does not differ significantly from that of the holotype in the general division into regions or the distribution of neurovascular foramina (Fig. 13). Its surface is mostly smooth, indicating a loose infilling by the brain. Vertical striations at the surface of the cavity for the olfactory tract probably result from a closer contact between the brain and its dura with the walls of the brain-case (Fig. 13A). Most sutures on the endocranial surface are obliterated; however, the basioccipital-otoccipital, basioccipital-parabasisphenoid, and prootic-otoccipital sutures are visible (Fig. 13A). The straight basioccipital-otoccipital sutures extend ventral to the internal foramina for CN XII and disappear towards the MF and la-genar fossae. The suture between the basioccipital and parabasisphenoid extends transversally between large, unossified lagenar fossae. The basioccipital apparently forms most of the ventral endocranial surface. Possible
prootic-otoccipital sutures extend as ridges on the preserved surface of both otic capsules.
A single transverse canal for CN II opens laterally on either side in a separate foramen (Fig. 13C). The hypophyseal cavity is comparatively shallow, nearly half the dorsoventral depth of the cerebral cavity above it. The tentorial crest separating the cerebral and cerebellar cavities appears to be prominent and sharp at its dorsal part; it is broken off ventrally on both sides and the canal for CN VI is exposed (Fig. 13B). The dorsum sellae preserves the central triangular projection. Posterodor-sally in the prootic, a foramen leads into two canals, one anteroventral (for CN VII) and the other posterodorsal into the otic capsule (for CN VIII) (Fig. 13C). The medial walls of both otic capsules are largely unossified, and two huge lagenar fossae are present on the floor of the endo-cranium (ves+lagf in Fig. 13B). Anterodorsal to the otic capsule, a distinct fossa is present on either side, which is not particularly evident in the holotype (floc, Fig. 13C). These depressions correspond in their position to the
floccular (auricular) fossae in other archosaurs (Gower, 2002; Sampson and Witmer, 2007; Witmer et al., 2008; Sobral et al., 2016). Each fossa has a pair of sediment-filled openings at the bottom that apparently connect to the inner ear cavities within the bone (the CT data offer insufficient resolution to trace these structures with confidence). These foramina likely transmitted vessels to the inner ear labyrinth. Although the external foramina for CN XII vary in number (two or three) between both sides of the specimen, there are three internal openings on either side (Fig. 13C).
Endocast. The digital
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The Basal Nodosaurid Ankylosaur Europelta carbonensis n. gen., n. sp. from the Lower Cretaceous (Lower Albian) Escucha Formation of Northeastern Spain
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[
"James I. Kirkland",
"Luis Alcalá",
"Mark A. Loewen",
"Eduardo Espílez",
"Luis Mampel",
"Jelle P. Wiersma"
] |
2013-08-31T00:00:00
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Nodosaurids are poorly known from the Lower Cretaceous of Europe. Two associated ankylosaur skeletons excavated from the lower Albian carbonaceous member of the Escucha Formation near Ariño in northeastern Teruel, Spain reveal nearly all the diagnostic ...
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PubMed Central (PMC)
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https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3847141/
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PLoS One. 2013; 8(12): e80405.
PMCID: PMC3847141
PMID: 24312471
The Basal Nodosaurid Ankylosaur Europelta carbonensis n. gen., n. sp. from the Lower Cretaceous (Lower Albian) Escucha Formation of Northeastern Spain
, 1 , * , 2 , 3 , 4 , 2 , 2 and 3 , 4
James I. Kirkland
1 Utah Geological Survey, Salt Lake City, Utah, United States of America
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Luis Alcalá
2 Fundación Conjunto Paleontológico de Teruel-Dinópolis, Museo Aragonés de Paleontología, Teruel, Spain
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Mark A. Loewen
3 Natural History Museum of Utah, Salt Lake City, Utah, United States of America
4 Department of Geology & Geophysics, University of Utah, Salt Lake City, Utah, United States of America
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Eduardo Espílez
2 Fundación Conjunto Paleontológico de Teruel-Dinópolis, Museo Aragonés de Paleontología, Teruel, Spain
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Luis Mampel
2 Fundación Conjunto Paleontológico de Teruel-Dinópolis, Museo Aragonés de Paleontología, Teruel, Spain
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Jelle P. Wiersma
3 Natural History Museum of Utah, Salt Lake City, Utah, United States of America
4 Department of Geology & Geophysics, University of Utah, Salt Lake City, Utah, United States of America
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Richard J. Butler, Editor
1 Utah Geological Survey, Salt Lake City, Utah, United States of America
2 Fundación Conjunto Paleontológico de Teruel-Dinópolis, Museo Aragonés de Paleontología, Teruel, Spain
3 Natural History Museum of Utah, Salt Lake City, Utah, United States of America
4 Department of Geology & Geophysics, University of Utah, Salt Lake City, Utah, United States of America
University of Birmingham, United Kingdom
Competing Interests: The authors have declared that no competing interests exist.
Conceived and designed the experiments: JIK LA MAL. Performed the experiments: JIK MAL. Analyzed the data: JIK LA MAL. Wrote the paper: JIK LA MAL. Oversaw the entire Ariño project: LA. Co-directed the excavation: EE LM. Oversaw the preparation of all the fossil materials: EE. Assisted with character evaluations and in constructing many of the figures: JPW. Coordinated quarry mapping and photography: LM EE.
Copyright © 2013 Kirkland et al
This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are properly credited.
Abstract
Nodosaurids are poorly known from the Lower Cretaceous of Europe. Two associated ankylosaur skeletons excavated from the lower Albian carbonaceous member of the Escucha Formation near Ariño in northeastern Teruel, Spain reveal nearly all the diagnostic recognized character that define nodosaurid ankylosaurs. These new specimens comprise a new genus and species of nodosaurid ankylosaur and represent the single most complete taxon of ankylosaur from the Cretaceous of Europe. These two specimens were examined and compared to all other known ankylosaurs. Comparisons of these specimens document that Europelta carbonensis n. gen., n. sp. is a nodosaur and is the sister taxon to the Late Cretaceous nodosaurids Anoplosaurus, Hungarosaurus, and Struthiosaurus, defining a monophyletic clade of European nodosaurids– the Struthiosaurinae.
Introduction
Ankylosaurs were first described from the Lower Cretaceous of England with Hylaeosaurus armatus (Valanginian) described in 1833 [1]–[3]. Hylaeosaurus is one of the three dinosaurs on which the Dinosauria were defined [4] and one of the first dinosaurs for which a full-sized life reconstruction was attempted at the Crystal Palace Park in London in 1854 [5]. Although first mentioned in an anonymous article in the September 16th 1865 issue of the “The Illustrated London News” by Sir Richard Owen [6], the Early Cretaceous (Barremian) Polacanthus was not described formally as Polacanthus foxii by Hulke until 1882 [7]–[10]. The abundant plates and spines of these ankylosaurs are characteristic of the Lower Cretaceous up into the lower part of the Aptian stage [11], [12]. In 1867, Huxley described the fragmentary Acanthopholis from the base of the Upper Cretaceous (Cenomanian) [13]–[15]. Additionally, in 1879, Seeley [16] described the juvenile nodosaurid Anoplosaurus curtonotus [17] from the uppermost Lower Cretaceous (upper Albian) Cambridge Greensand. Subsequent descriptions of the fragmentary remains of ankylosaurs from the Early Cretaceous of Europe have been tentatively assigned to the genus Polacanthus [18].
Only nodosaurids have been described from the Upper Cretaceous of Europe with Struthiosaurus austriacus described from the Campanian of Austria in 1871 [19]–[24] followed by Struthiosaurus transylvanicus [25], [26], [27] from the uppermost Cretaceous (upper Maastrichtian) strata of Romania. Until recently, all Late Cretaceous ankylosaur fossils in Europe have been assigned to Struthiosaurus [28]–[30] including Struthiosaurus languedocensis from the Campanian of southern France [31]. The primitive nodosaurid Hungarosaurus tormai [32], [33] from the mid-Late Cretaceous (Santonian) is now known from multiple specimens and has become the best documented ankylosaur in Europe.
Fragmentary ankylosaur remains are also known from a number of localities from the Middle to Upper Jurassic strata of Europe, but have been relatively uninformative as specimens are based largely on isolated skeletal elements [34].
Northeastern Spain has contributed many dinosaur discoveries from both Lower and Upper Cretaceous strata in recent years [35]. The Early Cretaceous dinosaurs discovered to date include numerous sauropods, iguanodonts, and ankylosaurs from the Barremian-lower Aptian, with all the fragmentary ankylosaur material assigned tentatively to the genus Polacanthus [25], [28], [36]–[40]. All the Late Cretaceous ankylosaurs from Spain have in turn been assigned to Struthiosaurus [28]–[30].
The earliest reported dinosaur remains from Spain were found in the Escucha Formation, few significant vertebrate fossils had been recovered from these rocks in the 140 intervening years [41], [42]. Current research on vertebrate sites in the Escucha Formation in the northern Teruel Province in the Community of Aragón, Spain, by the Fundación Conjunto Paleontológico of Teruel-Dinópolis has resulted in the discovery of an extensive new dinosaur locality in the open-pit Santa María coal mine near Ariño ( ) operated by Sociedad Anónima Minera Catalano-Aragonesa (SAMCA Group) [42]. The most abundant dinosaur identified is a distinctive iguanodontian ornithopod recently described as Proa valdearinnoensis [43]. Among the many other significant fossils excavated are two associated partial skeletons of a new species of ankylosaur, described herein as Europelta carbonensis n. gen., n. sp. This new taxon is the most completely known ankylosaur in Europe and adds considerable new information about Early Cretaceous ankylosaurian phylogeny and biogeography.
Geological Setting
Counterclockwise rotation of the Iberian Plate toward the end of the Early Cretaceous resulted in the development of a series of syndepositional sub-basins bounded by active faults within Ebro Basin south of the Pyrenean ranges, northeast of the Iberian Range, and northwest of the Catalan/Coastal Range [44], [45]. The new dinosaur locality is within the Oliete sub-basin on the northwest margin of the Escucha outcrop belt [42], [44]. The Formación Lignitos de Escucha and overlying Formación Arenas de Utrillas were initially described in 1971 [46]. These largely Albian-aged strata were deposited along the northwestern margin of the Tethys Sea during the fragmentation of this terrain, and overlie Aptian strata in the center of each sub-basin and unconformably overlie progressively older strata toward their margins. Initially, the Escucha Formation was divided into three members [47] and interpreted to be an unconformity-bounded lower to middle Albian depositional sequence, representing a progradational, tidally-dominated delta sequence [44], [48]–[52]. Recently, the upper “fluvial” member has been reinterpreted as an eolian depositional sequence separated from the underlying portions of the Escucha Formation by a regional unconformity [53]. We recognize this bipartite division of the Escucha Formation ( ).
The geologic age of the Escucha Formation has been considered to be early to middle Albian. It overlies Aptian strata in central basinal settings and is, in turn, overlain by the upper Albian Utrillas Formation [44]. However, both calcareous plankton (foraminifera and nanoplankton) [54] and palynomorphs [55], [56] indicate that the lower Escucha Formation is late Aptian in age. Both fresh and brackish coal-bearing strata are recognized below the regional unconformity within the Escucha [43]. However, reports on the microplankton restrict marine and marginal marine facies to the late Aptian in the lower Escucha Formation [54]–[56]. Marine ostracods have been reported from the upper Escucha Formation northeast of Teruel that confirm an Albian age for the upper portion of these strata in this area [57].
A sample of the matrix from the bonebed was processed for both palynomorphs and calcareous microfossils. The palynomorphs were exclusively of terrestrial origin and indicated an Albian age (Gerry Waanders, 2012, personal communication). The microfossils consisted exclusively of freshwater ostracods and charophytes. The ostracods represent new species and the charophytes are also reported from the Albian of Tunisia [58]. No arenaceous foraminifera were identified, which, along with the absence of dinoflagelates, indicates that the bonebed formed well inland of marine and brackish water influences ( ).
The bonebed is located immediately below the lowest mineable coal seam in the Santa María coal mine ( ), in a dark olive-gray to olive-black mudstone that preserves a high percentage of fossil plant debris. In overall appearance, the rock is much like the plant debris beds in the Wessex Formation on the Isle of Wight [59], [60] and, as in those beds, there is a great amount of pyrite (iron sulfide) disseminated through the matrix and in the fossils. Significant amounts of iron sulfide in the coals were found to decrease up section, away from marine and brackish-water environments. In addition to this depositional relationship, it has been speculated that detrital evaporites from exposed Triassic strata on the north and northwest sides of the basin have secondarily contributed significant amounts of sulfur to these coals [43], [61]. Additionally, the abundance of pyrite in the bones indicates that the long-term stability of the fossils is in question as pyrite breaks down in an expansive oxidation reaction that liberates corrosive sulfuric acid compounds that cannot be reversed [62]. The degradation by this pyrite is apparent on most of the bones soon after exposure to the surface. This is indicated by the rapid appearance of fine, powdery to crystalline gypsum coating bones and teeth, and by the expansion and shattering of some bones and teeth with internal gypsum formation ( ). Protocols are being developed to ensure the preservation of the primary data represented by these important fossils [42], [62].
The bonebed was located many tens of meters underground prior to strip mining operations in the Santa María coal mine. As mining operations proceed, more of the plant debris stratum containing the bonebed is exposed as simultaneous reclamation covers the previously exposed surface. Thus, with the help of mine managers, efficient methodologies for the documentation and extraction of significant fossils have been established [42]. By the end of 2012, an area of approximately 25 ha had been investigated and the areal distributions of 101 vertebrate concentrations were documented; 33 of these consisted of associated dinosaur skeletons (mostly iguanodonts) and 68 consisted of other vertebrate remains (mostly turtles and crocodilians). During this stage of the project, numerous dinosaurs (ornithischian elements and associated skeletons, and saurischian teeth), two types of turtle, crocodilians, fish (both ostheicthyians and selachiens), coprolites, molluscs (freshwater bivalves and gastropods), arthropods (ostracods), and abundant plant remains (logs, plant fragments, palynomorphs, and amber) have been excavated.
The bonebed designated AR-1 contains more than 5000 identifiable vertebrate specimens recovered from isolated skeletal remains and associated individual animals. All fossils receive a consecutive number from the site, each association is numbered as well. Thus:
AR-1-#fossil identifies each fossil found at the Ariño site (the ID written on each fossil);
AR-1/#concentration identifies a collection of bones belonging to a single skeleton;
AR-1-#fossil/#concentration identifies a fossil from a bone concentration # belonging or not belonging to a single skeleton.
The two associated ankylosaur skeletons described herein were separated by 200 meters. The location of the holotype AR-1/10 ( ) was still available for examination and sampling for microfossils in December of 2011 [58], while that of the paratype AR-1/31 ( ) was already inaccessible.
Materials and Methods
Paleontological Ethics Statement
All of the specimens described in this paper (AR-1/10 and AR-1/31) are reposited in the collections of the Fundación Conjunto Paleontológico de Teruel-Dinópolis/Museo Aragonés de Paleontología (FCPTD/MAP). Locality information is available from the registrar of the museum as per museum policy. All necessary permits were obtained for the described study, which complied with all relevant regulations. All of these specimens were collected under permits obtained from the Sociedad Anónima Minera Catalano-Aragonesa.
Nomenclatural Acts
The electronic edition of this article conforms to the requirements of the amended International Code of Zoological Nomenclature, and hence the new names contained herein are available under that Code from the electronic edition of this article. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information viewed through any standard web browser by appending the LSID to the prefix “http://zoobank.org/”. The LSID for this publication is: urn:lsid:zoobank.org:pub:9246FFA7-6271-4734-8E01-5590BE4A80C2. The LSID for Europelta carbonensis is: urn:lsid:zoobank.org:act:089040A3-1BCF-42D1-B99F-94840E2BB96D. The electronic edition of this work was published in a journal with an ISSN (1932-6203), and has been archived and is available from the following digital repositories: LOCKSS (http://www.lockss.org); PubMed Central (http://www.ncbi.nlm.nih.gov/pmc).
Terminology
We do not refer to the “armor” on the skull roof as caputegulae, as we consider these patterns in the Nodosauridae to reflect impressions of scale boundaries on the skull roof as opposed to thickened remodeled cranial bone. We use the term caudal rib instead of caudal transverse process. We employ the monophyletic clade Polacanthidae of Carpenter [63] to facilitate comparison with and discussion of a number of similar taxa (Gargoyleosaurus, Mymoorapelta, Hylaeosaurus, Polacanthus, Hoplitosaurus, and Gastonia). The most recent analysis of polacanthids as a monophylogenetic subfamily of nodosaurids was by Yang and others [64], who similarly defined them as the most inclusive clade containing Polacanthus foxii but not Ankylosaurus magniventris or Panoplosaurus mirus.
Institutional Abbreviations
AMNH, American Museum of Natural History, New York, New York, NHMUK, Natural History Museum, London, England, CEUM, Prehistoric Museum, Utah State University, Price, Utah, DMNH, Denver Museum of Nature and Science, Denver, Colorado, MPC, Geological Institute, Ulaan Bataar, Mongolia, FCPTD/MAP, Fundación Conjunto Paleontológico de Teruel-Dinópolis/Museo Aragonés de Paleontología, Teruel, Spain, FMNH, Field Museum of Natural History, Chicago, MPC, Institute of Geology, Mongolian Academy of Sciences, Ulaan Baatar, Mongolia; INBR, Victor Valley Museum, Apple Valley, California, IVPP, Institute of Vertebrate Paleontology and Paleoanthropology, Beijing, China, KUVP, Kansas Museum of Natural History, Lawrence, Kansas, MPC, Mongolian Paleontological Center, Ulaan Baatar, Mongolia; MNA, Museum of Northern Arizona, Flagstaff, Arizona, NMC, National Museum of Canada, Ottawa, Canada, NMW, National Museum of Wales, Cardiff, England, PIN, National Institute of Paleontology, Moscow, Russia, QM, Queensland Museum, Queensland, Australia, ROM, Royal Ontario Museum, Toronto, Canada, SDNHM, San Diego Natural History Museum, San Diego, California, SGDS, Saint George Dinosaur Discovery Site at Johnson Farm, St. George, Utah, SMP, State Museum of Pennsylvania, Harrisburg, Pennsylvania, SMU, Schuler Museum, Southern Methodist University, Dallas, Texas, USNM, National Museum of Natural History, Smithsonian Institution, Washington D.C.
Comparative Material
In addition to accessing the ever-expanding ankylosaur literature, the senior and third authors have had the opportunity to study firsthand much of the important ankylosaur material collected globally. From the basal thyreophorans: the type material of Scutellosaurus lawleri (MNA P1.175), the type material of Scelidosaurus harrisoni (NHMUK R 1111), and a large, exceptionally well-preserved, articulated Scelidosaurus specimen with intact armor, collected and owned by David Sole and currently exhibited at the University of Bristol. Also, a full cast of the left side of the skeleton (SGDS 1311) exhibited in southwestern Utah was examined.
In regards to Jurassic ankylosaurs: the extensive type and paratype material of Mymoorapelta maysi housed at the Museum of Western Colorado, Gargoyleosaurus parkpinorum (DMNH 27726), and the dentary of Sarcolestes leedsi (NHMUK R 2682) were studied.
Early Cretaceous polacanthine ankylosaur material examined includes Polacanthus foxii (NHMUK R 175, 9293), Hylaeosaurus armatus (NHMUK R 3775), Hoplitosaurus marshi (USNM 4752), and the extensive material of Gastonia burgei material housed at the Prehistoric Museum (including holotype CEUM 1307 and paratype material), and cranial material from a minimum of six individuals at Brigham Young University's Earth Science Museum, together with the postcranial skeleton of an unnamed new species of polacanthine (BYU 245).
Among basal shamosaurine-grade ankylosaurids, Cedarpelta bilbyhallorum (including CEUM 12360 and paratype material), Shamosaurus scutatus (PIN 3779/2), and a cast of the skull of Gobisaurus domoculus (IVPP 12563) housed at the Royal Tyrell Museum were studied.
Among derived North American ankylosaurs, Nodocephalosaurus kirtlandensis (SMP-VP-900), Ankylosaurus magniventris (AMNH 5214, 5859; NMC 8880), Anadontosaurus lambei (NMC 8530), Dyoplosaurus acutosquameus (ROM 784), Scolosaurus cutleri (NHMUK, R 5161), and several important examples of Euoplocephalus tutus, (AMNH 5404, 5409; RTMP 91.127.1) were examined.
Asian ankylosaur material researched include an adult skull of Tsagantegia longicranialis (MPC 100/1306), China, Pinacosaurus grangeri (AMNH 6523) and three undescribed skulls personally excavated by JIK from the Djadokhta Formation, Shabarakh Usu (Flaming Cliffs, Mongolia) and housed at MAS, Talarurus plicatospineus (composite skeleton made up of parts of many individuals assigned to PIN 557), cast skull of Saichania chulsanensis (PIN 3141/251), a relatively complete specimen referred to Saichania with in situ armor but lacking its skull (MPC 100/1305), Tarchia gigantea (PIN 3142/250), a cast skull of Minotaurasaurus ramachandrani (INBR 21004), and a cast skeleton of Crichtonsaurus benxiensis housed in the Museum at the Chaoyang Bird National Geopark, Liaoning.
Numerous nodososaurids were examined, including the Early Cretaceous nodosaurids Sauropelta edwardsi (AMNH, 3016, 3032, 3035, 3036; YPM 5502, 5529, 5499, 5178), Peloroplites cedrimontanus (CEUM 26331 and the extensive paratype material), and Pawpawsaurus campbelli (SMU 73203; = “Texasestes” pleurohalio USNM 337987). The early Late Cretaceous nodosaurids reviewed include Animantarx ramaljonesi (CEUM 6228), Silvisaurus condrayi (KUVP 10296), Nodosaurus textilis (YPM 1815), and Stegopelta landerensis(FMNH UR88) and the Late Cretaceous nodosaurids Panoplosaurus mirus (NMC 2759), Edmontonia rugosidens (USNM 11868; AMNH 5665), Edmontonia longiceps (NMC 8531), Denversaurus schlessmani (DMNH 468), casts of Struthiosaurus austriacus at the Carnegie Museum (PIUW 2349) and Struthiosaurus transylvanicus (NHMUK R 4966).
Enigmatic taxa such as the skull of Minmi paravertebrata (QM F18101), the skeleton of Liaoningosaurus paradoxus (IVPP V12560), and Aletopelta coombsi (SDNHM 33909) were also examined.
Results
Systematic Paleontology
Dinosauria Owen, 1842 [65]
Ornithischia Seeley, 1887 [66]
Thyreophora Nopcsa, 1915 [25]
Ankylosauria Osborn, 1908 [67]
Nodosauridae Marsh, 1890 [68]
Struthiosaurinae Nopcsa, 1923 [69]
Diagnosis
Nodosaurid ankylosaurs that share a combination of characters including: narrow predentaries; a nearly horizontal, unfused quadrates that are oriented less than 30° from the skull roof, and condyles that are 3 times transversely wider than long; premaxillary teeth and dentary teeth that are near the predentary symphysis; dorsally arched sacra; an acromion process dorsal to midpoint of the scapula-coracoid suture; straight ischia, with a straight dorsal margin; relatively long slender limbs; a sacral shield of armor; and erect sacral armor with flat bases. Struthiosaurinae is defined as the most inclusive clade containing Europelta but not Cedarpelta , Peloroplites, Sauropelta or Edmontonia .
Europelta Kirkland, Alcalá, Loewen, Espílez, Mampel, and Wiersma 2013 gen. nov.
urn:lsid:zoobank.org:act:62808E3D-85BE-4AE3-B771-9CFF2C6AC054
Etymology
“Euro” as a contraction for Europe in regard to its origin and “pelta” Greek for shield, a common root for ankylosaurian genera; “Europe's shield”.
Diagnosis
Same as for the only known species below.
Europelta carbonensis Kirkland, Alcalá, Loewen, Espílez, Mampel, and Wiersma 2013 gen. et sp. nov.
urn:lsid:zoobank.org:act:089040A3-1BCF-42D1-B99F-94840E2BB96D
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Etymology
The specific name “carbonensis” from the coal, is in honor of access to the fossil locality in the Santa María coal mine provided by Sociedad Anónima Minera Catalano-Aragonesa (SAMCA Group), which has been extracting coal in Ariño (Teruel) since 1919.
Holotype
AR-1/10, a disarticulated partial skeleton reposited at Fundación Conjunto Paleontológico de Teruel-Dinópolis/Museo Aragonés de Paleontología (FCPTD/MAP). The holotype consists of: a mostly complete skull (AR-1-544), isolated left and right nasals (AR-1-133, and AR-1-639), a dentary fragment (AR-1-362), 15 isolated teeth (AR-1-323 to AR-1-325, AR-1-343, AR-1-358, AR-1-417, AR-1-418, AR-1-423, AR-1-424, AR-1-428, AR-1-454, AR-1-482, AR-1-563, AR-1-564 and AR-1-567), an atlas (AR-1-649), five cervical vertebrae (AR-1-431, AR-1-449, AR-1-533, AR-1-637, AR-1-650), two cervical ribs (AR-1-450, AR-1-4452), AR-1-638 (possibly the first dorsal vertebrae), seven more posterior dorsal vertebrae (AR-1-154, AR-1-155, AR-1-322, AR-1-430, AR-1-448, AR-1-478, AR-1-535, AR-1-556), a section of synsacrum (AR-1-154), three isolated dorsal ribs (AR-1-331, AR-1-333, AR-1-476), seven dorsal rib fragments (AR-1-339, AR-1-341, AR-1-427, AR-1-534, AR-1-641, AR-1-642, AR-1-676), three caudal vertebrae (AR-1-562, AR-1-635, AR-1-636), four chevrons (AR-1-560, AR-1-561, AR-1-569, AR-1-4451), a coracoid with a small portion of scapula (AR-1-657), a scapular blade fragment (AR-1-429), two xiphosternal plates (AR-1-252, AR-1-4675), two partial humeri (AR-1-327, AR-1-655), a right ilium-ischium-pubis (AR-1-479), a left ischium-pubis (AR-1-129), and 70 osteoderms (AR-1-126 to AR-1-128, AR-1-192, AR-1-234, AR-1-241, AR-1-246, AR-1-247, AR-1-272, AR-1-276, AR-1-438, AR-1-444, AR-1-447, AR-1-461, AR-1-462, AR-1-464, AR-1-467, AR-1-472, AR-1-496 to AR-1-530, AR-1-553, AR-1-651 to AR-1-653, AR-1-659, AR-1-675, AR-1-4450, AR-1-4454 to AR-1-4463).
Paratype
AR-1/31, a partial skeleton deposited at Fundación Conjunto Paleontológico de Teruel-Dinópolis/Museo Aragonés de Paleontología (FCPTD/MAP). The paratype consists of a partial left jaw with dentary and surangular (AR-1-3698) and isolated angular (AR-1-2945), 10 teeth (AR-1-3432, AR-1-3495, AR-1-3524, AR-1-3650, AR-1-3699 to AR-1-3701, AR-1-3705, AR-1-3706, AR-1-3961), five cervical vertebrae (AR-1-3586, AR-1-3632, AR-1-3657, AR-1-3671, AR-1-3676), nine dorsal vertebrae (AR-1-3489, AR-1-3586, AR-1-3633, AR-1-3662, AR-1-3672 to 3675, AR-1-3677, AR-1-3704), three to four? dorsosacral vertebrae (AR-1-3450, AR-1-3451), a sacrum (AR-1-3446), a caudosacral vertebra (AR-1-3512), two sacral rib fragments (AR-1-3452, AR-1-3460), 14 caudal vertebrae (AR-1-2950, AR-1-3204, AR-1-3206, AR-1-3243, AR-1-3265, AR-1-3348, AR-1-3398, AR-1-3478, AR-1-3615, AR-1-3616, AR-1-3714 to 3717), a right ilium (AR-1-3490), two left ilium fragments (AR-1-3521, AR-1-3571), two ischia with fused pubes (AR-1-3648, AR-1-3649), a right femur (AR-1-3244), a right tibia (AR-1-3237), a right fibula (AR-1-3238), a calcaneum (AR-1-3239), four metatarsals (AR-1-3100, AR-1-3173, AR-1-3233, AR-1-3234), eight phalanges (AR-1-3032, AR-1-3066, AR-1-3174, AR-1-3179, AR-1-3324, AR-1-3234, AR-1-3292, AR-1-3356), nine unguals (AR-1-2952, AR-1-2986, AR-1-3172, AR-1-3181, AR-1-3182, AR-1-3288, AR-1-3291, AR-1-3386, AR-1-3711), and 90 osteoderms (AR-1-3024, AR-1-3030, AR-1-3074 to AR-1-3076, AR-1-3080, AR-1-3145, AR-1-3159, AR-1-3180, AR-1-3207 to AR-1-3209, AR-1-3216, AR-1-3223, AR-1-3226 to AR-1-3229, AR-1-3292, AR-1-3236, AR-1-3242, AR-1-3338 to AR-1-3340, AR-1-3390, AR-1-3438, AR-1-3447 to AR-1-3449, AR-1-3491, AR-1-3492, AR-1-3494, AR-1-3506, AR-1-3540, AR-1-3572 to AR-1-3576, AR-1-3587, AR-1-3588, AR-1-3590, AR-1-3597, AR-1-3598, AR-1-3608 to AR-1-3613, AR-1-3638, AR-1-3658, AR-1-3680 to AR-1-3684, AR-1-3687, AR-1-3708, AR-1-3720, AR-1-3721, AR-1-3932 to AR-1-3960).
Locality and Horizon
The type locality, Fundación Conjunto Paleontológico of Teruel-Dinópolis locality AR-1, is located east of Ariño, Teruel Province, Spain. The fossil horizon is below the lowest mineable coal seam at Sociedad Anónima Minera Catalano-Aragonesa Group's Ariño coal mine in a plant debris bed in the lower Escucha Formation [42]. The paratype AR-1/31 was located 200 m laterally from the holotype AR-1/10 in the same bed. Pyrite is common within the bone and the surrounding sediment of the bonebed, common also in plant debris beds in the older Wessex Formation on the Isle of Wight [58].
Age
Elsewhere, the Escucha Formation has been interpreted as late Aptian to early Albian in age based on nanofossils, planktonic foraminifera, dinoflagellates and palynomorphs [50], [52]. An analysis of the palynomorphs, ostracods, and charophytes from AR-1 indicates that the site is completely of early Albian age [57].
Diagnosis
The quadrate is shorter and mediolaterally wider than in any other ankylosaur. The posterior margin of the skull is concave in dorsal view. The sacrum is arched dorsally about 55° in lateral view. The pubis is fully and uniquely fused to the ischium with a slot-shaped foramen between the post-pubic process and the position of the pubic peduncle forming an ischiopubis. The tibia is longer relative to the length of the femur (90%) than in other ankylosaurs for which these proportions are known. Laterally compresed, flanged osteoderm with a flat plate-like base is present anteriorly on the pelvic shield.
Description and Comparisons
Skull
The skull (AR-1-544/10) was lying on its dorsal surface and is moderately well preserved although distorted through compaction ( ). The palate is crushed in toward the skull roof, resulting in the medial rotation of both maxillae with the posterior teeth displaced into the posterior palate. The sheet-like palatal bones are highly fragmented. The braincase is crushed along the plane of the cranial nerve openings and the fenestra ovalis completely obscures them. Unexpectedly, the right quadrate ( ) and associated portion of the palate was dislodged from the skull and subsequently crushed across the ventral side of the basicranium. This gives the impression that these bones had been expelled from inside the skull prior to compaction. Both the left and right nasals were separated from the skull and the premaxillae (whereas possibly present upon discovery) have not been identified.
The skull has a minimum length of 370.3 mm from the anterior end of the maxillae to the rear margin of the squamosals. The skull has a maximum width of 299.1 mm at the orbits and narrows to 203.7 mm at the posterior end of the skull at the squamosals, giving the skull the “pear-shaped” dorsal profile characteristic of derived nodosaurids [70], [71]. Although tapering posteriorly, there is no distinct post-temporal notch as in polacanthids and other nodosaurids [63].
The maxillae ( ) are irregularly sculptured externally with a flattened, horizontally oriented buccal recesses that are inset approximately 2 cm. The anterior margin of the maxilla appears to form the posterior margin of a relatively simple naris relative to derived nodosaurids and ankylosaurids. Medially, there is no evidence that the maxilla formed a portion of a secondary palate. The tooth row was arched ventrally with an estimated 22–25 alveoli increasing in size posteriorly as in Edmontonia [72]. In ventral orientation, the tooth rows are only moderately deflected medially, such that the palate would not have had a pronounced hourglass appearance typical of derived nodosaurs such as Pawpawsaurus, Edmontonia, and Panoplosaurus [73]–[75]. However, it is not dissimilar from that of the primitive nodosaurid Silvisaurus [76], [77].
The nasals (AR-1-133/10, AR-1-639/10) are relatively large and subrectangular, tapering somewhat anteriorly ( ). Both nasals extend laterally from their relatively straight, unfused midline suture before flexing down to a sutural contact with the maxillae that extends for most of their length. When rearticulated onto the skull, they appear to fit well, despite the skull's distortion. Most ankylosaurs have fused nasals except the nodosaurids Silvisaurus [76], [77] and Niobrarasaurus [78], although the nasals are unknown in European nodosaurids [24], [32], [33]. A distinct tongue-like process projects from the nasal's posterior margin and would have overlapped the frontals. The external surface is lightly textured and the internal surface is relatively smooth, suggesting the narial passage was large and simple, rather than convolute as in derived nodosaurids and ankylosaurids [79], [80].
The orbits are somewhat crushed and the sutures of the bones surrounding them are obscured by fusion. The orbits are subrectangular in shape, are slightly more elongate anteoposteriorly and are directed anterolaterally. The prominent and evenly rounded suborbital horn is formed mostly from the quadratojugal posterior to the ventral margin of the orbit, as in most derived ankylosaurs [81], [82] and unlike that in polacanthids such as Mymoorapelta, Gargoyleosaurus, and Gastonia where the suborbital horn is below the orbit and is formed exclusively by the jugal [83]–[85]. The suborbital horn appears to be unornamented and hides the head of the quadrate in lateral view.
The lateral wall of the skull extends posteriorly behind orbit with a dorsoventally wide posterior notch, such that the lower temporal opening is just visible in lateral view. There is no lateral wall of skull behind the orbits in polacanthids [70], [81] and most nodosaurids other than Peloroplites [86], Silvisaurus [76], Struthiosaurus transylvanicus [22], [23] and one specimen from the Dinosaur Park Formation assigned to Edmontonia (ROM 1215) [88], although in these taxa the lower temporal opening is still visible in lateral view as in Europelta. The lower temporal opening is completely obscured in lateral view in Cedarpelta [84], [86], Shamosaurus [89]–[91], Gobisaurus, [92] Zhongyuansaurus [93] and all derived ankylosaurids.
Although the palate is fragmented and crushed along the internal surface of the skull roof, the fragments of the vomer suggest it did not extend ventrally to the level of the tooth row. Additionally, the broad sheet-like pterygoids appear to have been flexed nearly dorsally against the anterior portion of the basicranium as in nodosaurids and not like the open transversely oriented pterygoids characteristic of ankylosaurids or polacanthids [94].
The posterolateral margin of the pterygoid is fully fused to the quadrate. There is a sutural contact between the straight, nearly vertical quadrates and the quadratojugal laterally. The quadrates are wide transversely and thin rostrocaudally as compared to the mediolaterally narrower quadrates of other ankylosaurs [82]. The contact with the squamosal is also transversely wide, unlike the narrow, rounded contact seen in many ankylosaurs such as Mymoorapelta (Kirkland, pers. obs.) and Cedarpelta [63], [86]. The mandibular articulation is proportionally wider than in any other ankylosaur examined as a part of this study and the medial condyle larger than the lateral condyle. The ratio of mediolateral quadrate width to dorsoventral quadrate length is 0.77 (94 mm/122 mm). The anteropostior length of the quadrate condyle is 31 mm. There is no fusion between the quadrates and the paroccipital processes.
Vertical compaction has obscured the posterior view of the skull, in particular the foramen magnum and the supraoccipital. However, even with compaction it is apparent that in occipital view the skull was subrectangular and wider than tall as in Gargoyleosaurus, Gastonia, and most other derived anklylosaurs, and unlike the narrow, highly arched occipital region of Struthiosaurus [22]. The paroccipital processes extend horizontally lateral to the foramen magnum and then flare dorsoventrally by approximately 100% of their minimum widths. They angle posteriorly at about 30 degrees when viewed ventrally ( ). In morphology and orientation, they are most similar to those in Gargoyleosaurus [95] although ventral twisting is not present. In most other ankylosaurs, the paroccipital processes extend straight laterally [81], [96] or may be flexed ventrally as in Gastonia [83]. A triangular wedge of bone of unknown identity is fused to the anterior ventrolateral margin of the paroccipital, separating it from the quadrate.
The subspherical occipital condyle ( ) has a width of 59.4 mm and height of 46.5 mm and lacks a distinct neck to separate it from the rest of the basicranium. Although no cranial sutures are visible, the occipital condyle does appear to be composed exclusively of the basioccipital. It is similar in overall morphology to that of the basal ankylosaurid Cedarpelta [88] except that the occipital condyle angles somewhat ventrally, but not as much as in more derived nodosaurids [71], [82]. The ventral surface of the relatively elongate basioccipital is broadly convex. Again, as in Cedarpelta [88], there are no distinct, separate basal tubera between the basioccipital and the short basisphenoid, but instead there is a prominent transverse flange extending across the ventral surface of the basicranium along the line of this suture. The pterygoid processes appear to be short, but are completely obscured by crushed pterygoids bone fragments that wall off the anterior part of the braincase as in most nodosaurids.
The skull roof ( ) is roughened texturally by remodeling of the bone surface as in Cedarpelta, the nodosaurids Sauropelta and Peloroplites, and the shamosaurine-grade ankylosaurids Shamosaurus and Gobisaurus [81], [86], [88]. Europelta differs from these specimens in that some of the margins of the scale impressions on the skull roof are visible, as seen in Edmontonia, Panoplosaurus and Struthiosaurus [22], [77]. These scale margins are represented by shallow grooves that are difficult to see relative to the textured surface of the skull and the cracks in the bone due to compaction. These grooves are particularly evident along the lateral margins of the skull roof above the orbit. An extensive median scale appears to have covered much of the central portion of the skull between and posterior to the orbits on the frontals and parietals as other nodosaurids [63], [82]. There does not appear to be any distinct nuchal ornamentation. The skull is thickened above the orbit, but there is not a distinct supraorbital boss, a condition similar to Peloroplites, Cedarpelta, Shamosaurus, and Gobisaurus [86], [88]–[90], [92]. Narrow grooves along the margin of the skull in this area above the orbits suggest that a particularly robust pair of scales were present in this area as indicated by a deep groove bisecting this ornamented area directly above the orbit. Weak grooves delineate a small scale without underlying ornamentation separating the posterior supraorbital scale from the squamosal horn forming the posteriolateral margin of the skull roof. The squamosal horn is ornamented by narrow grooves radiating from its apex onto the skull roof. Grooves on the anterolateral sides of the fronto-parietal scale appear to delineate two scales between the anterior supraorbital scales. Unfortunately, no distinctive scale boundaries are recognizable on the nasals, although the dorsal surfaces of the nasals are textured. Several elongate scales rimmed the lateral raised margin around the orbit.
In dorsal view, the posterior margin of the skull is concave, whereas it is nearly straight or convex in all other nodosaurids. This reflects the posterior angulation of the paraoccipital processes and the squamosal horns. Interestingly, the occipital condyle is barely visible, though not completely obscured in dorsal view. There is no evidence of any distinct nuchal sculpturing. The skull roof is relatively flat but a slight dome may have been present prior to crushing. However, it is clear that the skull roof is not as highly domed as in many other nodosaurids, such as Struthiosaurus [22], [26].
Attempts were made to image the skull using X-ray photography and CT scanning. The abundance of pyrite present in the skull ( ) presents a strong limitation in the use of these techniques as pyrite is opaque to X-rays.
Mandible
A small dentary fragment extending for only four complete alveolae (AR-1-133/10) was preserved from the holotype skeleton ( ). However, a robust left dentary and splenial are preserved together (AR-1-3698/31) from the paratype specimen ( ). The splenial is not in its posteriomedial position relative to the dentary, but is fused across the posterior portion of the tooth row transversely. Additionally, an isolated left angular with a distinct highly sculptured scale along its ventral margin (AR-1-2945/31), was recovered ( ).
The dentary is 184.7 mm long with a minimum of 21 tooth positions, with no possibility of more than two unpreserved alveoli as determined by the position of the suture with the angular and surangular. As with the maxillary teeth, the alveoli are more than twice as large posteriorly. There is only 1.5 cm between the anteriormost alveoli and the symphysis, suggesting that there may have been premaxillary teeth as at least nine anterior teeth would have been positioned to oppose the premaxilla. The primitive ankylosaurs Sarcolestes [34], [98], Gargoyleosaurus, [85], Silvisaurus [76], Animantarx [97], Sauropelta [99], Anoplosaurus [17], Hungarosaurus [33] and Struthiosaurus [22] have a short anterior diastema, and thus a narrow predentary, whereas this diastema is longer in ankylosaurs with wide predentaries. However, the symphysis in Europelta is robust and dorsoventrally deeper (45.0 mm deep and 29.00 mm across) than in ankylosaurs [82], and is most similar to the deep symphysis of Hungarosaurus [32], further suggesting a reduced predentary with a rudimentary ventral process. The symphysis is marked by two deep anteroposteriorly directed grooves. A row of foramina extends posteriorly on the lateral surface of the dentary from just dorsal to the buccal recess to the notch for the surangular, whereas nutritive foraminae are not clearly visible ventral to the alveolae on the medial side of the dentary as in other ankylosaurs. The recessed tooth row is deflected medially and forms a convex arch in lateral view. The dentary of Hungarosaurus is deeper dorsoventrally than that of Europelta [33].
The splenial ( ) is a thin bone with a convex ventral margin 156.6 mm long that contacts the angular. It has the appearance of an obtuse triangle in medial view. There is large, well-developed intermandibular foramen (7 mm long and 5.3 mm wide) 50 mm from its anterior end.
The angular ( ) has a maximum length of 175 mm. The lateral margin is highly rugose, because the bone is textured and remodeled to support a large scale, extending about 10–12 mm ventral to the ventral margin of the angular for most of its length. A distinct ridge marks the dorsal limit of the mandibular ornament medially, where it is in contact with the ventral margin of the splenial. Dorsal to this contact the bone is smooth. The ventral extent of the textured bone supporting the mandibular scale is similar to that observed in ankylosaurids such as Euoplocephalus [95] and Minataurasaurus [100], rather than the more lateral orientation found in Gargoyleosaurus [93] and in nodosaurids like Sauropelta [99] and Panoplosaurus [101].
Teeth
A large number of teeth are preserved from both the holotype AR-1/10 (20+) and the paratype AR-1/31 (15+) although many have drifted away from the alvaeolae. We assume that the teeth associated with the holotype pertain to the maxilla (several are preserved in the palate and in the maxilla) and those of the paratype pertain to the dentary (several are preserved in the dentary). In general, the cutting surfaces of the teeth are not well preserved, but a few exceptions exist. Wear facets were not observed on any of the teeth. The roots for both dentary and maxillary teeth are swollen lingually, are three to four times the length of the crowns, and are subquadrate in cross-section. One small tooth (AR-1-343/10) is more highly asymmetrical mesiodistally and may represent a premaxillary tooth ( ).
The isolated maxillary teeth ( ) have a weakly developed labial cingulum and a strongly developed lingual cingulum. The best preserved right tooth AR-1-324/10 is 11.50 mm wide, 9.99 mm tall with seven to eight mesial denticles and five to six distal denticles ( ). A large right tooth AR-1-564/10 is 17.23 mm wide and 12.95 mm tall with eight to nine mesial denticles and ∼six to seven distal denticles ( ).
The isolated dentary teeth ( ) are identical to the maxillary teeth and have a weak lingual cingulum and a strongly developed labial cingulum. The best preserved tooth AR-1-3700/31 is 14.03 mm wide and 12.69 mm tall with eight to nine mesial denticles and six to seven distal denticles ( ). The largest dentary tooth AR-1-3650/31 is 16.58 mm wide and 13.50 mm tall ( ).
With their relatively large size and well-developed cingula, the teeth of Europelta are most comparable to those of other nodosaurids [72]. They similar to the teeth of Cedarpelta, Sauropelta [34], [97], [102], Edmontonia and Panoplosaurus [72], but are not as high crowned as in the Jurassic ankylosaurs Sarcolestes and Priodontognathus [103], the Jurassic polacanthids Gargoyleosaurus [93] and Mymoorapelta (Kirkland, pers. obs.), the nodosaurids Peloroplites [84] or Hungarosaurus [33]. Additionally, the large teeth of Gobisaurus are more inflated labiolingually than in Europelta and other ankylosaurs. The teeth of Gastonia and putative Polacanthus teeth are also inflated, but are smaller proportionally [83], [103]. The teeth of Europelta differ from an isolated tooth from the Cenomanian of France which is about half the size, and proportionally is longer mesiodistally with more deeply divided denticles forming ridges on the labiolingual surfaces of the tooth [104]. Likewise, lower Cenomanian teeth assigned to “Acanthopholis” have more deeply divided denticles in what is a proportionally taller tooth [17]. The teeth of Struthiosaurus languedocensis [31] from the lower Campanian of France also differ in size and in having longer, lower tooth crowns.
Axial skeleton
There are numerous ribs and vertebrae preserved from the holotype (AR-1/10) and the paratype specimen (AR-1/31). Vertebral measurements are presented in .
Table 1
Europelta VERTEBRAL MEASUREMENTS IN MMAnteriorPosteriorOverallNeuralNeuralTotalNeuralTransverseTransverseCentrum FaceCentrum FaceCentrumCanalCanalVertebralSpineProcessesProcessesWidthHeightWidthHeightLengthWidthHeightHeightHeightWidthLength(above canal)AR/10* estimated Cervical Vertebrae AR-1-431109.278.8--*85.230.630.6186.186.4203.879.2AR-1-449100.174.3--66.331.622.4185.590.9198.261.1AR-1-53394.969.9--*81.7*23.1*22.9*218.9*133.2*203.172.5AR-1-637*81.5*60.5*78.3*60.5*96.8*25.8*17.1---47.3AR-1-63893.168.9*85.0*73.875.620.630.3--*160.286.7AR-1-64973.270.199.961.262.028.819.1---77.4AR-1-650*81.4*57.780.662.5*61.0*26.8*14.4122.5*56.0*104.1*29.0 Dorsal Vertebrae AR-1-154----79.4------AR-1-155*60.0*69.875.5*68.4*79.2--*159.5129.3--AR-1-32289.976.394.678.982.514.824.0-133.1-*85.2AR-1-43091.477.797.583.684.6*20.1*25.2222.9-*175.8*76.5AR-1-44890.378.595.579.090.7*16.8*23.7--*120.0*73.3AR-1-47891.478.294.884.0*86.116.722.1219.8139.2114.485.7AR-1-53598.581.792.8*82.093.522.426.3239.9-142.591.2AR-1-556*88.4*79.2*83.0*76.3*70.7*21.0*27.4---*89.8 Caudal Vertebrae AR-1-56276.273.881.679.372.414.326.0178.885.8193.964.8AR-1-63582.279.892.492.179.419.426.7192.280.7240.674.0AR-1-63682.780.789.194.2*66.2*23.10*21.3*193.0*88.8*211.377.4AR/31 Cervical Vertebrae AR-1-3632*76.5*63.4*66.0*63.9*52.9*9.0*18.5*154.8*68.2*139.6*60.6AR-1-365767.953.576.0-*51.813.721.1--*151.650.1AR-1-366269.160.3*67.160.2*53.3------AR-1-3671*69.0*49.5*78.0*52.4*53.9*25.2*11.8*134.1*57.3*120.5*41.3AR-1-3676*52.1*55.6*60.3*60.0*60.4*8.8*19.5*136.1*51.1*89.8*26.9 Dorsal Vertebrae AR-1-348965.659.365.061.879.0*12.0*15.1178.9105.0-68.0AR-1-358675.661.672.761.854.614.319.6157.574.7140.962.9AR-1-363376.460.167.058.662.713.118.9178.594.7*139.173.2AR-1-367268.752.777.157.873.5------AR-1-367366.660.566.555.472.5*11.8*15.5--*119.658.0AR-1-3674*59.1*65.7*53.7*63.8*72.9--*168.5*88.7-85.1AR-1-3675*64.656.966.763.266.315.922.3*171.8*104.4*133.974.2AR-1-370467.062.8*64.7*59.979.1-*14.4--*154.663.3 Caudal Vertebrae AR-1-295031.125.328.724.250.26.14.937.09.0--AR-1-3204---------49.8-AR-1-320639.029.535.430.050.6*8.04*7.0----AR-1-324343.038.838.731.751.05.36.047.320.7--AR-1-326545.1*30.045.232.052.6------AR-1-3348*60.749.5*55.3*45.5*53.7--73.115.1*99.038.2AR-1-339842.534.632.435.351.93.36.250.113.242.5-AR-1-347848.934.846.738.252.24.88.052.2-46.1-AR-1-361551.437.149.026.156.16.511.247.7*8.4--AR-1-3616*48.9*42.8*45.2*39.7*56.9--*58.9-49.3-AR-1-371430.223.7--42.15.74.8----AR-1-3715--25.622.237.7--31.6---AR-1-3716--*48.143.9---*70.115.5--AR-1-371761.440.058.139.654.27.9*5.1--95.429.7
The complete atlas (AR-1-649/10) from the holotype has a total width of 195.6 mm ( ). The neural arch is divided dorsally with the left side fused to the centrum and the right side unattached. The anterior face of the atlantal intercentrum is 73.7 mm wide by 71.7 mm tall and its posterior face is 99.9 mm wide by 61.2 mm tall with a length of 62.0 mm. The axis is not present in either associated skeleton.
There are five post-axis cervical vertebrae (AR-1-431/10, 449, 533, 637, 650) preserved from the holotype skeleton ( ) and five from the paratype skeleton; of which four are illustrated (AR-1-3586/31, 3632, 3671, and 3676) ( ). Overall, they are typical of most other described ankylosaur cervical vertebrae. The centra are amphicoelus, wider than tall, anterorposteriorly short, and medially constricted. Anterior and mid-cervical vertebrae have the anterior faces of the centra dorsally elevated relative to the posterior faces. This is in contrast to the posterior cervical centra which have horizontally aligned faces. The ventral sides of the anterior centra are characterized by two anteroposteriorly-oriented paired fossae separated by a low keel ( ), as observed in the primitive nodosaurid Animantarx [97]. The dorsal ends of the neural spines are expanded transversely. AR-1-638/10 may either be the last cervical vertebra or the first dorsal vertebra based on the position of the parapophyses.
There are two complete cervical ribs preserved for the holotype. AR-1-450/10 is a relatively anterior cervical rib ( ) and AR-1-4452/10 is a posterior cervical rib. There is no evidence of fusion of cervical ribs to the cervical vertebrae as in the ankylosaurid Saichania [105], [106] or Ankylosaurus [107]. The cervical ribs are Y-shaped overall and much like the cervical ribs of other ankylosaurs such as Silvisaurus [76], [78], [82].
Several amphiplatan to amphicoelus dorsal vertebra are preserved: eight for the holotype AR-1/10 and nine for the paratype AR-1/31. The diapophyses originate at the level of the post-zygopophyses at the dorsal extent of the neural canal. The more anterior vertebrae have large cylindrical amphiplatan centra which lack a constricted ventral keel with circular neural canals and fused ribs (AR-1-448/10, 478, and 535). The broad transverse processes are T-shaped in cross-section and angled dorsally, unlike the laterally directed transverse processes in Polacanthus [10], [38]. Two dorsal vertebrae from the holotype appear to be pathological with the centra overgrown by about 0.5 cm of lumpy reactive bone ( ). One of these pathologic vertebrae (AR-1-535/10) has fused ribs ( ) although the other (AR-1-430/10) does not ( ). Two additional dorsal vertebrae (AR-1-478/10, 448) with fused ribs are not pathologic ( ). More posterior dorsal vertebrae have shorter, taller, more medially constricted centra, laterally compressed neural canals, more dorsally directed transverse processes, and lack fused ribs (AR-1-155/10, 322, and 556). The neural spines are thin and rectangular with narrowly expanded dorsal ends as in Sauropelta [99]. The neural spines are oriented dorsally as opposed to the posteriorly inclined neural spines of some other ankylosaurs such as Sauropelta [97]. None of the paratype vertebrae (AR- 1-3489/31, 3633, 3662, 3672, 3673, 3674, 3675, 3677 and 3704) have fused ribs ( ), suggesting that this character is ontogenetic because the paratype AR-1/31 represents a somewhat smaller (and presumably younger) individual than the holotype AR-1/10. More expanded neural spines are present in Shamosaurus [91].
There are a number of rib fragments preserved with AR-1/10, but there are only three (AR-1-331/10, 333, 476) relatively complete ribs ( ). As with most other ankylosaurs, the ribs are sharply arched and L-shaped in cross-section proximally in anterior ribs and broadly arched and T-shaped in cross-section proximally in more posterior ribs.
The sacrum is not preserved in AR-1/10 other than an anteriormost centrum (AR-1-154/10) of the synscacrum ( ). However, for the paratype, AR-1-3466/31, there is a largely complete but fragmented synsacrum ( ) that includes an interpreted anteriormost synsacral centrum (AR-1-3451/31), more of the anterior synsacrum composed of two dorsal centra (AR-1-3450/31), four sacral vertebrae with the sacral ribs from the left side (AR-1-3446/31), two sacral ribs from the right side (AR-1-3452/31, 3460), and one caudosacral vertebra (AR-1-3512/31). Given that at least one intermediate and one anterior fused synsacral dorsal vertebra are missing, the vertebral formula for the synsacrum would be five or more dorsosacral vertebrae, four sacral vertebrae, and one sacrocaudal vertebra. The entire synsacrum would have been over 50 cm long and measures about 44 cm across the sacral ribs. The middle section of the preserved dorsal synsacrum thins anteriorly from about 7 cm wide to about 5.5 cm wide. It then expands again anteriorly as indicated by the anteriormost centrum of the synsacrum. This differs from the sacrum of Euoplocephalus [108] and Saichania [106] in which each centrum making up the synsacrum is constricted medially. The sacrum is distinctive in being more strongly arched anteroposteriorly than other described ankylosaur sacra. The neural spines are dorsoventrally shorter than the height of the centra and are fused into a vertical sheet of bone along the length of the sacrum. The caudosacral neural spine is longer and unexpanded, transitional in form between the sacral neural spines and those of the proximal caudal vertebrae. The neural spines are broken off the anterior end of the synsacrum. The ventral side of the sacrum and anterior synsacrum is longitudinally depressed. The distal ends of the sacral ribs are expanded and the most robust medial sacral rib is about 50% taller (9.4 cm) than wide (6 cm) at its attachment with the ilium. There is no sign of expansion of the dorsal termination of the neural spine on the sacrocaudal vertebra. Additionally, the caudal rib is reduced compared to the sacral ribs.
The sacrum of Struthiosaurus languedocensis [31] is similar overall, but based on the description is not so strongly anteroposteriorly arched as in Europelta. Similarly, the sacrum of Hungarosaurus, as exhibited at the Hungarian Natural History Museum, appears to be moderately arched. The moderate angulation of the faces of the sacral centra (somewhat wedge-shaped in lateral view) in Anoplosaurus [17] indicates that a moderately arched sacram may have been present in this taxon as well. Among North American nodosaurids, we have observed only a moderate anteroposteriorly arching of the synsacrum of Silvisaurus, which appears to be restricted to the posterior part of the sacrum and two sacrocaudals. In other ankylosaurs, the downward flexure of the tail from the hips is taken up in the proximal caudal vertebrae as in Mymoorapelta [84], [109] and Euoplocephalus [70], [82].
Only three proximal caudal vertebrae (AR-1-562/10, 635, 636) are present ( ). The proximal-most caudal vertebrae are not preserved for the holotype. The preserved vertebrae probably represent caudal vertebrae positions in the interval of about 3–7. The centra are anteroposteriorly shorter than dorsoventrally tall and somewhat wedge-shaped in anterior and posterior views. The posterior chevron facets are well developed. The neural spines are inclined posteriorly and the dorsal ends of the neural spines are only slightly expanded transversely as in Gargoyleosaurus [95] and some other ankylosaurs such as Cedarpelta [86], Edmontonia [110], Hungarosaurus [32] and Euoplocephalus [70], [82]. The neural spines are strongly expanded in most polacanthids such as Mymoorapelta [84], [109], Gastonia [83], and Polacanthus [10], and some North American nodosaurids such as Sauropelta [99], and Silvisaurus [76]. The neural spine of AR-1-562/10 is broken, erroneously giving it the appearance of being strongly inclined posteriorly. The caudal ribs (transverse processes) in Europelta originate high on the sides of the centrum and angle ventrally proximal to flexing laterally, giving them a dorsally concave profile in anterior view like Hungarosaurus, Struthiosaurus, and Peloroplites, and unlike the ventrally flexed caudal ribs of many polacanthids [10], [84], [109] and the caudal vertebra assigned to “Acanthopholis” [17] or straight caudal ribs of Gargoyleosaurus [95], Cedarpelta, Peloroplites [86], and Edmontonia [87]. The proximal caudal ribs of Hylaeosaurus differ in being swept back posteriorly [111]. The lateral terminations of the caudal ribs do not expand dorsoventrally as they do in Peloroplites [86] and Struthiosaurus, which actually appear to bifurcate [25], [26].
Additionally, there are four chevrons preserved from about the same region of the tail (AR-1-560/10, 561, 569, and 4451) of which three are illustrated ( ). The proximal chevrons are approximately as long as the neural spines as in most other ankylosaurs. They are relatively straight and expanded into teardrop shapes distally in lateral view. Unlike in many ankylosaurs, there is no fusion of proximal chevrons to their respective caudal vertebrae as in Pinacosaurus and Saichania [105], [106], Ankylosaurus [107], [112], and Edmontonia (ROM 1215) [87].
Several more distal caudal vertebrae are preserved in the paratype. The two most proximal of these (AR-1-3348/31, AR-1-3717/31) have centra of nearly equal height, width, and length, with a ventral groove, and caudal ribs shorter than the diameter of the centrum that extend laterally and angle posteriorly ( ). The chevron facets are well developed with the posterior facets more strongly developed than the anterior facets. The neural spines are not developed and the zygapophyeses only extend a short distance beyond the anterior and posterior margins of the centra. These vertebrae are interpreted to represent mid-caudal vertebrae. Two more distal mid-caudal vertebrae (AR-1-3616/31, AR-1-3716/31) are similar in morphology except that the caudal ribs are reduced to anteroposteriorly directed ridges on the lateral margins of the centra ( ). Their neural spines incline posteriorly, merging with the postzygapophyses as posterior processes extending laterally past the faces of the centra to overlie and articulate between the paired prezygapophyses of the immediatly distal vertebra. This morphology is retained in the distal caudal vertebra. More distally, as in AR-1- 2950/31, 3206, 3243, 3265, 3478, and 3615, the caudal ribs are lost and the centra become more elongate ( ). Unlike many ankylosaurs, the faces of the centra maintain a well-rounded to heart-shaped surface distally down the caudal series [82]. For many of these vertebrae, ventrally anteroposteriorly elongated skid-shaped (inverted T) chevrons are fused to the posterior chevron facets. Fusion of distal chevrons to their respective vertebrae is widespread among ankylosaurs [84], [106], [110] although it is not present in some, such as Nodosaurus [113]. One pair of distal caudal vertebrae is fused by their mutually shared chevron ( ) such as has been documented in Mymoorapelta [84]. The most distal four caudal vertebrae ( ) and their chevrons are fused together in AR-1-3204/31 to form a tapering, terminal rod of bone at the end of the tail somewhat similar to that of Sauropelta [71].
Pectoral Girdle
Parts of the right scapulocoracoid are preserved. A portion of the distal scapular blade (AR-1-429/10) is preserved with a portion of the distal ventral margin missing with a curved section broken away. There is no evidence of any distal expansion of the scapular blade as in many nodosaurids [94].
The coracoid (AR-1-657/10) is preserved with only the most proximal portion of the scapula fused on ( ). It appears to have been sheared off just dorsal to the suture between the coracoid and the scapula, perhaps in the process of removing the overlying coal seam. The coracoid is relatively equidimensional (201.3 mm long by 186.5 mm tall) relative to the elongate coracoids characteristic of many other nodosaurids [114] such as Peleroplites [86], Texasites [77], [115], and Animantarx [97]. The medial surface is concave and the lateral surface is convex giving it a bowl-shaped appearance. The ventral margin is evenly convex as in many polacanthids and nodosaurids and there is no anteroventral process as in all ankylosaurids, including Shamosaurus [91], [94]. The articular surface of the ventrally directed glenoid is wide, bounded by a flange that extends beyond the medial surface of the coracoid.
Both xiphisternal plates are preserved ( ). The best preserved xiphisternal is approximately 350 mm long. They appear to be arcuate flat bones. Xiphisternal plates are only known in a few nodosaurids, but those of Europelta, whereas similar in overall shape to other nodosaurid xiphisterna, are not fenestrate or scalloped along their margins as in North American nodosaurids for which they are known [82], [87], [116].
Forelimb
Parts of both humeri are preserved. The right humerus (AR-1-655/10) is represented by the proximal end ( ). It is 249.2 mm wide with a well-developed proximal head 91.9 mm wide that extends onto the posterior side of the humerus. Distinct notches separate both the laterally directed deltopectoral crest as in nodosaurids such as Sauropelta [70], [71], [99] and the internal tuberosity from the humeral head. The deltopectoral crest extends lateraly from the humerus and is not flexed anteriorly as in polacanthids and ankylosaurids [94].
The left humerus (AR-1-327/10) is represented by a midshaft for which both the proximal and distal ends appear to have rotted off and the core of the shaft has rotted away ( ). The shaft is deeply waisted relative to the proximal and distal ends. Although relatively uninformative, enough of this humerus is preserved to indicate that the deltopectoral crest would have made up less than 50% of the length of the humerus as in nodosaurids [71], [117] and in the basal ankylosaur Mymoorapelta (Kirkland, pers. obs.) compared to the longer deltopectoral crests of ankylosaurids [70], [71]. Overall, the humerus of Europelta is similar in proportions to Niobrarasaurus [118], [119]. The wide proximal end of the humerus figured by Ősi and Prondvai [120] as cf. Struthiosaurus is similar to that of Europelta, whereas the humerus of co-occuring Hungarosaurusis is more slender proportionally.
Among the nine unguals preserved for AR-1/31, one specimen (AR-1-3711/31) may represent a manual ungual. It is more equidimensuional than the other eight more elongate unguals.
Pelvic Girdle
The right ilium of AR-1/10 is fused with its ischium and pubis (AR-1-479/10) which are flexed medially due to compaction ( ). The acetabulum is completely enclosed as in all derived ankylosaurs [70], [71], [82], [94], [108]. Only Mymoorapelta is known to retain an open acetabulum [84], [109]. The acetabulum is directed verntrally and is situated medially near the contact of the ilium with the sacrum so that the ilium extends far out beyond the acetabulum laterally for a distance nearly equal to its width. The lateral and anterior margins of the laterally oriented ilium are broken away. The prepubic portion of the ilium diverges from the midline of the sacrum at about 30 degrees and is thickened ventrally along its midline. Large, fairly equi-dimensional, closely appressed osteoderms (7-10 cm in diameter) cover the dorsal surface of the ilium posterior to and medial to the acetabulum. As discussed below, this morphology of sacral armor compares well with “Category 3” pelvic armor of Arbour and others [121]. Anteriorly, the smooth dorsal surface of the ilium is exposed. The pubis is fully fused to the anterior margin of the ischium with no visible sutures; its presence is indicated by a slot-shaped foramen along the anterior side of the ischium. This foramen represents the obturator notch between the postpubic process and the main body of the pubis as in Scelidosaurus and stegosaurs [122]. The distal end of the ischium is broken away. Additionally, AR-1-129/10 is a poorly preserved, proximal left ischium with the pubis fully fused to its anterior margin ( ).
Beyond some relatively uninformative fragments of the ilium ( ), AR-1/31 includes both the right (AR-1-3648/31) and the left (AR-1-3649/31) ischia with fully fused pubes ( ). Both exhibit the slot-shaped foramen along the anterior side of the ischium formed by the obturator notch. The proximal ends appear enrolled such that the anterior and posterior margins are nearly parallel due to compaction. Both display an anterior kink at their distal end as in Cedarpelta [86], [88], but overall are straight-shafted as in the Ankylosauridae [70], [82], [123] and the other European nodosaurids Struthiosaurus [31] and Hungarosaurus [32]. The distal end of the left ischium is the best preserved and measures 299.9 mm long along its anterior margin, including the fully fused pubis forming an ischiopubis. Given the asymmetry of the proximal end of the fused ischium and pubis and the position of the obturator foramen, it appears that the pubis still makes up some of the acetabular margin. The contact between the ilium and the fused ischiopubis is straight with about one-fourth to one-third of the acetabulum formed by the fused ischiopubis.
A straight ischium has been considered to be the primitive character state for ankylosaurs, with the bent ischium of Polacanthus and nodosaurids, a derived character [63], [82], [83], [94], [114], [123]. It is possible that as opposed to being primitive, a straight ischium may be secondarily acquired in the ankylosaurids and European nodosaurids. The only known ischium from the Jurassic ankylosaur (Mymoorapelta) is bent, a trait that is also observed in some stegosaurs such as Kentrosaurus [124]. Stegosaur ischia, even when straight, have an angular thickening near the mid-point of the posterior margin [124] that is shared by the polacanthids Mymoorapelta (Kirkland pers. obs.) and Gastonia [83]. Europelta is the oldest known ankylosaur preserving a straight ischium. The slight kink in the distal end of the ischium of Europelta suggests the straight ischium in European nodosaurids and ankylosaurids is achieved by shortening the ischium distal to the bend.
Hindlimb
The right femur, tibia, and fibula were closely associated ( ). The robust right femur (AR-1-3244/31) is 502.9 mm long and 178.9 mm wide at the proximal end and has been flattened anteroposteriorly, with the most distortion to the mid-shaft region. The femoral head is distinct with much of its articular surface directed dorsally and only somewhat medially. It forms an angle of about 115° with the long axis of the femur. The femoral head is directed more dorsally under the ilium in polacanthids [7], [12], [82], [95], [125], and several nodosaururids. In addition, the femoral head of Europelta is expanded such that it overhangs the femoral shaft both anteriorly and posteriorly. The greater trochanter is well demarcated from the femoral head by a constriction across the proximal end of the femur, and the anterior trochanter forms a ridge ventral to the greater trochanter that is fully fused to the femur. The robust fourth trochanter overlaps the midpoint of the femoral shaft and its midpoint is located proximal at the midpoint of the femur. Polacanthids and nodosaurid ankylosaurs have this configuration, whereas in ankylosaurids the fourth trochanter is distal to the middle of the shaft [63], [82], [95], [120], [125]. The distal end of the femur is flattened and forms a planar articular surface relative to the straight femoral shaft. The intercondylar notch is not expressed ventrally, and is better developed posteriorly than anteriorly
The right tibia (AR-1-3237/31) and fibula (AR-1-3238/31) were closely associated ( ) and post-depositionally compressed. Compression has distorted the distal end of the tibia such that the wide posterior surface is twisted counterclockwise in line with the wide lateral side of the anterior end relative to the orientation of the proximal and distal ends of the tibia in most other ankylosaurs, such as Mymoorapelta [84] (Kirkland, pers. obs.). The fibula was taphonomically displaced ventrally and with the ventral end rotated posteriorly relative to its position in life with the tibia.
The tibia is 458.8 mm long and robust for its entire length ( ) as in Cedarpelta [86]. The proximal end is 169.2 mm wide by 93.1 mm wide and its distal end is 146.8 mm wide by 70.2 mm. It is significantly more narrowly waisted in Mymoorapelta [84], Gastonia [83], Polacanthus [7], [12], [18], Sauropelta [69], [71], [99], [108], Peloroplites [86], and in Zhejiangosaurus [126] and ankylosaurids like Saichania [106]. The cnemial crest is broadly rounded. The even curvature of the distal end of the tibia suggests that the astragalus was fully fused to it with no evident sutural contact as in most ankylosaurs [63], [82], [121]. The astragalus is not fused to the distal end of the tibia in Mymoorapelta [84], Gastonia [83], Hylaeosaurus [11], and Peloroplites [86].
Generally, ankylosaurids have tibiae that are less than two-thirds the length of their femora, as opposed to nodosaurids which have proportionally longer lower leg elements [127]. With a tibia to femur ratio of 0.91, Europelta has the proportionally longest tibia of any ankylosaur for which this ratio is known. Both Cedarpelta and Peloroplites have relatively longer tibiae than other ankylosaurs [86], with a tibia to femur ratio of 0.82 in both. Peloroplites differs in its proportionally more narrowly waisted tibial shaft.
The fibula is 395.5 mm long ( ) and laterally flattened. The proximal end is not expanded anteroposteriorly, such that the slender fibula changes little in size and shape from the proximal to distal end. In lateral view, the proximal end is rounded and the distal end is concave. In cross-section, it is flattened medially and convex laterally. It is longer relative to the tibia than in most other ankylosaurs [108].
A calcaneum (AR-1-3289/31) was identified in association with the lower right leg of AR-1/31. It is laterally compressed, convex laterally and concave medially ( ). Its dorsal margin is flattened where it would articulate with the fibula. Calcanea are practically unknown in ankylosaurs, but one has been identified in the juvenile specimen of the derived ankylosaur Anodontosaurus [128]. The type of Niobrarasaurus coleii preserves an articulated lower hind limb, with an astragalus fully fused with the tibia and possessing an articulation with the distal end of the fibula and an unfused calcaneum of similar morphology to that of Europelta [118]. The calcaneum is fully fused to the distal end of the fibula in Saichania [106].
A number of metatarsals and phalanges are associated with AR-1/31. The metatarsals have subrectangular proximal ends, indicating that they were closely articulated in a well-integrated pes in life ( ). The pedal phalanges ( ) are short, as in other ankylosaurs. There are eight relatively large, elongate, spade-like unguals ( ) of a morphology similar to pedal unguals in other ankylosaurs in which the unguals are nearly as long as the digits[82], which indicates that portions of both feet are present in AR-1/31. We interpret that the pes of Europelta possesses four pedal phalanges as in most other nodosaurids [80]. Liaoningosaurus has three digits on the pes. The eight similar unguals are interpreted as pedal unguals and the smallest ungual ( ) is interpreted as an isolated manual ungual. The overall proportions of the preserved pedal elements are similar to those of Niobrarasaurus [119], which also has pedal unguals nearly as large as its metatarsals.
Armor
There was an abundance of dermal armor recovered with both AR-1/10 and AR-1/31. On comparison with the quarry maps, none of the osteoderms appears to be preserved in situ with any of the skeletal elements or with each other, and there is no fusion between any of the osteoderms recovered. Therefore, the armor has been divided into several broad morphotypes for the purpose of description and comparison to armor described for other ankylosaurs. Although morphotypes and terminologies have been proposed [129], [130], no system fits for all armor types in all ankylosaurs. A number of researchers have divided armor into types as in Type 1, 2, etc. [131]; for this discussion the armor types are alphabetized to ensure minimal confusion with previous descriptions. The term osteoderm is used to describe relatively larger dorsal and lateral armor elements with the presence of an external keel or tubercle, whereas the term ossicle describes relatively smaller dermal armor lacking a keel, in the sense of Blows [130]. It is recognized that a consistent methodology for describing armor is achievable, but must be done within a phylogenetic framework to be of maximum utility.
Osteoderm surface texture may be broadly useful in differentiating ankylosaurids from nodosaurids [132], [133]. The vast majority of the osteoderms examined in Europelta has a moderately rugose texture with sparse pitting more in keeping with nodosaurids and basal ankylosaurids rather than more derived ankylosaurids. Whereas histological studies have proven useful in the study of thyreophorans [132], [134], [135], that is beyond the scope of this study.
It is noteworthy that no portions of distinct cervical rings were recovered, although cervical vertebrae are known for both skeletons of Europelta. Additionally, only one spine from the cervical or pectoral region was tentatively identified. We postulate that these elements were lost through the process of coal removal or may have been taphonomically removed from the skeletal associations. Only the discovery of additional specimens of Europelta can further reveal the presence of cervical half-rings.
Type A armor
An isolated fragmentary spine (AR-1-128/10), possibly from the cervical or pectoral region, is recognized from the holotype ( ). It appears to represent only the anterior half and may have been cut in two as the overlying coal was removed. This sharp, broken margin reveals an asymmetric, Y-shaped cross-section. The base flares more and is is less excavated than in a Type 2 caudal plate, suggesting that it was positioned on a broad flank of the body. From the possible anterior margin, the spine slopes posteriorly 15 cm to the broken margin in a gradual arc. There is no indication that the spine could not have been longer. The spine is compressed as in the cervical spines of Sauropelta [77], [99] and Edmontonia [110], [136], and the pectoral spines of Gastonia [83] and Polacanthus [7], [10]. The base is asymmetrical in a manner similar to the elongate osteoderms in Mymoorapelta [84], with one side of the base extending lower anteriorly and the other posteriorly. There is no evidence of a basal plate incorporated into fusion of the cervical half-ring as in mature ankylosaurs like Mymoorapelta [84] Gargoyleosaurus [85], [95], Gastonia [83], Polacanthus [10], [130], and Sauropelta [77], [99]. This may relate to the anchoring of larger elements into the dermis in Gastonia and Polacanthus [130]. We tentatively interpret AR-1-128/10 as a pectoral spine. However, if the complete element extends beyond the break for more than twice the length of the preserved portion, it would fall into the category of Type B armor, although that is unlikely because it is more massive form than the Type B elements.
Type B armor
Dorsoventrally compressed, hollow, asymmetric-based plate-like osteoderms with sharp anterior and posterior edges and lateroposteriorly directed apices are identified for AR-1/10 ( ) and AR-1/31 ( ). Similar large osteoderms have been described as caudal plate ostederms in Mymoorapelta [84], [109], Gargoyleosaurus [85], [95], Gastonia [83], and Polacanthus [8]-[10], [38], [130]. Similar, more anterorposteriorly symmetrical caudal plate osteoderms are also known in Minmi [137], [138] and several Asian ankylosaurids [131]. The few plate-like osteoderms of this morphology that are identified in Europelta are mediolaterally shorter and anteroposteriorly longer with a more posteriorly swept apices. Two pairs of similar plates are known for the holotype of Sauropelta (AMNH 3032), with one of the larger plates being illustrated [99]. One plate from the Yale collections of Sauropelta has a unique double apex (YPM 5490). Given the rarity of Type B armor in Sauropelta and Europelta we hypothesize that caudal plates in these nodosaurids ran down the sides of the tail but decreased in size more rapidly, such that long-keeled osteoderms of Type E morphology made up the lateral armor down most of the length of the tail. It is also possible that these large plate-like osteoderms were on the lateral margin of the sacrum as has been documented by Carpenter and others [106] in Saichania. Struthiosaurus preserves several osteoderms of this morphology that have been reconstructed as in Polacanthus as being medial, dorsally-projecting caudal osteoderms [25], [26]. The relative rarity of these plate-like osteoderms suggests that they were restricted to the base of the tail as well.
Type C armor
Both AR-1/10 ( ) and AR-1/31 ( ) preserve fairly large (∼15–25 cm long) subrectangular to subtrapezoidal, solid osteoderms with low, evenly developed keels running down the long axis of the osteoderm either medially or to one side of the mid-line. Their distal and medial surfaces are subparallel and the entire plate may be slightly flexed across the short axis perpendicular to the crest. The straight, longer margins of these plates appear to have been tightly affixed but not fused to adjoining osteoderms. Armor of Type C morphology is not common but is most similar to medial cervical osteoderms of half-rings, and most distinctively, across the mid-line of the pectoral region in some nodosaurids such as Stegopelta [138], Niobrarasaurus [140], [141], Panoplosaurus [74], [101], and Edmontonia [74], [110].
Type D armor
Both AR-1/10 and AR-1/31 preserve large (∼10-20 cm long) asymmetric, diamond ( ; ) to tear-drop shaped ( ) osteoderms with a long keel rising to an apex medially to posteriorly and in some specimens extending past the posterior margin of the base. They are distinguished from Type E osteoderms because they are wider than 50% of their length. The wider osteoderms are thinner and more solid than the narrower osteoderms with small pockets under the apices. The more diamond-shaped forms may be more closely appressed to each other in anterior bands similar to Type C armor.
Type D Armor is widely known in the nodosaurids such as Sauropelta [99], Panoplosaurus [101], and Edmontonia. Gastonia is documented to have similar armor [142], although more solid in cross section with less basal excavation, which occurs in oblique rows anterior to the sacrum with each osteoderm separated by a single row of small Type H ossicles. This pattern is similar to the dorsal dermal ornamentation documented for the ankylosaur Tarchia by Arbour and others [130], except that in Tarchia most of the intermediate scales lacked ossified cores. Similar armor is known from the lateral sides of the legs in some ankylosaurs such as Saichania [106].
Type E armor
Both AR-1/10 and AR-1/31 preserve large (10-15 cm long) moderately asymmetric osteoderms more than twice as long as wide with a long keel higher on the assumed posterior end ( ; ). These osteoderms have proportionally more deeply excavated bases than Type D armor, have chevron-shaped cross-sections, and are distinguished from Type D armor by their width being less than 50% of the length. Type E armor is gradational with Type D armor ( ; ) and may represent lateral or distal armor from the trunk of the body and along the sides of the tail. This armor type is present in Sauropelta [99] and Texasetes [115]. Similar armor is present on the sides of the limbs in Scelidosurus and Saichania [106].
Type F armor
Medium to large (∼5-15 cm long) oval to circular osteoderms of low profile with a median keel extending into an apex near or overhanging the posterior margin of the osteoderm are represented in both AR-1/10 ( ) and AR-1/31( ). The basal surface of the osteoderm is generally solid except for a small pocket under the apex, reminiscent of Type D armor. Less commonly, the base may be more extensively excavated. Armor of this morphology is abundant in many nodosaurids and makes up the major elements of the armor of Sauropelta anterior to the sacrum in AMNH 3036 [142] and is present in Panoplosaurus [101]. These osteoderms may reside within more expansive spaces among the larger dorsal armor as in Edmontonia (AMNH, 5665) and the polacanthids [81], [82], [93], [107], or may be major armor elements on the posterior portion of the sacrum as in Sauropelta (AMNH 3036). They may also lie on the tail between the Type B caudal plate-like osteoderms, or could be arranged along the lateral side of the limbs as in Saichania [106].
Type G armor
One piece (AR-1-192/10) of flat, oval to subtriangular armor (AR-1-192/10) from AR-1/10 is about 12 cm long and 7 cm wide and is about 0.5 cm thick throughout ( ). A pair of similar, osteoderms from the Sauropelta specimen AMNH 3032 was curated with a note from the collector, Barnum Brown, stating that these distinct osteoderms were associated with the forelimbs. Therefore, we suggest a similar position for Type G armor in Europelta.
Type H armor
Small (∼1-4 cm long) solid ossicles are abundant, with 71 examples from both AR-1/10 ( ) and AR-1/31 ( ) illustrated. These ossicles range in shape from round, to oval and even irregularly shaped, and are probably filling in the spaces between larger osteoderms. Small interstitial ossicles are not known for every ankylosaur taxon, but appear to be present in many nodosaurid taxa such as Sauropelta [99], [143] and Edmontonia [74], [136], in polacanthid ankylosaurs such as Gastonia [83] and in some ankylosaurids such as Tarchia [131], in which epidermal scales interstitial to osteoderms do not preserve deeper, interstitial ossicles. Their absence may be real, in that they never form deep to the epidermal scales, taphonomic, in that they are selectively transported away because of their small size and low density, or ontogenetic; in that they only ossify late in ontogeny. The surface texture of Gastonia ossicles is smoother than those of Europelta.
Sacral armor
Armor is present on the posterior margin of the ilium AR-1-479/10. It is composed of large, subequal-sized (7-10 cm) osteoderms that are tightly sutured together ( ) as in the poorly known Stegopelta [139], Nodosaurus [113], Aletopelta [127], and Glyptodontopelta [132], [144]. These low-relief ossicles lack a central apex or keel. The boundary between the margins of the osteoderms and the area devoid of osteoderms on the ilium is sharply demarcated along the margins of unbroken osteoderms, suggesting the armor was not coossified as in Aletopelta [127] and unlike the fully fused sacral armor in the polacanthids Polacanthus and Gastonia [63], [83]. This form of pelvic armor fits that of Arbour and others' Category 3 pelvic armor [121].
Additionally, there is a unique osteoderm AR-1-653/10 that has a large, posteriorly-curved, plate-like keel extending out from the surface that, considered in isolation, is comparable in size and morphology to Type B armor ( ). The base is smooth and gently convex, suggesting it may have been closely appressed to the more anterior portion of the ilium. In overall morphology, this large osteoderm is comparable to the spine-bearing armor plate-like osteoderm identified in Hungarosaurus and interpreted to be present in Struthiosaurus [33].
Unique armor pieces
Some irregularly shaped armor specimens are not represented by more than one element among this material or in the armor from other taxa. At this time, we can offer no positional interpretation of this armor. AR-1-447/10 is an irregular mass of what we interpret as an osteoderm, although it could be sacral armor ( ). AR-1-438/10 is a small, cap-shaped shaped with a small excavation in the center of the external surface ( ). Two small, deeply basally excavated, oval osteoderms ( ) were collected from AR-1/31(AR-1-3239/31, 3721). These osteoderms lack the external excavation.
Discussion
Europelta ( ) can be distinguished from any of the ankylosaurs assigned to the Polacanthidae (sensu Kirkland's Polacanthinae [83] and Carpenter's Polacanthidae [63] from the Upper Jurassic and Lower Cretaceous as defined by Yang and others [64]; see Terminology) by its rounded, tear-drop shaped skull and a suborbital horn developed on the posterior portion of the jugal and the quadratojugal posterior to the orbit, as opposed to a triangular-shaped skull that is widest at the posterior margin and a suborbital horn developed exclusively on the jugal (as seen in polacanthids). Post-cranially, it can also be distinguished from polacanthids, by its elongate lower hind limbs, the apparent rarity of cervical, pectoral, and thoracic spines, and reduction in the number of caudal plate-like osteoderms. Likewise, it has an abundance of Type D, asymmetric, tear-drop shaped osteoderms like those observed in many nodosaurids and absent in all polacanthids.
Europelta is also distinguished from derived ankylosaurids by its weakly ornamented teardrop-shaped skull in which the lower temporal opening is visible in lateral view. The absence of a tail club also distinguishes the taxon from these ankylosaurids. More basal “shamosaurine grade” ankylosaurids [63], [86] are more similar to Europelta, but also have the lower temporal openings completely obscured laterally by expanding the lateral margin of their skulls. “Shamosaurine grade” ankylosaurids also possess skulls that are approximately as wide mediolaterally between the orbits as they are across the posterior margin.
Europelta shares a number of derived characters with nodosaurids [71], [72], [83], [94], [114]. It has a tear-drop shaped skull that is longer than wide with its greatest width dorsal to the orbits, whereas the short, boxy skulls of Minmi and all anklosaurids are essentially as wide at the posterior edge of the skull, as are the elongate skulls of “shamosaurine-grade” ankylosaurids. Grooves in the remodeled textured skull roof define epidermal scale impressions, with the largest covering the frontoparietal area. Although poorly preserved, the laterally extensive pterygoids are pressed up against the anterior face of the braincase. All known nodosaurid scapulae have a prominent acromion process extending on to the blade of the scapula that terminates in an expanded knob. Unfortunately, this portion of the scapula is as yet unknown in Europelta.
Some character states considered typical of nodosaurids are absent in Europelta. Instead of having a distinct hourglass-shaped palate typical of nodosaurids [70], [71], [82], [83], [114], the upper tooth rows show less lateral emargination and diverge posteriorly. This is also true of Silvisaurus, which also shares an expanded lateral wall of the skull [76], [77]. The coracoid of Europelta is nearly as long as it is tall, whereas in other nodosaurids, for which the corocoid is known, it is expanded anteriorly and longer than tall [71], [72], [83], [94], [114].
The only other Early Cretaceous nodosaurid to have large cranial scales as in Europelta is Propanoplosaurus, known only from an embryonic to hatchling specimen from the base of the Potomac Group of Maryland [145]. However, only the anterior cranial scales are well defined in Propanoplosaurus, whereas only the posterior scale pattern in Europelta. The unusual preservation and extremely small size of Propanoplosaurus lead us to suspect that the fossil preserves the actual scales overlying the skull and not the remodeled skull roof, because this is such a young specimen and remodeling of the cranial bones is not expected to have occurred so early in ontogeny [129], [146].
Additionally, a number of important characters traditionally used to define nodosaurids are not known in Europelta, as yet, because of the missing anteroventral half of the scapula and the absence of premaxilla and surangulars. Thus, the presence absence of premaxillary teeth, if the tooth row joined the margin of premaxillary beak, the morphology of the naris, the height of the coronoid process, and the morphology of the acromion process are unknown for Europelta.
Europelta is distinguishable from European nodosaurids from the Albian through the Cenomanian. The juvenile Anoplosaurus from the Albian Gault Clays of southern England differs in a number of characters, such as possessing a proportionally longer coracoid, a narrower proximal end of the humerus, and a femur with a separate anterior trochanter [17] although the latter two characters are consistent with the juvenile nature of Anoplosaurus. No pectoral spines of the morphology described for “Acanthopholis” from the Cenomanian Lower Chalk in southern England by Huxley [13] are known in Europelta. Additionally, the tall teeth assigned to “Acanthopholis” are distinct in the long apicobasal ridges extending from the denticles to the root on medial and lateral faces of the teeth, and in the presence of caudal ribs that extend laterally and flex ventrally, whereas the caudal ribs in Europelta extend ventrolaterally and flex laterally [16], [17]. Europelta is like other Late Cretaceous European nodosaurids in having a short symphysis for the predentary, a mediolaterally wide and anteroposteriorly thin quadrate, an anteroposteriorly arched sacrum, and a straight ischium [21], [32].
The domed skull and elongate cervical vertebrae in Struthiosaurus clearly distinguish it from Europelta. Likewise, Hungarosaurus also has more elongate cervical vertebrae [32]. Both Hungarosaurus and Struthiosaurus possess a pair of spines on the anterior portion of the pelvis [33], whereas we interpret the presence of a pair of upright plate-like armor elements in this position in Europelta ( ).
The lateral wall of the skull in most North American nodosaurids is typically narrow [82], whereas in Europelta it is relatively wider, although a broad notch along its posterior margin permits the caudal margin of the lower temporal opening to be observed in lateral view. This morphology in Europelta is similar to that in the nodosaurids Silvisaurus [76], [77] and Peloroplites [86]. Although, the skull of Struthiosaurus transylvanicus is highly reconstructed [22], it appears that the lateral wall of the skull is expanded laterally, whereas not completely obscuring the lower temporal opening. This character state is not known in other species of Struthiosaurus, but appears to be moderately developed in Hungarosaurus [32].
Comparisons of Europelta with the Asian”nodosaurids” Zhongyuansaurus [93] and Zhejiangosaurus [126] from the lower Upper Cretaceous of China hinges partially on the question of whether those taxa have been validly referred to Nodosauridae. Carpenter and others [86] noted that the skull of Zhongyuansaurus is morphologically similar to that of a “shamosaurine-grade” (like Shamosaurus and Gobisaurus) ankylosaurids and was the first shamosaurine-grade ankylosaurid documented to not have a tail club. However, its distal tail is modified into a stiffened structure of the same morphology as the “handle” of the tail club in more derived ankylosaurids [147], [148]. Zhejiangosaurus was assigned to the nodosaurids based on characteristics of the femur and sacrum, together with the lack of a tail club [126]. We hypothesize that it lacked a knob as in basal ankylosaurids, polacanthids and nodosaurids because ankylosaurids with a full tail club have distal free caudal vertebrae bearing caudal ribs at the base of the handle. Most of the distal caudal vertebrae of Zhejiangosaurus have raised ridges on the sides of the centra as in the distal vertebrae of polacanthids and nodosaurids. Additionally, whereas the position of its most proximal preserved caudal vertebrae is not known, morphologically, they do not appear to represent the most proximal caudal vertebrae. Thus, while Zhejiangosaurus' 13 preserved caudal vertebra are more than the number of free caudals preserved in most ankylosaurs with tail clubs (10 in Saichania [106] and Dyoplosaurus [148]), the total number of free caudals in its tail would appear to be more than the 14 in Tarchia [130] and 15 in Pinacosaurus [129]. Unlike nodosaurids, Zhejiangosaurus has an exceedingly low ratio of femur to tibia length of 0.46 similar to that of with ankylosaurids and polacanthids rather than nodosaurids. Dongyangopelta [149] was described as a second nodosaurid from the same area and stratum as Zhejiangosaurus, which was found to be its sister taxon in their phylogeny [149]. With few overlapping elements, we feel that the proposed differences between these taxa may be due to preservation, individual variation, or ontogeny. Additionally, given the presence of a pelvic shield and numerous caudal plate-like osteoderms in Dongyangopelta, we suggest that both specimens may pertain to the same taxon and represent the first polacanthid described from Asia. Given the recent description of the polacanthid Taohelong from the upper portion of the Lower Cretaceous of Gansus Province in western China [64], this hypothesis has added support. We also do not think that the partial ankylosaur skull reported from the lower Upper Cretaceous of Hokkaido, Japan [150] can be diagnosed as a nodosaurid with any confidence at this time, due to the incomplete nature of the specimen. Thus, we do not presently recongnize the presence of true nodosaurids in Asia.
In his seminal paper defining a bipartite division of the Ankylosauria into Ankylosauridae and Nodosauridae, Coombs [71] hypothesized that Acanthopholis (as a nomen dubium in which he would have included Anoplosaurus) and Struthiosaurus might represent a separate lineage of European nodosaurids. Unlike Hylaeosaurus (in which he included Polacanthus), these taxa had a well-developed supraspinus fossa developed anteriorly on the scapula as did all North American nodosaurids. This European lineage was hypothesized based on their small body size, presence of premaxillary teeth, and their possessing an unfused scapula and corocoid. Although, none of the characters are valid in defining such a group, our research on Europelta has resulted in supporting the taxonomic hypothesis of Coombs [71], [72] as correct, just for the wrong reasons.
Relationships to Other Taxa
We use Struthiosaurinae to define the clade of European nodosaurs. Nopcsa [25] proposed Acanthopholidae as a family of relatively lightly built thyreophorans, that included Acanthopholis ( = Anoplosaurus), Polacanthus, Stegopelta, Stegoceras, and Struthiosaurus. In 1923, he divided the Acanthopholidae into an Acanthopholinae and a Struthiosaurinae without comment [69]. Subsequently, he relegated the Acanthopholidae to a subfamily of the Nodosauridae, in which he also included Ankylosaurus and restricted the Acanthopholinae to Acanthopholis, Hylaeosaurus, Rhodanosaurus, Struthiosaurus, Troodon [26], [151]. This artificial grouping included a polacanthid ankylosaur [72], [83], a pachycephalosaur [152] and Acanthopholis, now considered a nomen dubium [17], [82]. Thus, the term Acanthopholinae is not acceptable for this newly recognized clade of nodosaurids. Thus, Struthiosaurinae is the next published term available to use for this clade and is derived from the first described and youngest member of this clade. Struthiosaurinae is defined as the most inclusive clade containing Europelta but not Cedarpelta, Peloroplites, Sauropelta or Edmontonia.
In order to determine the systematic position of Europelta, it was found that previous cladistic analyses [71], [72], [82], [83], [114], did not include many of the character states that we identify as significant in our research on Upper Jurassic and Lower Cretaceous ankylosaurs. A major weakness of these analyses is the limited recognition of postcranial skeletal and dermal characters that restricts the testing the phylogenetic relationships for taxa for which skulls are either poorly known or not known at all.
We present a character based definition of Struthiosaurinae as: nodosaurid ankylosaurs that share a combination of characters including: narrow predentary; a nearly horizontal, unfused quadrate that is oriented less than 30° from the skull roof, and mandibular condyles that are 3 times transversely wider than long; premaxillary teeth and dentary teeth that are near the predentary symphysis; dorsally arched sacrum; an acromion process dorsal to midpoint of the scapula-coracoid suture; straight ischium, with a straight dorsal margin; relatively long slender limbs; a sacral shield of armor; and erect pelvic osteoderms with flat bases. This suite of characters unites Europelta with the European nodosaurids Anoplosaurus, Hungarosaurus and all species assigned to Struthiosaurus. This clade of European nodosaurids has not been previously recognized. Europelta represents the earliest member of the European clade Struthiosaurinae.
Biogeogeographic Implications
The near simultaneous appearance of nodosaurids in both North America and Europe is worthy of consideration ( ). Europelta is the oldest nodosaurid known in Europe, it derived from strata in the lower Escucha Formation that is dated to early Albian. The oldest nodosaurid from western North America is Sauropelta, which in the lower part of its range is in the lower Albian Little Sheep Mudstone Member (B interval) of the Cloverly Formation in northern Wyoming and southern Montana [99], [153] with an ash bed 75 meters above the base near the top of the member providing an age of 108.5±0.2 Ma [154]. Nodosaurid remains from eastern North America appear to be older. Teeth of a large nodosaurid Priconodon crassus are known from the Arundel Clay of the Potomac Group [77], [155], which palynology dates as near the Albian-Aptian stage boundary [156]. The hatchling Propanoplosaurus is from the base of the underlying Patuxent Formation of the Potomac Group of Maryland, which has been dated as late Aptian [157], [158], making Propanoplosaurus the oldest known nodosaurid. Polacanthid ankylosaurs characterize pre-Aptian faunas in both Europe [11], [12], [37]-[39] and North America [70], [95], [159]. We have not been able to document a specific example of Polacanthus in the Lower Aptian Vectis Formation of the Wealden Group, although Polacanthus has been reported to occur in those strata [10]-[12], [82], [160]. However, polacanthids are present in the lower Aptian Morella Formation of northeastern Spain [40]. Blows [10] illustrated a block with ankylosaur dorsal vertebrae with the uninformative ventral portion of a pelvic shield fragment and noted it as being from Charmouth, suggesting that there were upper Albian polacanthids in England [160]. However, the specimen NMW 92.34G.2 was actually found on the beach further west at Charton Bay and may have come from either the Aptian (Lower Greensand) or Albian (Upper Greensand). Only preparation of the dorsal surface of the pelvic shield would reveal if the specimen is a polacanthid or nodosaurid. A large polacanthid (BYU R254) occurs in the Poison Strip Sandstone Member of the Cedar Mountain Formation [156]. It is not a nodosaurid close to Sauropelta as reported by Carpenter and others [97], but a polacanthid that was initially described as cf. Hoplitosaurus [161]. These rocks have been dated as lower to middle Aptian by laser ablation of detrital zircons and by U-Pb dating of early diagenetic carbonate [162]. A fragmentary large nodosaurid with massive cervical spikes that may be referred to as cf. Sauropelta (DMNS 49764) has been recovered from the overlying Ruby Ranch Member about 20 m up section in the same region [163] in strata interpreted to be of Lower Albian age [162]. Thus, the youngest polacanthids occur in the lower to possibly mid-Aptian and the oldest documented nodosaurids occur in the upper Aptian or lower Albian in both Europe and North America with no discernible stratigraphic overlap ( ). Why this faunal discontinuity occurs is unknown. There are no documen
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A New Basal Ankylosaurid (Dinosauria: Ornithischia) from the Lower Cretaceous Jiufotang Formation of Liaoning Province, China
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[
"Fenglu Han",
"Wenjie Zheng",
"Dongyu Hu",
"Xing Xu",
"Paul M. Barrett"
] |
2014-08-10T00:00:00
|
A new ankylosaurid, Chuanqilong chaoyangensis gen. et sp. nov., is described here based on a nearly complete skeleton from the Lower Cretaceous Jiufotang Formation of Baishizui Village, Lingyuan City, Liaoning Province, China. Chuanqilong chaoyangensis ...
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https://www.ncbi.nlm.nih.gov/coreutils/nwds/img/favicons/favicon.ico
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PubMed Central (PMC)
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https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4131922/
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PLoS One. 2014; 9(8): e104551.
PMCID: PMC4131922
PMID: 25118986
A New Basal Ankylosaurid (Dinosauria: Ornithischia) from the Lower Cretaceous Jiufotang Formation of Liaoning Province, China
, 1 , 2 , * , 2 , 3 , 4 , 2 and 5
Fenglu Han
1 Faculty of Earth Sciences, China University of Geosciences (Wuhan), Wuhan, China
2 Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China
Find articles by Fenglu Han
Wenjie Zheng
2 Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China
3 University of Chinese Academy of Sciences, Beijing, China
Find articles by Wenjie Zheng
Dongyu Hu
4 Paleontological Institute, Shenyang Normal University, Shenyang, China
Find articles by Dongyu Hu
Xing Xu
2 Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China
Find articles by Xing Xu
Paul M. Barrett
5 Department of Earth Sciences, Natural History Museum, London, United Kingdom
Find articles by Paul M. Barrett
Peter Dodson, Editor
1 Faculty of Earth Sciences, China University of Geosciences (Wuhan), Wuhan, China
2 Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China
3 University of Chinese Academy of Sciences, Beijing, China
4 Paleontological Institute, Shenyang Normal University, Shenyang, China
5 Department of Earth Sciences, Natural History Museum, London, United Kingdom
University of Pennsylvania, United States of America
Competing Interests: The authors have declared that no competing interests exist.
Conceived and designed the experiments: XX. Performed the experiments: FLH. Analyzed the data: FLH. Contributed reagents/materials/analysis tools: XX DYH WJZ. Contributed to the writing of the manuscript: FLH PB XX.
Copyright © 2014 Han et al
This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are properly credited.
Associated Data
Supplementary Materials
GUID: 12FE3422-7959-49A2-997E-3369508387AE
Data Availability Statement
The authors confirm that all data underlying the findings are fully available without restriction. All relevant data are within the paper and its Supporting Information files.
Abstract
A new ankylosaurid, Chuanqilong chaoyangensis gen. et sp. nov., is described here based on a nearly complete skeleton from the Lower Cretaceous Jiufotang Formation of Baishizui Village, Lingyuan City, Liaoning Province, China. Chuanqilong chaoyangensis can be diagnosed on the basis of two autapomorphies (glenoid fossa for quadrate at same level as the dentary tooth row; distally tapering ischium with constricted midshaft) and also a unique combination of character states (slender, wedge-like lacrimal; long retroarticular process; humerus with strongly expanded proximal end; ratio of humerus to femur length = 0.88). Although a phylogenetic analysis places Chuanqilong chaoyangensis as the sister taxon of the sympatric Liaoningosaurus near the base of the Ankylosauridae, the two taxa can be distinguished on the basis of many features, such as tooth morphology and ischial shape, which are not ontogeny-related. Chuanqilong chaoyangensis represents the fourth ankylosaurid species reported from the Cretaceous of Liaoning, China, suggesting a relatively high diversity in Cretaceous Liaoning.
Introduction
Ankylosauria is a group of quadrupedal herbivorous dinosaurs characterised by parasagittal rows of osteoderms on the dorsolateral surface of the body and a heavily armored skull [1]. The earliest records of the group have been reported from various Early or Middle Jurassic localities, and include Bienosaurus lufengensis, Tianchisaurus nedegoapefererima, and Sarcolestes leedsi [2]–[4], although all of these records have been considered either nomina dubia or dubiously referable to Ankylosauria [1], [5]. Definitive ankylosaur taxa are known to occur from the Late Jurassic (e.g., Gargoyleosaurus from western North America: [6]) to the end of the Cretaceous and their remains have been reported from all continents except Africa [1]. In Liaoning Province, China, three ankylosaurian species have been reported: Liaoningosaurus paradoxus from the Lower Cretaceous Yixian Formation [7] and Crichtonsaurus bohlini [8] and C. benxiensis [9] from the Upper Cretaceous Sunjiawan Formation. Liaoningosaurus was originally considered to be a possible nodosaurid [7], but a recent study suggests that it is a basal ankylosaurid [10]. C. bohlini and C. benxiensis are also referable to the Ankylosauridae (and are probably basal ankylosaurines) [9], [10].
Here, we describe a fourth ankylosaur species from Liaoning based on a specimen collected from the Lower Cretaceous Jiufotang Formation. This specimen preserves a nearly complete skeleton, and it provides new information on the morphology and taxonomy of the Ankylosauria.
Materials and Methods
The permits for this research were obtained from the Chaoyang Jizantang Paleontological Museum of Liaoning, China. All of the materials described herein were collected from a single quarry by local farmers. Locality information was provided by staff at the Chaoyang Jizantang Paleontological Museum. The fossils are two-dimensionally preserved and visible in ventral view only. The material includes a skull and articulated postcranial material referable to a single individual. The skull and mandible are nearly complete, but have been strongly compressed dorsoventrally. The vertebral column includes the cervicals, dorsals, sacrals, and most of the caudals, but most of them are disarticulated. Both of the fore- and hind limbs are well preserved and articulated. Armor is preserved around the entire body but is only visible in ventral view.
In order to compare the ratios of humerus/tibia length to femur length with those of other ankylosaurs, the data were analysed in the software package SPSS 16.0 using the linear fit function. The best fit lines, regression equation and R2 values are presented in the Results.
The phylogenetic position of Chuanqilong chaoyangensis was inferred using parsimony analysis. The new taxon was incorporated into a previously published data matrix built to examine ankylosaurian interrelationships [10]. Liaoningosaurus paradoxus was also re-scored in the matrix based on our firsthand observations of the holotype specimen (Text S1). The modified data matrix consists of 170 characters and 52 taxa. The matrix was analyzed using TNT [11], and all of the characters were treated as equally weighted and unordered. The analysis was conducted using a heuristic search with 1000 replicates. TBR branch swapping was employed and 100 parsimonious trees were saved per replicate. A reduced consensus analysis was performed to identify wildcard taxa within TNT to provide maximum phylogenetic resolution for the new taxa [11]. Standard bootstrap values (absolute frequencies) were calculated using a traditional heuristic search with 1000 replications. Bremer supports were calculated by running the script “Bremer. run” automatically.
Nomenclatural Acts
The electronic edition of this article conforms to the requirements of the amended International Code of Zoological Nomenclature (ICZN), and hence the new names contained herein are available under that Code from the electronic edition of this article. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information viewed through any standard web browser by appending the LSID to the prefix “http://zoobank.org/”. The LSID for this publication is: urn:lsid:zoobank.org:pub:9D60475B-FA91-464B-8EBF-D335582AE23E. The electronic edition of this work was published in a journal with an ISSN (1932–6203), and has been archived and is available from the following digital repositories: PubMed Central (http://www.ncbi.nlm.nih.gov/pmc), LOCKSS (http://www.lockss.org).
Institutional Abbreviations
AMNH, American Museum of Natural History, New York, USA; BXGM, Benxi Geological Museum, Liaoning Province, China; CEUM, Prehistoric Museum, College of Eastern Utah, Price, Utah, USA; CJPM, Chaoyang Jizantang Paleontological Museum; DMNH, Denver Museum of Natural History, Denver, Colorado, USA; IVPP, Institute of Vertebrate Paleontology and Paleoanthropology, Beijing, China; LPM, Liaoning Paleontological Museum, Shenyang, Liaoning Province, China; MPC, Mongolian Paleontological Center, Ulaanbaatar, Mongolia; MTM, Hungarian Natural History Museum, Budapest, Hungary; MU, University of Missouri, Columbia, Missouri, USA; NHMUK, Natural History Museum, London, UK; PIN, Paleontological Institute, Russian Academy of Sciences, Moscow, Russia; QM, Queensland Museum, Brisbane, Australia; ROM, Royal Ontario Museum, Toronto, Canada; SMU, Southern Methodist University, Dallas, Texas, USA; TMP, Royal Tyrrell Museum of Palaeontology, Drumheller, Alberta, Canada; UALVP, University of Alberta Laboratory for Vertebrate Paleontology, Edmonton, Alberta, Canada; YPM, Yale Peabody Museum, New Haven, Connecticut, USA.
Results
Systematic Paleontology
Dinosauria Owen, 1842 [12]
Ornithischia Seeley, 1887 [13]
Thyreophora Nopcsa, 1915 [14]
Ankylosauria Osborn, 1923 [15]
Ankylosauridae Brown, 1908 [16]
Chuanqilong gen. nov. urn:lsid:zoobank.org:act:76DD6D3F-F23C-4AC0-B480-20AC73B50279
Type Species
Chuanqilong chaoyangensis gen. et sp. nov. urn:lsid:zoobank.org:act:EE6564A0-33AE-4CC7-B4EA-8CD682E1EB43
Etymology
The generic name is derived from Chinese Chuanqi (legendary, referring to western Liaoning providing a spectacular assemblage of Mesozoic terrestrial fossils) + long (dragon). The specific name is derived from the broader geographical area including the type locality.
Holotype
CJPM V001, a nearly complete skeleton missing only the distal portion of the caudal series. The specimen is housed in the Chaoyang Jizantang Paleontological Museum. A cast of the holotype specimen is housed in the Institute of Vertebrate Paleontology and Paleoanthropology as IVPP FV 1978.
Locality and Horizon
Baishizui Village, Goumenzi County, Lingyuan City, Liaoning Province, China ( ); the quarry is in the Jiufotang Formation (Lower Cretaceous, Aptian) [17] ( ). A detailed stratigraphical investigation of this quarry is required to establish its relationships to other exposures of the Jiufotang Formation in the area.
Diagnosis
An ankylosaur distinguished from other ankylosaurs by two autapomorphies: the glenoid fossa for the quadrate is at the same level as the dentary tooth row; and the distally tapering ischium is constricted at midshaft length. Chuanqilong also differs from all other ankylosaurians in having the following unique combination of character states: presence of a long retroarticular process (differs from all other ankylosaurians except Gargoyleosaurus); presence of a slender, wedge-like lacrimal (differs from all other known ankylosaurians except Minmi); ratio of humerus to femur length of 0.88 (notably higher than in most known ankylosaurians except Hungarosaurus and Liaoningosaurus); the width of the proximal end of the humerus is half of the length of the humeral shaft (substantially different from that of Liaoningosaurus, in which this ratio is 0.38); presence of subtriangular unguals (absent in all other ankylosaurs except Liaoningosaurus and Dyoplosaurus).
Description and Comparisons
The holotype skeleton is exposed mainly in ventral view ( ) and as result numerous anatomical details are not visible. In addition, the presence of some elements obscures large portions of the other bones present limiting the amount of information available. Nevertheless, preservation is generally good. Although the specimen represents a large animal (approximately 4.5 m in total body length; see for measurements of the holotype skeleton), it is likely to be a juvenile individual based on several features. For example, the vertebral centra are not fused to their neural arches in all visible vertebrae including cervicals, dorsals, and caudals. In addition, the sacral ribs are not fused to the sacral centra. Consequently, an adult individual is likely to have been greater than 4.5 m in length. Jurassic ankylosaurians are mostly relatively small in size with body lengths no greater than 4 meters. For example, Mymoorapelta and Gargoyleosaurus each have lengths of approximately 3 m [18], [19]. By comparison, most Cretaceous ankylosaurians have body lengths greater than 5 m. For example, the primitive ankylosaurid Cedarpelta has an estimated body length of 7.5–8.5 m [20].
Table 1
BoneL/RLengthWpWdWm1* 2* ScapulaL——19.59——R40—199——HumerusL3518145.5——R35———20—UlnaL18135———R17136———RadiusL236.56.54——R23884——FemurL4018179——R4020189——TibiaL—15197——R3612176——FibulaL32653.5——R35653.5——IliumL76———2015R————2015IschiumL421875——R411975.5——Metacarpal IL7.54.54.5———Metacarpal IIL844———Metacarpal IIIL845———Metacarpal IVL724———Metatarsal IIL137.553.5——Metatarsal IIIL15.57.563.5——Metatarsal IIR136.553.5——Metatarsal IIIR15663——Metatarsal IVR13561.5——
Skull
The skull and mandibles are strongly compressed dorsoventrally. The skull is triangular in ventral view, with a transverse width that was probably greater than its length, as in ankylosaurids ( , ) [21]. The maxilla is partially exposed in lateral view and exhibits a shallow, flattened buccal emargination. A large, triangular antorbital fossa or fenestra seems to be present, located in the caudodorsal region of the maxilla ( ) in lateral view. An antorbital fenestra is also present in juvenile ankylosaurids such as Liaoningosaurus and Pinacosaurus, though it is replaced by a small concavity in adult Pinacosaurus [7], [22]. A small antorbital fenestra is also present in the probably adult Minotaurasaurus [22],[23], whereas it is either unknown or absent in all other ankylosaurians [1]. It seems likely that the presence of an open antorbital fenestra in Chuanqilong chaoyangensis is due to its juvenile status. However, the orbit is circular in outline, and relatively small in comparison to skull length, contrary to what would be expected in a juvenile individual. The lacrimal is slender and wedge-shaped ( ), forming the rostral margin of the orbit, as in the basal ankylosaurid Minmi, whereas it is sub-rectangular in other known ankylosaurians, such as Pinacosaurus and Cedarpelta [1]. A long and dorsoventrally compressed supraorbital is visible in lateral view, contacting the lacrimal rostroventrally. Caudally, the postorbital is damaged and only partially exposed. The ‘squamosal’ may be composed of both the squamosal and a portion of the postorbital. It is subrectangular in outline and ornamented with subparallel grooves, as in Minmi [24] ( ). The left quadrate is exposed in rostral view. It is long and straight with a rectangular head, and there is no indication that the quadrate head was fused with the squamosal, thereby differing from the condition seen in nodosaurids [25]. Below the quadrate head, the shaft is transversely expanded to form a wide, shallow, and rostrally-opening depression. The quadrate constricts ventral to this point and is narrowest at midshaft length. A crescentic depression is present on the cranioventral surface of the quadrate for reception of the quadratojugal. The pterygoid process is thin, short, and sub-triangular in outline. The transversely expanded ventral end is composed of two well-defined mandibular condyles, which are separated by a shallow groove ventrally. The medial condyle is transversely wider and extends further ventrally than the lateral condyle, as in most other ankylosaurians. No other features of the skull are visible.
Mandible
The paired mandibular rami are preserved separately and both are visible in medial view only ( , ). The predentary is missing. The mandible is long and shallow, as in other basal ankylosaurids, but it differs from other taxa in the apparent absence of an osteoderm from its ventral margin. The lack of an osteoderm in this area may be either due to incomplete preservation, suggesting that they were not fused to the mandibular bones and thereby supporting the suggestion that the holotype Chuanqilong was not fully grown. In adult individuals of other ankylosaurians, such as Pinacosaurus and Saichania, osteoderms are fused to the lateral surface of the mandible [22], [26]. Alternatively, an osteoderm, if present, might be restricted to the lateral-most corner of the mandible and hence obscured from view (this condition is present in juvenile individuals of Pinacosaurus grangeri e.g., IVPP V16853; V. Arbour, pers. comm.), which would also represent a juvenile feature. The dentary tooth row is straight or slightly arched dorsally, but it is not as strongly sinusoidal as those of derived ankylosaurians, such as Euoplocephalus [27] or Pinacosaurus [26]. In dorsal view, the rostral end of the dentary tooth row is curved medially. At least 20 alveoli are present. The ventral margin of the mandible is relatively straight in its rostral and middle regions, but curves caudodorsally in its caudal part. The right dentary symphysis is preserved and is slightly downturned, short, robust, and sub-triangular in cross-section. The Meckelian canal is open, long, and deep. The coronoid eminence is prominent and projects above the level of the dentary tooth row, as in nodosaurids, whereas the coronoid eminence is situated at approximately the same level as the dentary tooth row in ankylosaurids, including Pinacosaurus [26], Euoplocephalus [27], and Ankylosaurus [28]. The splenial and prearticular are missing, exposing the adductor fossa, which is large and located below the coronoid eminence. The articular is small and oval in outline in lateral view. The retroarticular process is long and slender, as in Gargoyleosaurus, but differs from those of all other ankylosaurians, which possess relatively short and deep retroarticular processes [19]. Unusually, the glenoid fossa is situated in a relatively dorsal position, lying at approximately the same level as the dentary tooth row, unlike the condition present in all other ankylosaurians known from appropriate material, in which the glenoid fossa is situated at a level ventral to the dentary tooth row (e.g., Pinacosaurus: [26]). Unfortunately, this feature cannot be assessed in Liaoningosaurus [7].
Dentition
Premaxillary teeth are unknown due to breakage of the snout. Both maxillary and dentary teeth are preserved, with the former exposed labially and the latter exposed lingually.
There are at least 20 alveoli in the left maxilla. The maxillary tooth counts of most ankylosaurians are around 20: Ankylosaurus has the largest number of maxillary teeth (34–35) [28], whereas Liaoningosaurus has the smallest number (about 10: [7]). Tooth count probably increases during growth [7] and adult tooth count also varies between species [28]. The teeth and their marginal denticles are small in comparison to the size of skull, as occurs typically in ankylosaurids [21].
The four preserved rostral maxillary teeth are smaller than the caudal teeth. The crowns are as tall as their width with sub-triangular outlines, as in most ankylosaurians ( ). The base of the crown is strongly swollen with a weak cingulum, as in ankylosaurids [21]. There is no shallow notch at the base of tooth crowns, unlike the notched condition present in the ankylosaurid Crichtonsaurus [8] and the nodosaurid Edmontonia [29]. Additionally, the teeth of C. bohlini bear much larger denticles and a more distinct cingulum than those of Chuanqilong. There is no distinct primary ridge, and secondary vertical ridges and grooves are present on the labial surfaces of the tooth crowns. These ridges usually terminate apical to the cingulum, but some ridges extend across the cingulum to the basal margin of the crown. The crescentic cingula seen in some ankylosaurs, such as Texasetes, are absent. Small denticles and cusps are present on one rostral maxillary tooth crown. The denticles are small and tapering with a round cross-section at their base. However, most of the teeth do not bear these structures, though it is not clear if this absence is due to poor preservation or tooth wear. Most of the dentary teeth are missing. Those that are preserved are only partially exposed in the left dentary. Dentary teeth seem to have been similar in size and shape to the maxillary teeth.
Axial Skeleton
Several cervical and dorsal vertebrae are scattered on the slab. The cervical centra are spool-like and shorter than they are wide: their articular surfaces are all obscured. The dorsal centra are also spool-like in ventral view, slightly amphicoelous, and possess concave lateral surfaces. Dorsal centra are longer than tall. A ventral keel is absent from all of the preserved dorsal centra. One sacral vertebra is exposed on the slab in ventral view. Its centrum is wider than it is long, and its exposed articular surface is rugose, suggesting that it had not yet fused with the other sacrals. Several sacral ribs are preserved separately. They are robust and dumbbell-shaped in outline. Approximately 20 caudal vertebrae are preserved, but most of them are disarticulated. The centra of the proximal and middle caudal vertebrae are shorter than they are wide. Deep longitudinal grooves are present on the ventral surfaces of the proximal caudal vertebrae. Chevron facets are well developed, with the caudal facets more prominent than the cranial ones.
One middle caudal vertebra is well preserved. Its centrum is relatively longer and transversely narrower than those of the cranial caudals and in lateral view it has a square outline. The transverse process is reduced to a small nodular process and is located on the dorsal part of the lateral surface of the centrum. The neural spines are elongate, oriented caudodorsally, and possess arc-shaped outlines. The prezygapophyseal facets are oval in outline and face craniomedially, whereas the postzygapophyses are positioned near the tip of the neural spine and face caudolaterally.
Caudually, three additional distal caudal vertebrae are tightly articulated. Their centra are more elongate and transversely narrower than those of the proximal and middle caudals. In these vertebrae, the neural spine has merged with the postzygapophyses to form a single midline caudal process that extends caudally. The caudal process terminates cranial to the midpoint of following vertebra, as in nodosaurids, but unlike the condition in derived ankylosaurids, in which the process is longer [21]. The prezygapophyses are short and reduced in size, corresponding to the size reduction of the postzygapophyses. No chevrons are preserved.
Several ossified hypaxial tendons are present near the distal region of the tail ( ). During preservation, they have moved from their original positions and arrangement so that they are now aligned in different orientations. On the basis of the morphology of the preserved caudals, which does not conform with that of the tail club handle morphology seen in ankylosaurine taxa, a tail club knob was probably absent, as in all nodosaurids and some basal ankylosaurids (e.g., Minmi: [30]). Presence of a tail club was formerly an important diagnostic feature of ankylosaurids [21], but recent work has indicated that tail clubs may have been present in ankylosaurines only, a clade that includes Ankylosaurus [28], Euoplocephalus [31], Pinacosaurus [32], Talarurus [33], Saichania [34], Tarchia [35], Tianzhenosaurus [36], Dyoplosaurus [37], and Nodocephalosaurus [38]. It is unknown whether tail clubs were present or not in the basal ankylosaurids Crichtonsaurus [8], [9], Cedarpelta [20], Gobisaurus [39], and Shamosaurus, but the ankylosaurids Minmi, Liaoningosaurus, and Zhongyuansaurus lack tail clubs [7], [40], [41]. The likely absence of a tail club in Chuanqilong chaoyangensis adds further support to the hypotheses that the tail club is a derived feature that appears only in derived, and currently only Late Cretaceous taxa.
Pectoral Girdle and Forelimb
Scapula
Both scapulae are preserved. The scapula and coracoid are not co-ossified, contrary to the condition in most ankylosaurians (e.g., Ankylosaurus: [28]), but this may be another indication of specimen immaturity [42]. The scapula and coracoid are unfused in all known juvenile ankylosaurs, including Liaoningosaurus [7], an indeterminate nodosaurid hatchling from the Paw Paw Formation of Texas, juvenile Pinacosaurus [43], and Anoplosaurus [42], but they are fused in most adult or sub-adult individuals ( ). The scapula blade is slender, deflected caudoventrally, and has a rhomboidal outline ( , ). The dorsal margin of the scapula blade is relatively straight, whereas its ventral margin is concave. The caudal margin expands dorsoventrally. The shaft of the scapula blade is narrowest caudal to the glenoid fossa. A transverse flange is positioned along the craniodorsal margin of the scapula, as in ankylosaurids, whereas in nodosaurids the acromion is positioned ventrally, near the glenoid, and overhangs the coracoid [21]. There is no distinct enthesis present on the ventral edge of the scapula, which probably marks the insertion of the M. triceps longus caudalis (See [28]), and has been reported in derived ankylosaurids, including Crichtonsaurus [9], Ankylosaurus [28], and Euoplocephalus [31]. The absence of a distinct enthesis also suggests that CJPM 001 is not fully grown (K. Carpenter, pers. comm.). The glenoid fossa is large, oval in outline, and faces ventromedially. Both of the coracoids are missing or concealed.
Table 2
TaxonSpecimen numberFemur lengthTibia lengthHumerus lengthScapula & coracoidRatio H/FRatio T/FReference Liaoningosaurus paradoxus IVPP V125602.52.52.5unfused11.00 [7] nodosaurid scuteling from Paw Paw FormationSMU 724447.3–6.84unfused0.94– [52] Anodontosaurus lambei AMNH 526625.5190–unknown–0.75 [31] Crichtonsaurus benxiensis BXGMV0012-1322923fused0.720.91 [9] Crichtonsaurus bohlini LPM 10134.3–24unfused0.700.00 [8] cf. Pinacosaurus MPC 100/13053820.326.5–0.700.53 [66] Chuanqilong chaoyangensis CJPM 001403635unfused0.880.90this study Pinacosaurus granger PIN 614402730unknown0.750.68 [66] Animantarx ramaljonesi CEUM 6288R41.5–29.8fused0.72– [20] Gargoyleosaurus parkpinoorum DMNH 2772646.5–29.2unknown0.63– [19] Talarurus plicatospineus PIN 557-34724.833.5fused0.710.53 [33] Hungarosaurus tormai MTM 2007.2549–45.5fused0.93– [55] Euoplocephalus tucki UALVP 3151.5–37.7probablyunfused0.73– [31] Euoplocephalus tucki AMNH 540453.542.140.3fused0.750.79 [31] Polacanthus foxii NHMUK R1755334.5–unknown–0.65 [53] Dyoplosaurus acutosquameus ROM 78456.2––unknown–– [37] Scolosaurus cutleri NHMUK nR51616041.544fused0.730.69 [14] Ankylosaurus magniventris AMNH 521467–54.2fused0.81– [28] Sauropelta edwardsi AMNH 303670–49.5fused0.71– [67]
Humerus
Both humeri are well preserved, except that the ventral part of the deltopectoral crest is damaged on the left humerus. The left humerus is exposed in cranial view, and the right humerus in lateral view ( , ). The humerus is short and robust. The deltopectoral crest is large and rounded in outline in cranial view, unlike in Crichtonsaurus benxiensis which possesses a straight lateral margin [9] ( ). The deltopectoral crest and the transverse axis through the distal condyles are in the same plane, and the deltopectoral crest extends for more than half of the length of the humerus as in ankylosaurids, but unlike the condition in nodosaurids, which possess a relatively short deltopectoral crest [21]. There is no distinct separation between the humeral head and the deltopectoral crest as in most ankylosaurians, but in contrast to Cedarpelta and Ankylosaurus in which the dorsal surface of the deltopectoral crest is lower than the humeral head [28], [44] ( ). The width of the proximal end is much greater than the distal width as in most ankylosaurids except Zhongyuansaurus, in which both ends are of equal width [40]. The laterally placed radial condyle is oval and more prominent than the medial ulna condyle. The lateral epicondylar ridge is not well developed.
Ulna and Radius
Both of the ulnae and radii are complete ( , ). The olecranon process of the ulna is low and wedge-shaped as in Liaoningosaurus [7] and juvenile specimens of Pinacosaurus [45], whereas it is tall and strongly developed in most large ankylosaurians, such as Pelorolites and Cedapelta [44] ( ). The low olecranon process may represent an ontogenetically variable character of ankylosaurians [7]. The humeral notch is moderately developed. The radius is slender in comparison with the ulna. It is rod-like with a flat proximal articular surface and a rugose, convex distal end. The distal end is wider transversely than the proximal end.
Manus
The left manus contains four complete but disarticulated metacarpals and their identifications are based on the well preserved metacarpals of Peloroplites [44] and Pinacosaurus [45], [46] ( ). All of the preserved metacarpals are slender, as in Pinacosaurus [45]. Metacarpal III is the longest. Metacarpals I and II are sub-equal in length. Metacarpal IV is significantly shorter than other metacarpals. Metacarpal I is the most robust of the metacarpals, as in other ankylosaurids [1]. Metacarpals II and IV are more slender than metacarpals I and III. All of the metacarpals have expanded proximal and distal ends. The proximal articular surfaces are slightly concave, whereas the distal articular surfaces are strongly convex. There are no distinct ginglymi at the distal end. The phalanges are proximodistally short and transversely wide. The ungual phalanges are triangular in outline with sharp point in dorsal view. Their ventral surfaces are flattened and their proximal surfaces have a round outline and are slightly concave.
Pelvic girdle and hind limb
Ilium
Both ilia are well preserved and exposed in ventral view ( , ). As in other ankylosaurs, the preacetabular process rotated medially, making the ‘original’ lateral surface face dorsally, whereas the postacetabular process rotated in apposition and the original surface faces ventrally [47]. The preacetabular process is very long and transversely wide, and diverges laterally from the vertebral column at an angle of approximately 45°. The lateral margin of the preacetabular process is straight in ventral view, as in ankylosaurids, but unlike the condition in most nodosaurids, such as Sauropelta [25], Struthiosaurus [48], and Zhejiangosaurus [49], in which it is laterally curved. The postacetabular process is subtriangular in outline. It is very short, with a length less than that of the acetabulum, as in ankylosaurids [1]. The acetabulum is imperforate with a concave articular surface for accepting the femoral head, as in all ankylosaurians except Mymoorapelta [18]. The pubic peduncle is well developed with a sub-rounded profile and is dorsoventrally compressed. The ischial peduncle is rudimentary.
Ischium
The ischium is long, slender, and mediolaterally compressed ( ). The proximal end is expanded craniocaudally and contributes to half of the medial wall of the shallow acetabulum. There is no obturator process, which is absent in all ankylosaurians. The shaft of the ischium is slender and slightly curved ventrally, as in the ankylosaurid Zhongyuansaurus [40], whereas it straight in most ankylosaurids [28] and significantly curved ventrally near the distal end in nodosaurids [1]. The shaft of the ischium is unique in being narrower in its mid-shaft region and widening towards to the distal end, prior to tapering again further distally, whereas in other ankylosaurians the shaft either tapers distally along the whole shaft (e.g. Ankylosaurus [28], Sauropelta, Edmontonia [48]) or remains sub-equal in size along the whole shaft (e. g., Euoplocephalus: [31]), or is just slightly expanded at the distal end (e.g., Cedarpelta: [44]) ( ). The proximal end of the ischium is straight in lateral view. This is unlike the convex and fan-like ischium in the ankylosaurids Ankylosaurus [50] and Euoplocephalus [51], and also unlike the concave proximal ischia of the nodosaurids Struthiosaurus and Edmontonia [48]. The proximal end of the ischium lacks the medial wall present in the basal ankylosaurid Cedarpelta [20], [44].
Femur
The femur is robust and straight, as in other ankylosaurians ( , ). The femoral head is robust and expanded forming a spherical articular surface. It forms an angle of about 145° with the long axis of the femur. Both the cranial and greater trochanters are present, and they are separated from the femoral head by a prominent constriction. The cranial trochanter is slender, finger-like, and separated from the greater trochanter by a vertical cleft. The cranial trochanter is present in juveniles, such as the Paw Paw nodosaurid scuteling and Anoplosaurus, but fused with the greater trochanter in most adult ankylosaurs [52]. However, the cranial trochanter is also present in some large primitive nodosaurids, such as Polacanthus [53] and Texasetes [54]. So the presence of a cranial trochanter is likely to be a primitive character of ankylosaurids, as well as being under ontogenetic control in some taxa. The fourth trochanter is a prominent rugosity that is located distal to femoral mid-length, as in typical ankylosaurids [21].
Distally, a shallow cranial intercondylar fossa is present. A deep caudal intercondylar groove divides the medial (tibia) and lateral (fibula) condyles, and the former is slightly larger than the latter. The ratio of humerus to femur length is 0.83, similar to the condition in Ankylosaurus, but lower than the ratios in Liaoningosaurus, the indeterminate juvenile nodosaurid from Paw Paw Formation, and Hungarosaurus, and greater than those of other known ankylosaurians ( ; ). Juveniles may have proportionally longer forelimbs than adults [52]. The juvenile Liaoningosaurus and indeterminate Paw Paw nodosaurid have humerus to femur length ratios of 1.0 and 0.93, respectively, whereas the ratio is about 0.7 in most large ankylosaurians ( ). However, this ratio is also substantially higher in adult Hungarosaurus (0.92: [55]), Ankylosaurus (AMNH 5214, 0.81, [28]), and in the late juvenile Chuanqilong (0.88) ( ). This suggests that the ratio of humerus to femur length may represent a valid taxonomic difference.
Tibia
The tibia is shorter than the femur ( , ). The ratio of tibia to femur length is approximately 0.9. This is similar to the ratio in Crichtonsaurus benxiensis (0.91: [9]), Europelta carbonensis (0.91: [56]), Liaoningosaurus paradoxus (0.95: pers. observ.) and greater than in all other known ankylosaurians ( ; ). The tibia is straight, robust, and greatly expanded mediolaterally both proximally and distally. The transverse expansion of the proximal end is relatively weaker than that of the distal end in cranial view.
Fibula
The fibula is slender and slightly shorter than the tibia. The proximal end is expanded craniocaudally and compressed laterally. The whole shaft is relatively equal in size and oval in cross-section. The distal end is slightly expanded mediolaterally with a flattened cranial surface.
Proximal tarsals
The calcaneum and astragalus are not preserved, and they are inferred to have remained unfused to the distal end of the tibia. The calcaneum and astragalus are usually fused with the distal end of the tibia in most ankylosaurians [1], but they are unfused in juveniles of Anodontosaurus lambei (AMNH 5266; [31]) Liaoningosaurus [7], and Pinacosaurus [46], suggesting that this was under ontogenetic control in ankylosaurians. However, the astragalus is not fused to the distal end of the tibia in the early ankylosaurians Mymoorapelta (DMNH 15162: [57]), Peloroplites (CEUM 11319: [44]), and possibly Hylaeosaurus (NHMUK R2615: [53]), which indicates they may have been unfused primitively in adult ankylosaurians.
Pes
Metatarsals II, III, and IV are well preserved and in articulation in the right foot ( , ). The possible presence of metatarsals I and V cannot not be excluded due to the preservation of the specimen. Metatarsal III is much longer (187.5% of the length) and more robust than metacarpal III. This ratio is similar to that seen in the primitive ankylosaurid Gargoyleosaurus (184.4%: [19]), greater than in most ankylosaurians, such as Pinacosaurus grangeri (113.5%: [45]) and Talarurus plicatospineus (132.8%: [45]), but smaller than that in Liaoningosaurus, which has even longer metatarsals (more than 200%: [7]). Metatarsals II and IV are sub-equal in length, and metatarsal III is longer and more robust than the other two metatarsals. They all have expanded proximal and distal ends. The proximal end of metatarsal II is dorsoventrally deeper than it is wide transversely and it has a concave and oval articular surface for the distal tarsals. Metatarsal III has a square cross-section proximally, and metatarsal IV is transversely wider than deep craniocaudally. The distal ends of all metatarsals are transversely expanded and bear no, or very weak, ginglymi.
The unguals are robust and sub-triangular in outline with sub-rounded distal ends in dorsal view. This is unlike the pedal unguals of Liaoningosaurus, which have much sharper distal ends [7]. Coombs [51] noted that in ankylosaurs the pedal unguals are widest at a point approximately one-third of the distance from their proximal ends in juveniles, whereas in adults they are widest proximally. However, in juveniles like Liaoningosaurus and Chuanqilong, the pedal unguals are widest proximally and taper distally, contrary to this observation. Ungual shape may, therefore, be a useful character for taxonomic or systematic purposes, and similar sub-triangular unguals are also known in Dyoplosaurus [37].
Armor
Cranial armor is not visible on the slab. The cervical armor usually comprises two cervical half rings in ankylosaurids and three cervical half rings in nodosaurids. These half rings consist of a superficial layer of osteoderms fused to an underlying band of bone. The osteoderms are usually pitted and rugose, similar to body osteoderms, whereas the connecting band is usually smooth and plate-like [58]. In Chuanqilong, only one cervical half ring is present in ventral view. The band appears to be fused into a single large plate, as in ankylosaurids [58]. However, it is compressed dorsoventrally, damaged and separated into four sections ( ). The right three sections are thickened, arched dorsally, and subrectangular in outline with smooth ventral surfaces. The left section has a wide medial edge and tapers caudolaterally with a subtriangular profile, and this section has a straight craniolateral edge as also seen in Gargoyleosaurus [19], but which is unknown in other ankylosaurians. Two large armor plates are preserved in the shoulder region. They are thickened, subrectangular in outline, have flat, smooth surfaces, and are thicker along their margins than centrally. The larger plate is twice the length of the smaller one. Both of them are similar to the osteoderms of the cervical half ring, and they may represent separate cervical armor plates. A relatively small triangular armor plate is present between the proximal end of the left ulna and radius ( ). It is dorsoventrally compressed, wide at the base and tapers distally. One nearly complete oval armor plate is present near the left ischium in dorsal view, which is sharply keeled along its midline ( ). A variety of small irregular osteoderms and ossicles are preserved over the whole body in ventral view ( ), as in most ankylosaurians. Dermal armor is absent in the indeterminate Paw Paw nodosaurid ([52]: SMU 72444), the juvenile specimen of Anoplosaurus [42], and the hatchling dinosaur Propanolosaurus [52], [59]. However, dermal armor is present in the small specimen Liaoningosaurus, suggesting that the ability to produce dermal armor had already appeared by this early growth stage [7], although armor is absent in the even smaller specimen of Propanolosaurus [59].
Discussion
Ankylosauria is traditionally divided into two families, Ankylosauridae and Nodosauridae, which are distinct in many features [21]. A third group, Polacanthinae [60] or Polacanthidae [5], has been proposed, and it is normally defined as all ankylosaurians more closely related to Gastonia than to either Edmontonia or Euoplocephalus [5]. However, some phylogenetic analyses, including ours (see below) do not recover this group as a separate clade [1], [10], [26], and here we follow traditional ankylosaurian taxonomy in our discussion.
Chuanqilong chaoyangensis possesses many ankylosaurid features, including cheek teeth with a strongly swollen tooth crown with a weak cingulum, a long deltopectoral crest that extends for more than half of humeral length, a straight lateral margin of the preacetabular process, a very short postacetabular process that is shorter than the length of the acetabulum, a slender ischial shaft that is curved slightly ventrally, and a distally located fourth trochanter. However, it lacks several features shared by derived ankylosaurids, such as the presence of a tail club. This character combination suggests that Chuanqilong chaoyangensis represents a basal ankylosaurid.
In order to confirm our hypothesis regarding the systematic position of Chuanqilong chaoyangensis, we conducted a phylogenetic analysis by adding Chuanqilong chaoyangensis to a recently published dataset on ankylosaurian phylogenetic relationships [10]. Our analysis produced 15902 most parsimonious trees (MPTs), with tree lengths of 542 steps (Consistency Index = 0.34, Retention Index = 0.66). The strict consensus tree (not shown) of these 15902 MPTs lacks resolution, with the only clear result being recovery of ankylosaurid monophyly. A reduced consensus tree was calculated a posteriori which excluded seven wildcard taxa (Zhejiangosaurus, Niobrarasaurus, Hungarosaurus, Antarctopelta, Anoplosaurus, Polacanthus rudgwickensis, and Stegopelta) [61], and this shows considerably greater resolution ( ).
Our results indicate that Chuanqilong chaoyangensis is a basal ankylosaurid and that it is the sister taxon of the sympatric Liaoningosaurus. However, only two unambiguous synapomorphies support this relationship: presence of an antorbital fossa or fenestra ([10]: character 1) and scapula glenoid oriented ventrally ([10]: character 121).
It should be noted that both Chuanqilong chaoyangensis and Liaoningosaurus paradoxus are represented by specimens at a relatively early ontogenetic stage, although the much larger size, relatively smaller orbit, and higher tooth count of Chuanqilong suggest that the latter is at a more advanced ontogenetic stage. Although Chuanqilong and Liaoningosaurus are sister taxa, they can be distinguished on the basis of the following characters, which are probably not ontogenetically variable:
Cheek tooth crown morphology. In Chuanqilong, the cheek teeth are relatively small compared to the skull, and there are more than 20 maxillary teeth, whereas in Liaoningosaurus, the cheek teeth are significantly larger in comparison to the skull, and there are only approximately 10 maxillary teeth. Although Xu et al. [7] noted that the low tooth number of Liaoningosaurus may be due to its juvenile status, the ontogenetic variation in teeth number among all known ankylosaurians is less than 10. For example, the variation in Euoplocephalus and Pinacosaurus cheek teeth number is five and three, respectively [28]. Additionally, the cheek tooth crowns of Chuanqilong bear small denticles and cusps, with approximately 12 denticles per tooth, whereas in Liaoningosaurus, the denticles are large cusps that are also relatively large with respect to the size of the tooth crown, and there are approximately seven denticles per tooth. The tooth crowns of Liaoningosaurus are similar to those of another ankylosaurid from Liaoning Province, Crichtonsaurus bohlini, but differ from those of Chuanqilong.
The proximal end of the humerus is strongly expanded in comparison to humeral length in Chuanqilong (the ratio of proximal width to whole length is 0.51), whereas it is only moderately expanded in Liaoningosaurus (the ratio of proximal width to whole length is 0.38). Additionally, the distal end of the deltopectoral crest extends for more than half of the length of the humerus in Chuanqilong, as in typical ankylosaurids, whereas in Liaoningosaurus, the deltopectoral crest is less developed and does not extend to the mid-length of the humerus, as in other nodosaurids.
The lateral edge of the ilium is straight or slightly convex in ventral view in Chuanqilong, whereas it is slightly concave above the acetabulum in Liaoningosaurus.
The ischial shaft of Chuanqilong has a constriction at mid-length and tapers distally, whereas the ischial shaft is relatively equal in width along the whole length and slightly expanded at the distal end in Liaoningosaurus.
The ratios of metatarsus to metacarpus length in Chuanqilong is substantially less than that of Liaoningosaurus ( ). The metatarsus is more than twice the length of the metacarpus in Liaoningosaurus and this is probably an autapomorphy of this taxon [7]. It is unknown whether the ratio of metatarsus to metacarpus length changes during ontogeny. However, the metatarsus is less than twice the length of the metacarpus in the hatchling Propanoplosaurus ([59]: ).
The pedal unguals of Chuanqilong are widest at a point approximately one-third of the distance from the proximal end and are slightly constricted at the proximal end, whereas the pedal unguals are sub-triangular and widest at the proximal end in Liaoningosaurus [7] and Dyoplosaurus [37].
Several juvenile ankylosaurians have been recognized and provide important ontogenetic information [7], [26], [42], [51], [52], [62]. These studies indicate that some features used for species diagnosis are probably under ontogenetic control, such as some fusion characters, including fusion of the scapula and coracoid, fusion of the calcaneum and astragalus, and fusion of the cranial and greater trochanters.
Ontogenetic variation may affect phylogenetic reconstruction (e.g. [63]). Many derived features found in adult specimens are rudimentary or undeveloped in juvenile specimens, making the latter appear more basal than adult individuals in phylogenetic analyses. Therefore, ideally, ontogenetically variable characters should be excluded from phylogenetic analysis or such analyses should be based upon adult specimens only. However, many ankylosaurians are only partially preserved and it has been difficult to document their ontogenetic variation. Euoplocephalus and Pinacosaurus, which are known from multiple individuals, may provide more insights into this problem, but the taxonomy of Euoplocephalus has been controversial and many formerly referred specimens are now thought to represent other distinct taxa [31]. In order to retain as many taxa in our analysis as possible, we were unable to exclude ontogenetic characters from our phylogenetic analysis. Further ontogenetic precision could be gained from aging individuals using bone histology, which has not been widely applied to ankylosaurians. As Liaoningosaurus and Chuanqilong are represented by juvenile specimens only, more material, especially adult specimens, will help to further elucidate their phylogenetic relationships.
Chuanqilong was moderate in size compared with other known ankylosaurians ( ). However, it still larger than adult Jurassic ankylosaurians, including Mymoorapelta and Gargoyleosaurus [18], [19]. The juvenile Chuanqilong is similar in size to most Cretaceous ankylosaurians, including adult Hungarosaurus [55] and Europelta [56], but is smaller than Cedarpelta (7.5–8.5 m: [20]) and Polacanthus (5–7 m: [53]). However, as the holotype of Chuangqilong is not fully-grown, based on the above-mentioned features, this suggests that the adults of this taxon may have been among the largest ankylosaurians. This suggests in turn that ankylosaurs has already evolved large size by the late Early Cretaceous. Bone histology should be used in future to gain a more accurate understanding of the ontogenetic age of this specimen.
Supporting Information
Text S1
Updated character scores for Chuanqilong , and additional scores for Liaoningosaurus .
(DOC)
Acknowledgments
We thank Haijun Li for his invitation to work on the material and for his hospitality in Chaoyang, Liaoning province. We thank Hailong Zang for providing the photographs of this specimen. We thank Xulong Lai and Richard Butler for their useful comments. Many thanks to editor Peter Dodson, and reviewers Victoria Arbour and Kenneth Carpenter for their helpful and constructive reviews of an earlier version of this article.
Funding Statement
This project was supported by the National Natural Science Foundation of China (41120124002; 41172026) and 973 program (2012CB821900). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
Data Availability
The authors confirm that all data underlying the findings are fully available without restriction. All relevant data are within the paper and its Supporting Information files.
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https://static.famousfix.com/img/ff/favicon.ico
|
FamousFix.com
|
https://www.famousfix.com/list/nodosaurids
|
1.
Panoplosaurus
Extinct genus of dinosaurs
Overview: Panoplosaurus is a genus of armoured dinosaur from the Late Cretaceous of Alberta, Canada. Few specimens of the genus are known, all from the middle Campanian of the Dinosaur Park Formation, roughly 76 ...
0 0
2.
Acanthopholis
Genus of reptiles (fossil)
Overview: Acanthopholis (meaning "spiny scales") is a genus of ankylosaurian dinosaur in the family Nodosauridae that lived during the Late Cretaceous Period of England. A single species, A. horrida, exists.
0 0
3.
Borealopelta
Extinct genus of reptiles
Overview: Borealopelta (meaning "Northern shield") is a genus of nodosaurid ankylosaur from the Lower Cretaceous of Alberta, Canada. It contains a single species, B. markmitchelli, named in 2017 by Caleb Brown and ...
0 0
4.
Propanoplosaurus
Extinct genus of dinosaurs
Overview: Propanoplosaurus is a genus of herbivorous nodosaurid dinosaur from the Early Cretaceous Patuxent Formation of Maryland, USA. Its type specimen is a natural cast and partial natural mold of a hatchling ...
0 0
5.
Silvisaurus
Extinct species of reptile
Overview: Silvisaurus, from the Latin silva "woodland" and Greek sauros "lizard", is a nodosaurid ankylosaur from the middle Cretaceous period.
0 0
6.
Pawpawsaurus
Extinct species of reptile
Overview: Pawpawsaurus, meaning "Pawpaw Lizard", is a nodosaurid ankylosaur from the Cretaceous (late Albian) of Tarrant County, Texas, discovered in May 1992. The only species yet assigned to this taxon, Pawpa ...
0 0
7.
Niobrarasaurus
Extinct genus of reptiles
Overview: Niobrarasaurus (meaning "Niobrara lizard") is an extinct genus of nodosaurid ankylosaur which lived during the Cretaceous 87 to 82 million years ago. Its fossils were found in the Smoky Hill Chalk Member ...
0 0
8.
Hylaeosaurus
Ankylosaurian dinosaur genus from Early Cretaceous Period
Overview: Hylaeosaurus ( hy-LEE-o-SOR-əs; Greek: hylaios/ὑλαῖος "belonging to the forest" and sauros/σαυρος "lizard") is a herbivorous ankylosaurian dinosaur that lived about 136 million years ago, in the late ...
0 0
9.
Vectipelta
Genus of ankylosaurian dinosaurs
Overview: Vectipelta (meaning "Isle of Wight shield") is an extinct genus of ankylosaurian dinosaur recovered from the Early Cretaceous Wessex Formation of England. The genus contains a single species, V. barretti ...
0 0
10.
Horshamosaurus
Extinct genus of dinosaurs
Overview: Horshamosaurus is a genus of herbivorous ankylosaurian dinosaur from the Early Cretaceous of England. It lived during the Barremian of the Cretaceous and the type species is Horshamosaurus rudgwickensis ...
0 0
11.
Sauroplites
Extinct genus of dinosaurs
Overview: Sauroplites (meaning "saurian hoplite") is a genus of herbivorous ankylosaurian dinosaur from the Early Cretaceous of China.
0 0
12.
Tatankacephalus
Extinct genus of dinosaurs
Overview: Tatankacephalus is a basal genus of nodosaurid dinosaur that lived during the Early Cretaceous. It's length has been estimated at 7 meters (23 ft).
0 0
13.
Peloroplites
Extinct genus of dinosaurs
Overview: Peloroplites (from Greek pelor "monster", and hoplites, "armoured soldier") is a genus of nodosaurid armored dinosaur from Lower Cretaceous rocks of Utah, United States. It is known from a partial skull ...
0 0
14.
Hierosaurus
Extinct genus of reptiles
Overview: Hierosaurus (meaning "sacred lizard") is an extinct genus of nodosaurid ankylosaur which lived during the Cretaceous 87 to 82 million years ago. Its fossils were found in the Smoky Hill Chalk Member of ...
0 0
15.
Stegopelta
Extinct genus of dinosaurs
Overview: Stegopelta (meaning "roofed shield") is a genus of armored dinosaur. It is based on a partial skeleton from the latest Albian-earliest Cenomanian-age Lower and Upper Cretaceous Belle Fourche Member of ...
0 0
16.
Hungarosaurus
Extinct species of reptile
Overview: Hungarosaurus tormai is a herbivorous nodosaurid ankylosaur from the Upper Cretaceous (Santonian) Csehbánya Formation of the Bakony Mountains of western Hungary. It is the most completely known ankylosaur ...
0 0
17.
Gargoyleosaurus
Extinct species of reptile
Overview: Gargoyleosaurus (meaning "gargoyle lizard") is one of the earliest ankylosaurs known from reasonably complete fossil remains. Its skull measures 29 centimetres (11 in) in length, and its total body ...
0 0
18.
Patagopelta
Genus of nodosaurid dinosaurs
Overview: Patagopelta (meaning "Patagonian shield") is an extinct genus of nodosaurine dinosaur from the Late Cretaceous (upper Campanian–lower Maastrichtian) Allen Formation of Argentina. The genus contains a single ...
0 0
19.
Zhejiangosaurus
Extinct genus of dinosaurs
Overview: Zhejiangosaurus (meaning "Zhejiang lizard") is an extinct genus of nodosaurid dinosaur from the Upper Cretaceous (Cenomanian stage) of Zhejiang, eastern China. It was first named by a group of Chinese ...
0 0
|
||||||
622
|
dbpedia
|
3
| 43
|
https://dokumen.pub/the-princeton-field-guide-to-dinosaurs-course-booknbsped-9781400836154.html
|
en
|
The Princeton Field Guide to Dinosaurs [Course Book ed.] 9781400836154
|
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[
""
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[] | null |
This lavishly illustrated volume is the first authoritative dinosaur book in the style of a field guide. World-renowned...
|
en
|
dokumen.pub
|
https://dokumen.pub/the-princeton-field-guide-to-dinosaurs-course-booknbsped-9781400836154.html
|
Table of contents :
CONTENTS
Preface
Introduction
Group and Species Accounts
Dinosaurs
Theropods
Sauropodomorphs
Ornithischians
Additional Reading
Index
Citation preview
|
|||||
622
|
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3
| 0
|
https://blog.everythingdinosaur.com/blog/_archives/2007/08/17/3163933.html
|
en
|
New Dinosaur Species Found in Eastern China
|
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[
"Mike"
] |
2007-08-17T00:00:00
|
A new dinosaur species from eastern China has been announced by Japanese and Chinese scientists. The dinosaur, a nodosaur has been named Zhejiangosaurus
|
en
|
https://www.everythingdinosaur.com/wp-content/uploads/2016/08/favicon.ico
|
Everything Dinosaur Blog | All about dinosaurs, fossils and prehistoric animals by Everything Dinosaur team members.
|
https://blog.everythingdinosaur.com/blog/_archives/2007/08/17/3163933.html
|
New Dinosaur Species Found in Eastern China – Zhejiangosaurus
A joint Japanese and Chinese team of palaeontologists have announced the discovery of a new dinosaur species that roamed eastern China approximately 100 mya (the Albian stage mid Cretaceous). The animal was found in the south-western region of Zhejiang province, when workmen building a road in 2000 close to the city of Lishui unearthed the first of a series of well preserved bones.
Post cranial bones, parts of the pelvis, the two hind-limbs plus tail and back vertebrae were recovered from the site, enough to permit the scientists to identify this as a brand new species of dinosaur. The animal has been named Zhejiangosaurus lishuiensis (in honour of the province and the nearby city). It was a nodosaurid, an armoured dinosaur similar to the better known ankylosaurs but without the characteristic tail club. The animal believed to be a fully grown adult was over 6 metres long but with a squat gait, typical of a nodosaur only reaching a height of 1 metre at the shoulders.
A Drawing of a Typical Nodosaur
Drawing courtesy of Everything Dinosaur
This peaceful herbivore is a rare find. Nodosaurs are much better known from North America with very finds from what was eastern Laurasia, the only other Chinese nodosaur remains found to date are from the Henan province in central China.
The team’s work has just been published in an English language academic quarterly magazine produced by the Geological Society of China. The fossils (classification code ZNHM M8718), are on display at the Zhejiang Provincial Museum of Natural History and it is hoped that a life size reconstruction of Zhejiangosaurus will be added to the exhibit in the near future.
|
||||
622
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|
https://opendino.wordpress.com/2010/03/05/behold-the-mighty-cladogram/
|
en
|
Behold, The Mighty Cladogram
|
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2010-03-05T00:00:00
|
It's taken awhile, but I finally put together a basic phylogeny to provide the evolutionary backbone for our phylogenetically-informed analyses. References (in very abbreviated format - we'll have to flesh it out more for the paper) are given at the end of this post. I created a "pseudo-supertree", based on a number of what I…
|
en
|
https://secure.gravatar.com/blavatar/30bf5d96fa452ab4bb08005c66a1e27c8f6f745b4c37bbd1122641895441bffa?s=32
|
The Open Dinosaur Project
|
https://opendino.wordpress.com/2010/03/05/behold-the-mighty-cladogram/
|
It’s taken awhile, but I finally put together a basic phylogeny to provide the evolutionary backbone for our phylogenetically-informed analyses. References (in very abbreviated format – we’ll have to flesh it out more for the paper) are given at the end of this post.
I created a “pseudo-supertree”, based on a number of what I (in my opinion) consider to be some of the better and more up-to-date published cladistic analyses. The vast majority of our species of interest are included here, and a draft is shown later in the post. But, we need your help in order to get to the final product.
Here’s What You Can Do
Below is a list of taxa that are not yet placed on the cladogram. In all cases, they were not featured in any of the cladistic analyses I looked at for my first pass. In order to incorporate them into the analysis effectively, though, we need to figure out where they belong. So. . .I’m looking for anything – a reference, personal opinion, cladogram I missed – to place them on the tree. Please mention it in the comments, and we’ll be able to get the tree updated! At the minimum, a reference would be helpful, so that we can be as explicit as possible in our reasoning for the paper. I focused my efforts so far on just ornithischians, so many of the problematic taxa are recently-published animals that are outside Ornithischia.
Also, if you see anything in the tree that you consider to be in gross error, please put a note in the comments section, and we can talk about it. We want the best phylogeny possible. But at the same time, studies have shown that some phylogeny – any phylogeny – is better than no phylogeny at all, so we don’t have to sweat things too much if new data later force us to revise.
Taxa to Place
Ankylosauria: Aletopelta coombsi, Dracopelta zbyszewskii, Dyoplosaurus acutosquameus, Hungarosaurus tormai, Niobrarasaurus coleii, Nodosaurus textilis, Polacanthus foxii, Zhejiangosaurus lishuiensis
Ceratopsia: Graciliceratops mongoliensis, Psittacosaurus major, Psittacosaurus mongoliensis, Psittacosaurus neimongoliensis, Psittacosaurus ordosensis, Psittacosaurus sibiricus, Psittacosaurus sinensis, Psittacosaurus xinjiangensis, Xuanhuaceratops niei
Crurotarsi: Gracilisuchus stipanicicorum, Hallopus victor, Protosuchus richardsoni, Saurosuchus galilei, Pseudolagosuchus major, Scleromochlus taylori
Hadrosauroidea: Anatotitan copei (is there a consensus that this is just Edmontosaurus?), Barsboldia sicinskii, Claosaurus affinis, Edmontosaurus saskatchewanensis (how does it relate to other Edmontosaurus species?), Hadrosaurus foulkii, Mandschurosaurus amurensis, Shantungosaurus giganteus
Basal ornithischians: Eocursor parvus, Geranosaurus atavus
Ornithopods: Draconyx loureiroi, Gongbusaurus wucaiwanensis, Oryctodromeus cubicularis, Xiaosaurus dashanpensis
Parasuchia: Machaeroprosopus gregorii
Sauropodomorpha: Aardonyx celestae, Gyposaurus sinensis, Panphagia protos, Pantydraco caducus, Sellosaurus gracilis
Stegosauria: Chialingosaurus kuani, Lexovisaurus durobrivensis
Theropoda: Guaibasaurus candelariensis, Podokesaurus holyokensis
Logic Behind the Tree
In the interest of Open Notebook Science, I have provided full references and justifications (if any) for the topology of the illustrated tree. All of this should go into the final paper, so that others can reproduce our work.
Overall topology of Ornithischia:
From Butler et al. 2008, Figures 2, 3, and 4, with additional modifications from Butler et al. 2009, Figure S4
Contents of Thyreophora (Lesothosaurus, Scutellosaurus, Emausaurus, Scelidosaurus, Stegosauria, Ankylosauria) based on this phylogeny
Placement of Stenopelix within Pachycephalosauria based on this phylogeny (2009).
Position of Heterodontosauridae follows this reference (2008), as do positions of Stormbergia, Agilisaurus, and Hexinlusaurus.
The position of Othnielia (Othnielosaurus) follows Figures 2 (50% majority rule part), 3, and 4. (2008)
The position of Orodromeus follows Figures 3 and 4. (2008)
The position of Hypsilophodon relative to Jeholosaurus, Yandusaurus, Orodromeus, and Zephyrosaurus follows Figure 2 (50% majority rule part). (2008)
Jeholosaurus and Yandusaurus are arbitrarily placed as sister taxa. Their position basal to Hypsilophodon follows Fig. 2 (50% majority rule part), and Jeholosaurus‘s position more derived than Orodromeus follows Figure 4. (2008)
Bugenasaura is excluded from the tree following its synonymization with Thescelosaurus by Boyd et al. 2009.
The positioning of Thescelosaurus, Parksosaurus, and Gasparinisaura as more derived than Hypsilophodon and outside of the remaining ornithopods follows Figure 2 (50% majority rule part), Figure, and Figure 4. Thescelosaurus is arbitrarily placed as more derived than Parksosaurus+Gasparinisaura. (2008)
The positioning of Talenkauen follows that of Figure 2 (50% majority rule part). (2008)
The remainder of Ornithopoda is consistent across all versions of the cladogram, and this phylogeny is followed for the positions of Rhabdodontidae, Tenontosaurus, Dryosauridae, and Ankylopollexia.
The problematic taxa Zephyrosaurus, Echinodon, Lycorhinus, were excluded (and do not have postcrania, anyhow).
Topology of Stegosauridae:
From 50% Majority-Rule Consensus Tree of Mateus et al. 2009 (Miragaia paper, supplementary information, Figure S7B)
Topology of Ankylosauria:
From Vickaryous 2004, Figure 17.20 (strict consensus tree)
Pawpawsaurus, Sauropelta, and Silvisaurus were in a polytomy; their positions were assigned arbitrarily. This will not matter ultimately, because Pawpawsaurus and Silvisaurus do not have multiple postcranial elements known, and are thus excluded from the quantitative analysis. Saichania and Talarurus had polytomy arbitrarily resolved, too.
Topology of basal Iguanodontia:
Follows Norman 2004, figures 19.21 (strict consensus) and 19.22 (single most parsimonious tree following taxon deletion).
Tenontosaurus is treated as monophyletic, following Figure 19.22.
Euijuubus is arbitrarily treated as more basal than Lurdusaurus.
Jinzhousaurus and Nanyangosaurus are arbitrarily placed as sister taxa, more basal than Probactrosaurus+Ouranosaurus.
Eolambia+Altirhinus are placed between Ouranosaurus and Protohadros, following Figure 19.21 in Norman 2004.
Topology of basal hadrosaurids, hadrosauroids, and other derived iguanodontians:
Follows Dalla Vecchia 2009, Figure 8B (50% Majority Rule Tree). Bactrosaurus and Gilmoreosaurus are arbitrarily resolved from their polytomy with more derived hadrosauroids. Mantellisaurus and Dollodon are given as sister taxa, following this analysis.
Topology of lambeosaurines:
Follows Evans and Reisz 2007, Figure 9
Corythosaurus arbitrarily resolved as closer to Hypacrosaurus than Olorotitan
Pararhabdodon as a basal lambeosaurine after Dalla Vecchia 2009
Topology of hadrosaurines:
Follows Fig. 16 of Gates and Sampson 2007
Edmontosaurus, Prosaurolophus, and Saurolophus are split into their species, assuming that each genus is monophyletic
Topology of Pachycephalosauridae:
Follows Schott et al. 2009 (Colepiocephale description in JVP 29(3)), Figure 8B. Stygimoloch is removed, following synonymization with Pachycephalosaurus by Horner et al. 2009. Colepiocephale is arbitrarily resolved as sister taxon to Stegoceras.
Topology of Ceratopsia:
For non-ceratopsids, follows Makovicky and Norell 2006, Figure 20A (strict consensus tree)
Placement of Cerasinops, and its relationships with Udanoceratops, Leptoceratops, Montanaceratops, and Prenoceratops, based on Figure 6 of Chinnery and Horner 2007
Placement of Yinlong and Micropachycephalosaurus after Figure S4 of Butler et al. supplementary information
Topology of Ceratopsidae:
Chasmosaurinae after Figure 23.8 of Sampson et al. 2004
Centrosaurinae after Figure 12 of Ryan 2007
Placement of Avaceratops based on unpublished analysis
Placement of Turanoceratops after Farke et al. 2009
Topology of Heterodontosauridae:
Follows Figure S4 of Butler et al. 2009 supplementary information.
Abrictosaurus and Tianyulong are placed arbitrarily.
Topology of basal dinosaurs, dinosauriforms, and dinosauromorphs
After Nesbitt et al. 2010, Figure S1
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It's #SciArt week on Twitter!
I think we often downplay or take for granted the role that art plays in science. High quality art is obviously a hugely important aspect of public science communication. A paper describing a new species of dinosaur will have much more impact on the public if it's accompanied by an excellent life restoration of that dinosaur. Astronomers and their spacey kin use illustrations to show us satellites, the solar system, and far-off planets we can't photograph. Biologists dealing with the very small need illustrators to show us the cells in our bodies, what's inside those cells, what DNA looks like and how it works – the list is endless.
But the #SciArt tweet storm happening this week got me thinking again about the role that art plays in my own daily scientific activities. While I don't consider myself an artist, I was always drawing while I was growing up (for a while I entertained the idea of becoming an animator!). And I'm still drawing! Every time I go to a museum, I draw pretty much everything I look at. Why draw when I've got easy access to digital photography? Well, I take tons of photos, too, but drawing makes me LOOK at the specimen.
LOOKING AROUND YOU IS VERY IMPORTANT.
Sketching slows me down, in a good way. What's that weird texture in this part of the bone, how far does this groove extend, what's with this unusual hole in this spot? Is there symmetry? Asymmetry? What's missing, and what's been filled in with plaster? What exactly was I measuring when I say 'length' or 'width'? I've filled many notebooks with drawings, stream-of-consciousness-style notes, measurements, and other bits of data. Mostly I use regular ol' pencils, but I also really like coloured pens and usually travel with a set for annotating my pencil drawings. I would love to be the kind of person that could do watercolour sketching, or proper graphite drawings.
These are some of my earliest notes from my MSc research, from a 2007 visit to the Royal Ontario Museum.
I think, as scientists, we do ourselves a disservice by not teaching students more about art skills and visual design. Being able to quickly and confidently sketch something in front of you is a useful skill to have! And understanding some of the principles of visual design – lines, shapes, negative space, colour combinations, and the like – can only make you a better communicator of science, especially in scientific papers. In addition to just being personally rewarding, drawing makes me a better scientist!
If you're a Twitterer, you should really check out the #SciArt hashtag this week (and into the future), to see the variety of techniques and approaches people take to science art.
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What's up at Wapiti River?
The world can always use some more Pachyrhinosaurus bonebeds. So hooray to my friends and colleagues Federico Fanti and Mike Burns, and my PhD supervisor Phil Currie, for publishing a description of the Wapiti River Pachyrhinosaurus bonebed (currently in 'early view' accepted manuscript form at the Canadian Journal of Earth Sciences).
A friendly Pachyrhinosaurus lakustai greets students at Grande Prairie Regional College!
Most of the time, dinosaur palaeontologists look for bones in dry, barren landscapes – the badlands of Alberta, the Gobi Desert, etc – places that have lots of rocks and not much covering them up, like inconvenient forests or cities. But sometimes, you don't have vast expanses of outcrop. In Nova Scotia, we dig up dinosaurs on the beach. In the area around Grande Prairie, Alberta, you look for bones in the outcrops along rivers and streams.
The very first summer I went out with the University of Alberta crew (way back in the halcyon days of 2007; the first Transformers movie was 'good', everybody read the last Harry Potter book overnight to avoid spoilers, and...apparently not much was happening in my musical spheres, but my, how time has flown), there wasn't a Wapiti River bonebed. We knew that there were bones coming out of the riverbank somewhere, but it took the better part of a day to trace them up the hill to the bone layer.
See if you can spot Phil for scale way up on the hill there, and remember that Phil is about 3x as tall as most humans. That's where the bone layer is!
It's a pretty steep hill, and so those first few days excavating the bone layer meant hacking out little footholds and gradually making enough of a ledge for us to sit on and walk around each other without plummeting to our death.
The last time I was there, in 2011, the ledge had expanded significantly, although you can see it's still a pretty narrow slice! It's a scenic place to work, with the river and boreal forest stretching away below; bear sightings were not uncommon (and occassionally closer than we'd all prefer), and I remember a hummingbird came down to check on us one day, buzzing around my head for a few moments!
In this bonebed, there's a layer of bones in a crazy, mixed-up layer of folded mudstones, and those are pretty easy to excavate.
Here's a dorsal vertebra. Nice and easy.
But down beneath that, the skulls and larger bones are encased within super hard ironstones. We can't really do much with these in the field, so we need to take them out in huge pieces.
And here's what the skulls look like. The circular depression down towards my left foot is the narial opening. The UALVP has like 15 of these suckers and they each take about 2 years to prepare with a crack hammer and chisel.
But the bonebed is also about halfway down into the river valley on a steep slope that's hard enough to just haul yourself up, let alone a huge boulder. So we've been very lucky to have helicopter support to carry out some of the heaviest pieces at the end of each field season.
Up, up and away!
Sometimes we were even visited by Aluk the Pachyrhinosaurus, mascot of the Arctic Winter Games in 2009!
This was probably the strangest day in the field.
There's still much more work to be done on this bonebed – we still aren't exactly sure what species of Pachyrhinosaurus is present. The age is right for P. canadensis, but only time will tell. And with two Pachyrhinosaurus bonebeds in Grande Prairie – the Pipestone Creek bonebed with P. lakustai, and the slightly younger Wapiti River bonebed – there's bound to be much more to learn about the evolution and biology of this unusual ceratopsian.
Previously in Pachyrhinosaurus:
Wapiti River Fieldwork, Part 1
Wapiti River Fieldwork, Part 2
And don't forget to check out:
Fanti F, Currie PJ, Burns ME. 2015. Taphonomy, age, and paleoecological implication of a new Pachyrhinosaurus (Dinosauria: Ceratopsidae) bonebed from the Upper Cretaceous (Campanian) Wapiti Formation of Alberta, Canada. Canadian Journal of Earth Sciences, early view.
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#MuseumWeek Retrospective!
Last week's #MuseumWeek tweetstorm was an awful lot of fun, especially following the #SciArt event just a few weeks earlier. I thought I'd share a couple of photos and thoughts for each day's theme – I didn't manage to post something for each day on Twitter, but I'll fill in some thoughts and photos here!
Day 1: Secrets
One of the nice things about working in the Paleontology & Geology Research Lab at the North Carolina Musuem of Natural Sciences is that "behind the scenes" is part of the scene. You can actually stare at me while I'm working away at my computer each day, if you desire to do such a thing. More interesting, probably, would be to watch our staff, students, and volunteers preparing fossils in the main lab space - secrets waiting to be revealed. But hey, whatever floats your boat!
If you're in Raleigh, stop by and say hi to Carnufex!
Day 2: Souvenirs
I am kind of a Stuff Person and also have a Thing for Museum Gift Shops. As such, I have loads of doodads from my various museum visits. One of the things I like picking up are postcards, especially those that have non-Tyrannosaurus dinosaurs featured on them. For a while, I had these up on my wall at my apartment in Edmonton. Those who have visited my UofA office will also be familiar with my embarassingly large collection of ankylosaur toys, or as I prefer to refer to them, 'scientific models for grown-ups'.
Recognize any museums from your own travels?
Day 3: Architecture
I had a lot of fun with this one on twitter because I LOVE interesting museum architecture. A couple of favourites:
Permian Hall at the Moscow Paleontological Museum:
...which also had custom door hinges, like plesiosaurs!
Dinosaur museum in an old castle in Lerici, Italy:
I wasn't sure about the ROM Crystal at first, but it's grown on me:
And I think the SECU Daily Planet at the NC Museum is pretty swell (on the inside, it's a theatre!):
Day 4: Inspiration
Some non-dinosaur stuff for inspiration day: I really like learning about Canadian art and its history, and one of my very favourite museums on the entire planet is the Museum of Anthropology at the University of British Columbia. If you're in Vancouver, DO NOT MISS THAT MUSEUM. It's an emotional experience to step into the exhibits at this museum and be surrounded by so much creativity and history and skill. Here's a sample to sharpen your brain.
Day 5: Family
I'm lucky to have had great parents that fed my dinosaur obsession as a kid with trips to museums near and far. I'd love to dig out some photos from the before time, but for now, I'll leave this day for my own memories. What are some of your favourite museum memories from your childhood?
Day 6: Favourites
I like busy museums that are crammed full of stuff, especially when that takes the form of a Wall of Stuff or a Hall of Stuff. Here's a few of my favourites.
Hall of Stuff at the Museo de La Plata
Wall of Stuff at the Natural History Museum of LA County
Day 7: Pose
I don't like posting pictures of myself very much, so I'll just include one here to finish off: here's Pinacosaurus (nee "Syrmosaurus") at the museum in Moscow, with me for scale.
That's it for now! What did you share for Museum Week?
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A Brontobyte of Sauropods
Palaeontology emergency alert! This is not a drill! Brontosaurus is back!
YES I FINALLY GOT TO USE THIS ON THE BLOG. Success!
I mean, Brontosaurus never really left. That's the nice thing about taxonomy – once a name is out there, it's there forever, even if we decide later on that it might represent the same kind of animal that another name does. And so every now and then, we get to bring an old name back from the dead. Today, Tschopp and colleagues have published some very good support to indicate that Brontosaurus really is distinct from Apatosaurus after all, and we can all use that name and stop telling people that Brontosaurus isn't real. OMG, WHAT A RELIEF.
To recap: Brontosaurus has not been an accepted name in the palaeontological community for more than 100 years, but because of its use in some museum exhibits, and things like the 1964 World's Fair and the "Rite of Spring" passage in Fantasia, for example, the name has become entrenched in the popular consciousness in a way few other dinosaur names have. It is very disappointing to learn that palaeontologists don't call that big dinosaur Brontosaurus, but the decidedly less evocative name Apatosaurus instead.
Click for sauropod-size. With many thanks to the authors and PeerJ for creating such a useful diagram, which I'm sure will be reproduced often and with much gratitude by palaeontologists, teachers, and other science communicators.
The new paper is staggering in its length (almost 300 pages!) and the amount of work it represents, and I'm not a sauropod specialist, so I'll summarize it here without delving into sauropod anatomy very much:
Two of the Big 3 diplodocids: Apatosaurus (in the back) and Diplodocus (foreground) face-off at the Carnegie Museum.
Tschopp et al. did a specimen-level phylogeny of diplodocids, the sauropods like Apatosaurus and Diplodocus, but not Brachiosaurus or Camarasaurus. This means that individual specimens were coded, rather than species. Often, phylogenetic studies have just looked at the 'classic' diplodocids Apatosaurus, Barosaurus, and Diplodocus (the 'Big 3', shall we say?). And most of those studies elide the many species represented by these three genera. So a specimen-level phylogeny is a much-needed approach to resolve some questions about diplodocid diversity.
They then used some techniques to quantify differences among specimens – pairwise dissimilarity, and apomorphy counts – that would help justify dividing clusters of individuals into different genera. There isn't a rule in palaeontology that individuals need to be a certain amount 'different' from each other in order to be a new genus or species, so the authors looked at how many unique characters separate some sauropods that everyone seems pretty comfortable calling different species and genera. Apatosaurus ajax and Apatosaurus louisae had 12 different features, and Diplodocus carnegii and Diplodocus hallorum had 11 different features. So 13 different characters was set as the baseline for separating out genera in the specimen phylogeny. Using the same approach, they also set 6 differences as the baseline for separating species within a given genus. These numbers only apply to this particular analysis, but it's an interesting approach that I think would be worth considering for other dinosaur phylogenies.
Using this, they wind up doing some taxonomic reshuffling:
a.Diplodocus longus lacks any diagnostic features at the species level and is a nomen dubium, which is bad because it's also the type species for Diplodocus. A petition to the ICZN to switch the type species to D. carnegii is in the works. Diplodocus includes the species D. carnegii and D. hallorum (née Seismosaurus)
b.Dinheirosaurus (from Portugal) is a junior synonym of Supersaurus, and so Supersaurus is a cross-continental genus represented by two species.
c.Diplodocus hayi passes the threshold for generic distinctiveness from Diplodocus and gets a new name, Galeamopus hayi. Specimens of Galeamopus are actually more complete than Diplodocus, which means that Diplodocidae is best represented by Galeamopus at present if you need a diplodocid for whatever you're working on.
d. And finally, and arguably most significantly, Brontosaurus passes the threshold for generic distinctiveness from Apatosaurus. There are three species within Brontosaurus: B. exelsus ('classic'Brontosaurus), B. parvus(née Elosaurus), and B. yahnahpin (née 'Eobrontosaurus').
The third of the Big 3 diplodocids, the iconic rearing Barosaurus at the American Museum of Natural History.
I really hope this taxonomic shuffling gains wide acceptance, because 1) I think their approach and reasoning are pretty sound, and 2) it's going to be SO MUCH EASIER not to have to constantly 'debunk'Brontosaurus with non-palaeontologists.The oft-repeated story that "Brontosaurus" wasn't real because it had the head of one animal and the body of another is wrong, but the real story, about the rules of taxonomy and how we define species, is much more difficult to explain. (It's interesting, but it's not as easily parsed to a lay audience.) And let's face it, Brontosaurus was a really good name and it was sad that it had to be synonymized. The story of Brontosaurus now has a new and interesting chapter – our ideas about the biology of Brontosaurus have changed, but now we can talk about changes to how we think Brontosaurus looked and lived, rather than just focusing on a quirk of taxonomy. So let your Brontosaurus flag fly high, dinosaur fans, because Brontosaurus is back and that's awesome.
Old-timey sauropod in the little diorama at the Smithsonian, back in 2011.
Big taxonomic revisions are hard and important but often don't feel as 'sexy' as some of the other research that gets publicized. I like thinking about alpha taxonomy (uh, perhaps obviously) and I like doing this kind of research, and I think it's really important that we recognize how important this kind of work is – alpha taxonomy is really foundational to a lot of other studies. If you don't know how many species you have, or where they lived, or what anatomy belongs with each species, how can you do projects that look at the evolution of certain features through time, or understand changing ecosystems?
For example, given that there's at least 14 species of diplodocid in only 11 million years of Morrison Formation, it's unlikely that there's a slice of time in there in which there's only one diplodocid species. (And remember, diplodocids weren't the only sauropods in the Morrison – this is also the home of Brachiosaurus and Camarasaurus and Suuwassea and who knows what else.) This is a pretty good reason to reject what I like to call the "Highlander hypothesis", i.e. There Can Only Be One ___(ankylosaur, tyrannosaur, whatever)___ in a given formation, something that I've encountered in conversations on occasion. It's understandable that we would feel unease at the idea of high species/generic diversity in such massive dinosaurs, because how are they dividing up ecosystem space? But over and over again it seems like lots of similarly-shaped dinosaurs were occupying similar times and spaces in terms of what we see in the rock record, which I find very interesting indeed. (Now what we need is a really good stratigraphic framework for putting all of these diplodocids into chronological and geographical context.) We can only do a good job of addressing these kinds of questions by having good data to put into those studies, and that data comes from taxonomic revisions like this one.
And revising taxonomy is probably a never-ending job, because we need to keep reassessing our definitions of genera and species as we get more information through new specimens. Let's make sure we all support this kind of research as palaeontology continues to evolve with new techniques, questions, and approaches. Bully for Brontosaurus, and bully for alpha taxonomy.
Stray observations:
The concept of a 'relatively small' animal that is 12-15 metres long amuses me. (re: Kaatedocus, page 2)
The 'brontobyte' image at the top of this post is an old joke from my Currie lab days; a brontobyte is actually 10^27 bytes. But I think it would be a good collective noun for sauropods, and it also feels appropriate given the large number of sauropod species recovered by Tschopp et al. In fact, we need more collective nouns for dinosaurs, and so I'd like to propose brontobyte for sauropods and armada for ankylosaurs, to join terror of tyrannosaurs.
Go read the paper! It's open access!: Tschopp E, Mateus O, Benson RBJ. 2015. A specimen-level phylogenetic analysis and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda). PeerJ 3:857.
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May your mountains dark and dreary be.
Just wanted to give a quick shout out to some old fossil friends of mine. Horton Bluff/Blue Beach is a pretty cool place and I have fond memories of field trips out there during my Dalhousie days. Between this new paper and the recent paper describing the Permian to Jurassic assemblage of tetrapods, it's been a good time for Nova Scotia palaeontology.
Your friendly neighbourhood ankylosaur palaeontologist, in the before time (i.e. 2003), at Horton Bluff, following in her tetrapod ancestor's footprints. It's goopy there.
Which of course makes me miss it all terribly.
Anderson JS, Smithson T, Mansky CF, Meyer T, Clack J. 2015. A diverse tetrapod fauna at the base of 'Romer's Gap'. PLOS ONE 10:e0125446.
Sues H-D, Olsen PE. 2015. Stratigraphic and temporal context and faunal diversity of Permian-Jurassic continental tetrapod assemblages from the Fundy rift basin, eastern Canada. Atlantic Geology 51:139-205.
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On Surprises
I love surprises. Which is unfortunate for me, because I am extremely bad at being surprised. And it's hard to be surprised by things as you get older, and as easier access to more and more information becomes available to us every day.
But boy, when a surprise comes along that actually takes me by surprise, what a thing to be able to savour.
An extraordinary painting of Yi qi by Emily Willoughby, CC-BY.
So, enter Yi qi. In many ways, it's hardly a surprise at all – numerous artists produced wonderful speculative art about scansoriopterygids predicting membranous wings and/or gliding abilities, and there was even this neat hypothetical Archaeopteryx ancestor that I found in a paper a few years ago. At the time, I wrote on Facebook: "I had not realized that a bat-winged proto-bird was an idea that was on the table!"
(I also wrote, "I like his smirk, lack of neck, and skinny skinny tail", and I agree with Past Victoria about all of those things.)
While Yi qimight not really be a proto-bird, it's still an amazing discovery that shows there was a lot of experimenting with flying and gliding going on back in the Mesozoic, which is perhaps unsurprising, given that lots of disparate groups of animals use gliding to their advantage today – fish, frogs, rodents, marsupials, dermopterans, you name it. And yet, even though there's lots of precedent for gliding vertebrates, and others had predicted something kind of like Yi qibefore, I was still genuinely shocked when I saw the paper and press images. What a great feeling.
What I love best about Yi qi, apart from it's extremely meme-able name, is that it's a great example of maybe what I'll call an 'expected surprise'. A surprise that, as soon as you see it, it seems so obvious and like it should have been there all along. It's like the opposite of a failure of imagination. Surely there is a long German word that captures this specific emotion? What other expected surprises are lurking out there in our futures? What things have we speculated on today, dismissed as being way too out there to take seriously, and yet will pop up as really-for-real things later on?
I guess I need to get to work on some ankylosaur speculative biology! Maybe we'll find the real Yee?
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On Failures of Imagination
Yesterday I talked about 'expected surprises' with regards to Yi qi. Yi qi is a surprise because its anatomy is so unlike other theropods, and it suggests that dinosaurs were experimenting with flight and/or gliding in some ways that were quite different from our current understanding of feather and bird wing evolution. But it was also not entirely unexpected, because scansoriopterygids had super weird anatomy to begin with that gave us enough information to speculate about possible gliding adaptations in those dinosaurs, even though the general consensus was that it was pretty far-fetched.
But today I wanted to talk about a related feeling, which I like to call the Failure of Imagination. Last summer I was working my way through a DVD set of classic sci-fi, fantasy, and adventure movies that I had picked up at some point. I wound up watching a lot of these with friends and basically Mystery Science Theatre 3000-ing the films, and in particular the old space adventure movies from the 40s-60s provided much entertainment. It's really fun to take a look back and see what sorts of things people envisioned the future holding for us – space travel, exoplanet exploration, robots. But what also struck me was the things that the filmmakers and storywriters couldn't even imagine.
They could imagine spaceships and robots, but they couldn't imagine wireless technology. Or storing information in digital form rather than on spools of tape.
They couldn't imagine non-button-and-dial-based instrumentation.
And they definitely couldn't imagine women in roles other than administrative assistants (or as the bad guys). SO MANY SPACE SECRETARIES.
I kept thinking to myself – what sorts of failures of imagination are we having in palaeontology today? We can imagine so many things. But I wonder what kinds of things we won't even know we don't know. When we try our hand at speculative biology, what will scientists 80 or 100 years from now think was charming, or quaint, or ahead of its time. Failures of imagination are one of those things that make me nervous as a scientist, because I don't like the idea that I won't even know what I'm not imagining.
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Crystal Geyser Quarry Quest
I just got back from my first stint of fieldwork for the year, and my first time doing fieldwork in the States. This was just a brief jaunt out to Utah and Colorado for two weeks, but it was a nice sampling of some interesting and different field localities compared to my previous experiences. Today's post: Crystal Geyser quarry in Utah!
So scenic, so majestic. Such altitude.
Crystal Geyser is a non-geothermal, carbon dioxide geyser near Green River, Utah; although we didn't visit the geyser itself, it lends its name to a series of quarries of a massive bonebed in the Yellow Cat Member of the Cedar Mountain Formation (about 125 million years ago). The bonebed is mostly composed of the early therizinosaur Falcarius. The bones in this quarry are incredibly delicate - sometimes even using just a brush through the sediment felt like it was too aggressive! Definitely a challenging site to work at.
Ominous clouds brewed up frequently and then dumped rain and hail on us.
But then sometimes there were rainbows, so I guess it was ok.
We camped in the Morrison Formation and walked up to the Cedar Mountain Formation each day, which was kind of fun.
I'm not accustomed to walking through such a dramatic shift in time and faunas: the Morrison is characterized by lots of classic dinosaurs like Allosaurus and Apatosaurus and Stegosaurus from about 156 to 146 million years ago, but the dinosaurs of the Cedar Mountain Formation have only recently begun to receive much attention and are still poorly known. There's a gap of about 20 million years between the two formations, and in the Yellow Cat Member we find dinosaurs like Falcarius, the ankylosaur Gastonia, the iguanodontians Hippodraco and Iguanacolossus, and dromaeosaurs like Utahraptor and Geminiraptor. The world was changing.
We'll be returning to Utah later in July to work in the Mussentuchit Member. Up next: jackhammering in the Mesaverde Formation of Colorado!
Epilogue: I made a friend at lunchtime one day. D-:
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Mad Max: Fury Quarry
There's a hadrosaur underneath this cliff. (Probably.)
For the second week of our field expedition, we drove from Utah to a site near Rangely, Colorado, to help the Colorado Northwestern Community College with a specimen poking out of a cliff. We've shifted into the Upper Cretaceous here, and are working in the Mesaverde Formation. There's not very much known about the dinosaurs from this formation, so hopefully this will shed some more light on the dinosaurs in this region!
In between Utah and Rangely, we spent a night in Grand Junction and went to see Mad Max: Fury Road, which was way more awesome than any of us had really anticipated and which was basically all we could talk about all week. And so I must also share this great photo that Lindsay took! According to buzzfeed, my Mad Max name is Roop Duststorm, which seems appropriate given the dustiness of working around a rock saw all day.
I did a lot of jackhammering last week, which was great fun if terrible for my lower back, but my favourite thing to do is to pop off the blocks made using the rock saw. You cut a grid into the rock, position your chisel at just the right point, give a couple of hearty whacks with a crack hammer, and off pop these incredibly satisfying 'brownies' of sandstone. It's still slow going, but you can move a lot of rock a lot more quickly this way. (Thanks to Lindsay again for snapping this fun photo!)
Early in the week we were plagued with constant large thunderstorms that rolled in every few hours and made things kind of cold and miserable. Thankfully, this was the last one and it missed us! Instead, it just looked dramatic, which is fine by me.
By last Saturday we had made a lot of progress, although there is still a long way to go to get down to the bone level and (hopefully) find a good dinosaur down there. Best of luck to the crew as they keep working furiously away!
*'Fury Quarry' is also shamelessly stolen from Lindsay.
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Cornelius says hello
Say hello to Cornelius! I got to meet him during a brief visit to the ROM last week, and he seems like a pretty nice guy.
This cool new ceratopsian is on display in the Age of Dinosaurs gallery in an exhibit called "New Dino Discovered", and was also featured in Dino Hunt Canada, which aired earlier this year. It should have a new name soon, but for now Canada voted to nickname it Cornelius. The really nice skeletal mount was put together by Research Casting International based on about a 50% complete disarticulated skeleton.
Here's a close-up of that winning smile. This new dude is a centrosaurine ceratopsid with some pretty neat ornamentation going on at the back of the frill.
I really liked the inclusion of a quarry map on the floor, which highlights some of the bones that are on display. The skeleton was found in southern Alberta in the Milk River area, and comes from the Oldman Formation.
The mounted skeleton is a cast, but there are some original bones on display, like the radius and ulna shown here.
In particular, I liked this set of panels on the wall showing differences in frill ornamentation between centrosaurines, and how we identify different species. On the right is the original frill material for Cornelius, the bottom left is Centrosaurus, and the top left is Styracosaurus.
And look, there was even an ankylosaur osteoderm on display! These are some of the fossils found in the Milk River area, which tell us a bit about the ecosystem that the new centrosaurine lived in.
It's a cool new dinosaur and a nice exhibit, so definitely don't miss it if you're visiting the ROM anytime soon!
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Why does Jurassic World hate dinosaurs?
I have some Thoughts and Feelings about Jurassic World! Spoiler alert, I'm going to talk about details and plot points and this post is really for people who have seen the film. Also, while I'm going to talk about the dinosaurs a bit, this isn't really a review of the science of the film, because that's already been done to death. Ok, onwards and upwards into something that wound up being way too long!
Does Jurassic World hate dinosaurs?
I think the answer to that question is yes. Jurassic World keeps making these little homages and throwbacks to the earlier films (there are lots of shots that echo iconic moments in the earlier films, and some of the plot points mirror the original film almost exactly), and yet I feel like we could consider the theme of Jurassic World to be about rejecting nostalgia and childhood. It's buried under an interesting discussion of the role of the military in funding scientific research, and why some kinds of research are prioritized over others, and it may actually be unintentional, but it's the theme I took away most immediately from this film.
There are two characters that I think are supposed to represent the audience, and neither are treated particularly well by the other characters. And by 'the audience', I'm going to be really self-centered and say that I mean the 30-somethings like myself who saw the original film when we were in that 8-12 year old bracket, or 'peak Jurassic Park' age, and who this film is clearly pandering to. Firstly, we have Gray Mitchell, a 10-ish year old who represents us when we first saw Jurassic Park: he's a dinosaur geek and is one of the only characters to show unrelenting enthusiasm for dinosaurs while visiting Jurassic World. Secondly, we have Lowery, the 30-something computer room dude, who wears an original Jurassic Park shirt and has dinosaur toys on his desk and is obviously super into the dinosaurs in the dinosaur theme park. He is us, now, grown up and nostalgic for the original film. Multiple times throughout the film, Gray's older brother tells him he needs to grow up, and points out that many of the things are for little kids. Claire makes fun of Lowery's shirt, and I think in general we're supposed to think he's kind of a weird man-child who hasn't really grown up.
There's a moment in the film where Gray and his brother Zach stumble upon the old Jurassic Park visitor center building. The T. rex cast skeleton lies on the ground covered in vegetation, and a little piece of the "When Dinosaurs Ruled the Earth" banner is visible. Zach uses it to make a torch so they can investigate the rest of the suspiciously-well-lit ruins. Visiting the old building felt like some gratuitous fan-service to me, but then burning the banner felt like a purposeful statement about rejecting the nostalgia of the original film.
Jurassic World is constantly setting up little nostalgic moments and then seemingly stomping all over them. It's like the filmmakers wanted to pay tribute to Jurassic Park but then were embarassed to show that they liked it – or maybe they didn't really like that movie at all, but wanted to make lots of money (success!). I don't know, but I find it thematically problematic and a bit sad, since the excitement over DINOSAURS! in the first movie is one of the defining aspects of that film, and that sense of wonder and grandeur has rarely been replicated. Jurassic World feels jaded, and like it's too cool for dinosaurs.
Can we talk about ladies in this movie for a moment?
Did we really need to introduce our main female character with the camera sweeping up her legs to her face? Was that absolutely necessary? Also, could we just not use the 'frigid, uptight workaholic woman needs to learn to loosen up and become sexually free with a man, and also needs to remember that all women will have children eventually' stereotype? COULD WE JUST NOT?
It's an intriguing throwback to the original Jurassic Park movie, which I feel successfully used the kids as a character development point for Alan Grant. But Sam Neill managed to portray Grant's discomfort with kids in a more organic way, and the movie gave that plotline a bit of breathing room to develop during some of its quieter moments AND its action sequences (see: sitting in the tree feeding Brachiosaurus; escaping the falling car in the tree; the fence). It's less believable with Claire Dearing, because she doesn't even spend any time with the kids in peril until almost the very end of the movie, at which point she basically worried herself into liking kids? Or something?
Look, not every movie is going to have (or should have) a Strong Female Character(TM), because there are lots of ways to be a lady just like there are lots of ways to be a dude. But the first two Jurassic Park movies had some cool female characters: Ellie Sattler, a palaeobotanist, who was brave and curious and smart! Lex Murphy, who knew those UNIX systems! Sarah Harding, who was a bit foolish but was also brave and curious! Kelly Curtis Malcolm, who gymnastic-ed a Velociraptor to death! In Jurassic World, we get a woman who has great power and authority (she runs a theme park full of dinosaurs!) being told she should be different at almost every opportunity, and we get a distracted babysitter who is killed in the most gratuitous, drawn-out sequence of all. Thanks, movie.
Ok, now let's actually talk about dinosaurs (and other prehistoric creatures) in Jurassic World.
Other palaeontologists have already beaten me to much of this, but I still had a few thoughts I wanted to share. Ultimately I don't have a big problem with the 'retro' dinosaurs of 1993 appearing in this film, because I'm willing to go with the flow in terms of continuity. But there were some pretty dumb things in this film:
· The pterosaur sequence was pretty godawful and brought the action to a screeching halt. I can't suspend disbelief that the pterosaurs would immediately rampage and murder a bunch of people, and I can't suspend disbelief over the physics of that sequence. Refrigerating that babysitter lady was also pretty awful. Sweet jeepers, Jurassic World, you're going to make me say something horrible: this sequence was better in Jurassic Park III. THERE. I hope you're happy.
·I never really bought Indominus rex as anything more than a really big Allosaurus or Saurophaganax. (Sorry, theropod people! Allosaurus is cool, but not, like, THAT cool.) I did, however, like the incorporation of the camouflage idea from the Carnotaurus in the Lost World book, something that I had missed from the film adaptation. Overall, I'm frustrated that Indominus exists mostly so they had a dinosaur they could trademark. Because that's totally what that is, and everything else is secondary to that, including its incorporation into the plot.
·That mosasaur is just so gigantic. I'm on board, but that was starting to stretch credulity as well.
·Why doesn't Rexy eat Blue after the fight? The mind boggles.
Ok, things I liked!
·The Ankylosaurus gives Indominus the old what-for and doesn't immediately die like everything else! Indominus needs to really work at murdering that poor fellow. The design of the Ankylosaurus themselves is pretty terrible (wrong osteoderms, tail too curly, nostrils in the wrong spot, head generally a bit off), although I think it's meant to be consistent with Jurassic Park III.
Here's what Ankylosaurus REALLY looks like!
·Dinosaur petting zoo! It should be for all ages!
·The big kaiju battle between Indominus and Tyrannosaurus was pretty well matched. I liked the little kick to JPIII when the Tyrannosaurus busts through the Spinosaurus skeleton on the way to the fight.
·"Are they safe?""Oh no, under no circumstances, not even a little."
Some final Thoughts and Feelings
I haven't decided yet if I liked Jurassic World. I can't help but think back to the original Jurassic Park with its iconic visual moments and charming, if hokey, dialogue. While it was fun to see an operational Jurassic Park with rides and attractions, I don't feel like Jurassic World had much visual flair. It's really hard to beat dramatic, symbolic visuals like this:
Interesting camera angles like this:
Or quiet moments of terror like this:
And I miss the yellow and green and red colour palette of the original park, replaced here with chrome and blue and silver like every other washed out movie in theatres lately. It is also interesting that all of the big sweeping themes from the original soundtrack are used not for the dinosaurs, but for the manmade structures of the park itself. It really does feel like Jurassic World doesn't care about dinosaurs.
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Dinosaurs Unearthed!
Growing up in Nova Scotia, despite its many excellent and significant palaeontological treasures, meant that there weren't many dinosaur fossils for me to gawp at regularly. The Nova Scotia Museum of Natural History (which I loved) had only a few small fossils on display, and besides an exciting appearance by the Dinosauroid when I was small, did not have any big traveling dinosaur exhibits come through. But when I was in Grade 1 or so, DINAMATION came to town and seared its robotic dinosaurs all over my brain forever. And so I think I will forever have a soft spot in my heart for animatronic dinosaur displays.
Mike Burns and I ran into some of the old Dinamation robots puffing away at the New Mexico Museum back in 2012!
Like Jurassic Forest and Dino Dino Dreampark, the traveling exhibit Dinosaurs Unearthed (at Telus World of Science in Edmonton for the summer) mostly features large animatronic dinosaurs, as well as some casts and interactive displays.
One of the main focuses of the exhibit is showcasing Chinese dinosaurs and fossils, and talking about recent research on the evolution of birds from dinosaurs. Probably one of the best parts of the exhibit is the large number of casts of feathered dinosaurs from China, including Sinosauropteryx, Caudipteryx, Microraptor, and Confuciusornis. These are still not particularly household names, so it's nice to see these on display, especially given the lack of feathers in Jurassic World's dinosaurs!
Just past the casts we have a diorama of Jehol Biota feathered dinosaurs, including (from left to right) the dromaeosaur Microraptor, the compsognathid Sinosauropteryx, the tyrannosaur Dilong, and the bird Confuciusornis. Some of the animatronics are better than others, and all are kind of weirdly oversized, but I think if there was a sign that said this was a diorama at 4x life size or something like that, that it would work pretty well.
My favourite cluster of dinosaurs was the set of Mongolian dinosaurs, including the first time I've ever seen Gigantoraptor anywhere! There's also a pretty dapper Alxasaurus in the front there.
I would have liked to see more cast fossils rather than sculpted reconstructions, and perhaps more fossils overall and a couple fewer animatronics. But generally the information presented in the exhibit was pretty good and had been recently updated, with references to the new research on Brontosaurus, and lots of recent behavioural, biomechanics, and ecology facts as well. Here's a nice display showcasing some of the cool imaging work done by the WitmerLab!
Until next time...watch out for that Shantungosaurus as you leave!
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Woe betide those who summon the Galactic Coelacanth
A couple of years ago I had an existential crisis when I realized that, in the time one of my papers had been in review (almost 8 months!), I could nearly have physically created an entirely new human being in my body, if I had so chosen. Thus began the saddest game in the universe that I like to play when I submit a paper: "What kind of animal could have been gestated in the time this paper has been in review?". And this became an even better running joke when one of my colleagues had a highly unusual review experience that lasted for several years, which completely exhausted the gestation times of real animals.
My amazing and lovely sister saw us talking about this on Facebook and went ahead and wrote an R script that tells you exactly what kind of animal you could have birthed while waiting for reviewer comments. And because I am always forgetting to save this amazing piece of code, I've gotten permission from Jessica to post it here for posterity. My sincere apologies to anyone who gets the Space Whale, and my deepest condolences to anyone who is graced by the presence of the Galactic Coelacanth.
Click here for the R script!
Updated 30 June 2015: If you don't have R, you can also download a text file to see the code!
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Know Your Ankylosaurs: China Edition
I'm in Utah digging up dinosaurs! But also, one of the last big chunks of my PhD thesis has just been published online at the Journal of Systematic Palaeontology. They are generously allowing free access to the paper through the end of August, so head on over and grab a copy while it's free! This time, I'm taking all of the knowledge gained from my previous taxonomic revisions, adding in some more taxa, and doing a revised phylogenetic analysis building on previous analyses to see how everyone shakes out and to learn a little bit more about ankylosaurid biogeography. I'll cover some of the taxonomic stuff over the next few posts, and finish off with the big picture of ankylosaurid evolution.
Pinacosaurus!
I've talked previously about the ankylosaurs of Mongolia, but I've also had the opportunity to study some of their friends from across the border in China. In particular, I got to see lots of specimens of Pinacosaurus, both from the Alag Teeg bonebed in Mongolia, and from Bayan Mandahu in China. Because Pinacosaurus specimens are relatively abundant and usually well preserved, there has already been lots of descriptive work on this taxon, including on the skull (and here, and here), hands and feet, and general postcrania.
Baby Pinacosaurus are so teeny tiny! This one is from Bayan Mandahu and was collected during the Canada-China Dinosaur Project back in the 1980s.
I've discussed just a few new points about Pinacosaurus, especially about how we tell the two species of Pinacosaurus apart. Pinacosaurus grangeri is known from lots of specimens, almost all of which are juveniles; it has relatively short horns at the back of its skull, a constriction in the snout between its nose and its eyes, and a notch in the rough ornamentation above each nostril. Pinacosaurus mephistocephalus is known from just one specimen (also a juvenile), and it has long squamosal horns, no constriction in its snout, and no notch in the ornamentation above each nostril (it looks like it does on one side, but I think this is just damage given that it is not present on the other side). Both species are known from Bayan Mandahu, and so it is reasonable to ask whether or not these could represent the same taxon – given the differences in skull morphology, I suspect we're not looking at intraspecific variation here, although more specimens of P. mephistocephalus would be very helpful in this regard!
Crichtonsaurus becomes Crichtonpelta
Crichtonsaurus is another cool ankylosaur that has received surprisingly little attention given its Jurassic Park affinities. Two species have been named: Crichtonsaurus bohlini (the type species), and Crichtonsaurusbenxiensis. Crichtonsaurus bohlini is, unfortunately, a very incomplete jaw that does not bear any diagnostic features, and so we argue that Crichtonsaurusshould be considered a nomen dubium. Crichtonsaurus benxiensis, on the other hand, is a great specimen with a really good skull and a fair bit of the postcrania, and the skull has some unique features that make it easy to distinguish from other taxa, most specifically the upturned quadratojugal horns. We've proposed the new name Crichtonpelta benxiensis for this material – Crichtonsaurus was a good name and we wanted to keep the replacement name similar, so now we have Crichton's shield instead of Crichton's lizard.
During the Flugsaurier symposium in 2010, while I was visiting Beijing and the IVPP, we took a field trip out to Liaoning and visited the Sihetun Fossil Site. It has a cool museum, including a mounted Crichtonpelta skeleton! I don't think this specimen has been described, but it does corroborate certain aspects of the holotype skull. Crichtonpelta seems to lack discrete caputegulae (tile-like ornamentation) on its skull, which gives it a similar appearance to Pinacosaurus. I don't think the osteoderms have been placed quite correctly on this skeletal mount – I think they've been tipped on their sides so that the keel forms part of the 'base', giving it a somewhat stegosaur-like appearance.
Liaoningosaurus and Chuanqilong
I'm going to talk more about Liaoningosaurus in a few months, but it is one cool little ankylosaur! At only about 30 cm long, the holotype is one of the smallest known ankylosaur specimens and probably represents a very young individual. There may be a few osteoderms in the cervical/scapular region, but that's about it. I've previously argued that the putative plastron in this specimen is more likely skin impressions, which is still pretty cool because we don't have a lot of belly skin for ankylosaurs.
Liaoningosaurus! YAY!
I also wanted to give a shout out to here to Chuanqilong, a larger ankylosaur from Liaoning that was described last summer and which didn't make it into my thesis but which I did include in the revised phylogenetic analysis in the final paper.
Here's Chuanqilong, from Han et al. (2014).
Dongyangopelta, Taohelong, and Sauroplites
Let's finish off this post today with a triad of interesting but enigmatic ankylosaurs. Dongyangopelta and Taohelong are relatively new entries to the world of ankylosaurs, with both taxa appearing in 2013. Neither are particularly complete, but they are interesting because both species possess chunks of fused osteoderms, which would have been found over the pelvis and which are most commonly encountered in nodosaurids and 'polacanthids/polacanthines', and are presently unknown in ankylosaurids – and indeed, Yang et al. described Taohelong as the first example of a polacanthine from Asia. Nodosaurids (including 'polacanthines' as basal taxa within this clade) have been tentatively identified from Asia previously (an interesting but fragmentary specimen from Japan may be a nodosaurid), but to find a Polacanthus-like animal in Asia is unexpected and very interesting. The two species can be differentiated based on the morphology of these pelvic shield pieces. Dongyangopelta comes from the Chaochuan Formation, and another ankylosaur, Zhejiangosaurus, had been named from that formation in 2007; it may eventually shake out that Dongyangopelta is a junior synonym of Zhejiangosaurus, but in the absence of overlapping diagnostic material we opted to keep these taxa separate for now.
Pelvic shield fragments - Dongyangopelta redrawn from Chen et al. (2013), Taohelong redrawn from Yang et al. (2013), and Sauroplites redrawn from Bohlin (1953).
Sauroplites, on the other hand, is a very old name that has been largely overlooked in recent assessments of ankylosaurs. The material was originally described by Bohlin in 1953, but sadly the whereabouts of the original material is unknown today (although there are casts at the American Museum of Natural History). I think Sauroplites was overlooked for a while because it's based off of osteoderms alone, and it's hard to assess diagnostic characters in osteoderms sometimes because they vary so much along the body. This is partly why I like cervical half rings and pelvic shields – in these structures, you can understand the positions of the osteoderms on the body and directly compare patterns and morphologies across different taxa. Supposedly, the osteoderms for Sauroplites were preserved in their original positions when the specimen was excavated, and if so, it's a bit surprising that more of the skeleton was not preserved. Bohlin correctly identified some of these pieces as elements of the sacral armour, and the morphology of these pieces can be used to differentiate Sauroplites from Taohelong and Dongyangopelta, and we consider Sauroplites to be a valid, but poorly known, taxon. It's good to revisit poorly figured and fragmentary taxa from time to time, because new discoveries might help put those pieces in context.
Next time: we head south! See you then!
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Know Your Ankylosaurs: Gondwana Edition
Last time, I talked about the ankylosaurids of China, and today we're talking about Gondwanan ankylosaurs. Gondwana basically refers to the continents of today's southern hemisphere; when the supercontinent Pangaea broke apart, it split into two large continents – Laurasia in the north, and Gondwana in the south. Gondwana includes South America, Africa, Australia, and Antarctica, and, somewhat nonintuitively, India (India kind of beelined into Asia from Australia and that's why we have the Himalayas). Almost all of the ankylosaurs we know about are from the Laurasian continents, which means that the few found in Gondwana are phylogenetically and biogeographically interesting: do they represent southern branches of the ankylosaur family tree, or new migrations into Gondwana from Laurasia? Let's take a closer look:
Minmi paravertebra and Minmi sp.
Minmi is the iconic Australian ankylosaur. Most people, when they think of such things, think of the spectacular referred skeleton with agood skull and in situ armour.
The Smithsonian has a cast of the specimen - here's a section of the ribcage, showing some of the osteoderms in their original arrangement.
Sadly, the holotype is extremely fragmentary and has few elements to make a diagnosis with. Originally, one of the most striking features of Minmi paravertebra was the presence of paravertebral elements, thin rod-shaped bones along the dorsal vertebrae. These were originally interpreted as ossified tendons of the dorsal muscles, and although these are cool to see in Minmi, they are not really unique to Minmi or even to ankylosaurs, since ossified tendons are ubiquitous throughout Ornithischia. One unusual aspect of these ossified tendons is that one set has a flattened, expanded front end. These were interpreted as possible ossified aponeuroses (aponeuroses are sheets of connective tissue in between muscles and tendons). This particular aspect of the ossified tendons IS very unusual, because ossified aponeuroses are extremely rare in animals. While I was hunting around for information about ossified aponeuroses, I came across a very odd case study about mouse deer (Tragulus)– the males completely ossify the aponeuroses above their pelvis and back, creating a carapace-like structure! This is super weird and I would love to investigate this further at some point.
Ossified aponeuroses have since been identified in the European nodosaur Hungarosaurus, which poses a bit of a problem for Minmi: since this feature was one of the only diagnostic characters for Minmi, and since it is now found in an animal that is very unlikely to be Minmi given the spatial and temporal distance between the two, Minmi paravertebra is left without diagnostic characters. A sticky situation that will hopefully be resolved in the future by people who have spent time with the original fossil material!
Antarctopelta
Did you know that the first dinosaur discovered in Antarctica was an ankylosaur? Cryolophosaurus might get all the buzz, but Antarctopelta was first to the press. Antarctopelta is a very interesting little ankylosaur, which I had the chance to study during my visit to Argentina back in 2011. The material is fragmentary but tantalizing, with some pieces of the pelvic armour that are reminiscent of ankylosaurs like Stegopelta and Glyptodontopelta from North America. Unfortunately, in the course of my research I noticed that some of the bones attributed to Antarctopelta and used to help diagnose the taxon didn't quite seem like they came from an ankylosaur. The material was found on an ancient beach strandline with some marine fossils mixed in, and it looks like some of the material originally interpreted as ankylosaurian might be better interpreted as belonging to a mosasaur and a plesiosaur. In the end, we weren't left with any diagnostic characters for Antarctopelta and we should consider that a nomen dubium for now, but there's definitely an Antarctic ankylosaur and I hope at some point some better material is recovered so we can determine the best name for this guy.
The Argentinian ankylosaur
Finally, I also had the chance to study the only described ankylosaur from Argentina. This is also a fairly fragmentary specimen, and it came from a channel lag deposit so it's possible that more than one individual is represented. There are osteoderms, some vertebrae, and a femur, and all are very small – about the same size as the juvenile Anodontosaurus (originally described by Coombs as Euoplocephalus) from Alberta. The femur is interesting because it has some very prominent ridges running lengthwise on it, which seem to be intermuscular lines; these are present but very faint on some other ankylosaurs, and I haven't encountered anything like that in other ankylosaurs. There also may be fragments of the cervical half rings preserved as part of this specimen, since there are some unusual curved osteoderms with multiple peaks and keels. These don't bear any resemblance to other half rings I've looked at, and cervical half ring morphology seems to be taxonomically informative for ankylosaurs. Together, the weird intermuscular lines and unusual cervical half ring fragments might be enough to diagnose the Argentinian specimen as a new taxon, although we withheld from doing so at present.
Here's the specimen on display at the Museo Carlos Ameghino in Cipoletti!
There have been reports of some possible ankylosaur material from India and Madagascar, although much of this material is either very fragmentary (a single tooth from Madagascar), or has not been described (material from India). Stay tuned to find out more about how these rare ankylosaurs fit into the big picture of ankylosaur evolution!
Next up: a grab bag of everybody else!
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Know Your Ankylosaurs: North American Odds and Ends Edition
I've covered many of the North American ankylosaurs in my previous papers and blog posts. In 2013, I argued that what we thought was Euoplocephalus was more likely 4 taxa– Anodontosaurus, Dyoplosaurus, Scolosaurus, and Euoplocephalus proper. Then in 2014 we described a newankylosaurid, Ziapelta, from New Mexico. There are a few other taxa that had previously been proposed to be ankylosaurids, so let's take a look at them here.
Aletopelta, Stegopelta and Glyptodontopelta
Aletopelta is one of the more tantalizingly enigmatic ankylosaurs from North America. It's from a weird place – California – which may have been much further south 75 million years ago compared to its current position. It was also found in marine sediments, and the decaying carcass had formed a little reef, with oysters encrusting the ribs. The only known specimen of Aletopelta is relatively complete, all things considered, with the osteoderms in situ over part of the pelvis, the legs partially articulated, and with various odds and ends like osteoderms and vertebrae. Unfortunately, the ends of the bones are often chewed apart, and some of the material is a bit hard to interpret.
Here's the articulated pelvis and hindlimbs, and some other armour pieces, on display at the San Diego Museum of Natural History.
Regardless, Aletopelta is a very interesting ankylosaur. It has an unusual osteoderm morphology over the pelvis, with small hexagonal osteoderms closely appressed to each other. Ankylosaur pelvic armour seems to come in two major flavours: fused rosettes, like we saw in Dongyangopelta and Taohelong (and perhaps most famously in Polacanthus), and interlocking hexagons, like in Stegopelta, Glyptodontopelta, and Aletopelta. Tracy Ford suggested that ankylosaurs with these hexagonal pelvic shields might represent a clade (dubbed Stegopeltinae) of ankylosaurids. Glyptodontopelta has since typically been interpreted as a nodosaurid, as has Stegopelta, but the most recent interpretation of Aletopelta was that it was an ankylosaurid. In the revised phylogeny in my new paper, we found Stegopelta and Glyptodontopelta as nodosaurids, but Aletopelta as a very basal ankylosaurid. However, although Ford and Kirkland reconstructed Aletopelta with the typical ankylosaurid tail club, I don't think that it possessed one: the preserved distal caudal vertebrae don't show any of the lengthening or other modifications that are characteristic of ankylosaurid handle vertebrae.
An updated restoration of the known elements in Aletopelta - the main differences between this and Ford and Kirkland's reconstruction are the absence of a tail club, and uncertainty over what the head should look like.
Cedarpelta
Cedarpelta is an important taxon for understanding the biogeography and evolution of ankylosaurids, and I wish we had more specimens! I don't have many new comments to add about this taxon, since Ken Carpenter published a great description of the disarticulated skull back in 2001. Cedarpelta has been interpreted as a shamosaurine ankylosaur, as a relative of taxa like Gobisaurus and Shamosaurus (which I'll talk about in the next post) from Asia, and thus may point towards a mid Cretaceous faunal interchange between these two continents. In our revised phylogenetic analysis, we didn't find Cedarpelta as the sister taxon to either Gobisaurus or Shamosaurus, but it does come out as a basal ankylosaurid in their general neighbourhood, and I honestly wouldn't be surprised if future analyses or new taxa show support for it as a shamosaurine ankylosaur after all.
Nodocephalosaurus
Nodocephalosaurus! What a fun ankylosaur. It's really quite unlike the other ankylosaurids from North America, which typically have flat, hexagonal cranial ornamentation. Instead, Nodocephalosaurus has bulbous, conical cranial ornamentation. Bulbous cranial ornamentation is typical of Campanian-Maastrichtian Mongolian ankylosaurs like Saichania and Tarchia, but in those taxa the ornamentation is pyramidal rather than conical. The front end of the snout in Nodocephalosaurus is also unusual, because there's no obvious narial opening and instead the ornamentation has a stepped appearance. Hopefully better specimens with more complete snouts will resolve this weird morphology. I've also reinterpreted the position of the quadratojugal horn compared to Sullivan's original figures – the horn should be rotated forward so that the bottom margin of the orbit is complete.
Nodocephalosaurus holotype skull in dorsal and left lateral views.
Tatankacephalus
I don't have much to say about Tatankacephalus because I didn't look at the original material myself, but the previous phylogenetic analysis by Thompson et al. recovered it as a nodosaurid rather than an ankylosaurid as originally suggested by Parsons and Parsons, and we found the same result. Overall, Tatankacephalus is VERY similar to Sauropelta, so this is perhaps not surprising.
Up next: More odds and ends, but after I return from Utah!
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Know Your Ankylosaurs: Mongolian Odds and Ends Edition
I'm back in civilization, so let's get back to ankylosaurs! Ready Set Go!
Gobisaurus, Zhongyuansaurus, and Shamosaurus
Shamosaurus is a really interesting ankylosaurid from the Zuunbayan Formation of Mongolia. Unlike later ankylosaurids, it still has a relatively long snout like you see in basal ankylosaurs and nodosaurids, and it lacks the distinctive tile-like skull ornamentation of ankylosaurs like Euoplocephalus or Saichania, instead just having a granular, pebbly texture on the skull surface. Gobisaurus, from the Ulansuhai Formation of China, is nearly identical in appearance, and only a few features distinguish these two taxa, namely the length of the tooth row relative to skull length and the orientation of the pterygoids. (Indeed, I think you could make an argument for subsuming Gobisaurus into Shamosaurus as Shamosaurus domoculus, but I'm generally reluctant to start making new combinations given that generic separation is pretty arbitrary anyway.)
Shamosaurus and its too-cool-for-school cervical half rings, on display in Moscow.
Gobisaurus and Shamosaurus are sister taxa; the name Shamosaurinae was proposed at one point and there's no reason to discard it at present even though it only contains two taxa. Shamosaurinae is the sister taxon to Ankylosaurinae. I also identified one new character that links Gobisaurus and Shamosaurus together which isn't present in other ankylosaurids: both taxa have a distinctive groove on each premaxilla, the purpose of which is unknown but there you go. There have been some suggestions that Cedarpelta (from North America) is also a shamosaurine ankylosaurid, and while I find the overall morphology of Cedarpelta to be pretty compelling for placing it in a clade with Gobisaurus and Shamosaurus, I didn't recover it with those taxa in my analysis (it came out more basally-positioned). However, I wouldn't be surprised if Cedarpelta winds up in Shamosaurinae at some point in the future as we find more specimens of both it and Gobisaurus and Shamosaurus.
Zhongyuansaurus was originally described as a nodosaurid ankylosaur partly because of its long snout, but it's indistinguishable from Gobisaurus (except for being smashed and flattened). The holotype is also a subadult (or at least not fully skeletally mature), since some of the cranial sutures are still visible towards the back of the skull. There are some interesting things going on with the postcrania of Zhongyuansaurus, but that's a story for a few weeks from now so STAY TUNED NO SPOILERS IF YOU'VE READ MY THESIS.
Tsagantegia
Of all of the more obscure ankylosaurs I looked at during my PhD, Tsagantegia might be my favourite for being the most surprising in person compared to what I had read about it. Tumanova included a line drawing of the specimen in her original description, which has been oft reproduced, but interestingly it doesn't really do justice to the original specimen (despite being a pretty nice drawing). The line drawing shows a long-snouted ankylosaur with amorphous cranial ornamentation, not dissimilar to Shamosaurus, but with a wider premaxillary beak more typical of later ankylosaurs. In person, however, the skull has distinct cranial caputegulae like we see in Euoplocephalus and Ankylosaurus! It's a pretty cool ankylosaur and I think it's probably really important to understanding the dispersal of ankylosaurs from Asia into North America and the diversification of ankylosaurids in the Campanian-Maastrichtian of Asia, but it's really hard to pin down the age of the Bayan Shiree Formation, and we don't have any postcrania for this taxon. I'm sure I'll be revisiting this guy in the future.
Heck yeah Tsagantegia!
Here it is again but in a more different view!
Talarurus
Way back when I originally started this blog in 2010, I had travelled to Korea to spend some time working in the Hwaseong paleo lab preparing Talarurus material and generally studying the ankylosaur material they had collected from the Gobi. Talarurus, like Tsagantegia, is also from the Bayan Shiree Formation but is clearly distinct. The holotype skull has very subtle cranial ornamentation that takes the form of small cones, rather than flat hexagonal tiles like Euoplocephalus, or bulbous pyramids like Saichania. Weirdly, this configuration is also present in the North American taxon Nodocephalosaurus – either this ornamentation style has convergently evolved, or, as I recovered in my analysis, these two taxa are closely related despite being fairly widely separated geographically and temporally. This is another ankylosaur that I'm sure we'll talk about again.
Talarurus butt in Moscow. The skeleton on display is a composite of several individuals from the same locality, and the skull is totally sculpted and a bit out of date.
Here's the holotype skull, with its weird, weird ornamentation.
Saichania
I've talked about Saichania fairly extensively here last year, but there were a few new things added in this most recent paper: Tianzhenosaurus and Shanxia (both from China) are, most likely, junior synonyms of Saichania, making this the most geographically widespread of the Asian ankylosaurids. Tianzhenosaurus has a nearly identical cranial ornamentation pattern when compared to Saichania, and I couldn't identify any differences that were outside of the usual ornamentation pattern variation we see in something like Euoplocephalus. Shanxia is known from the same formation but from a less well preserved skull, but the morphology of the squamosal horn is consistent with that of both Tianzhenosaurus and Saichania and therefore it probably represents the same taxon.
Next up: what's the big picture here, anyway?
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Know Your Ankylosaurs: Everybody's in this Together Edition
So with all of those posts about ankylosaur taxonomy over the last few weeks, what have we learned about the evolution of this group? Over the course of my PhD research, I was able to identify a bunch of new characters that seemed useful for understanding ankylosaur phylogenetic relationships, including characters related to the cranial ornamentation, pelvis, and osteoderms. Although ornamentation and osteoderms can be tricky, they can still yield useful information if you're careful about how you construct the characters.
Here's a sampling of some of the new characters from the supplementary file that goes along with the paper. Long live rainbow ankylosaur skulls.
With all the new information, here's what the results of the analyses gave us (click to embiggen):
From this, we can take away some interesting points:
1.Gondwanan ankylosaurs are probably not ankylosaurids, but they also don't form a single evolutionary group. Whatever "Minmi" is, it's a very basal kind of ankylosaur, possibly outside the split between Ankylosauridae and Nodosauridae. It's a little bit harder to say what's going on with "Antarctopelta" (previously considered an ankylosaurid), and the Argentinian ankylosaur: both came out as relatively derived nodosaurids, but my dataset isn't designed to test the interrelationships of nodosaurids. I wouldn't be surprised if future analyses incorporating more nodosaurids and more nodosaurid-based characters found that these two species were closely related. It would also be interesting to know which lineage of nodosaurids (probably a lineage from North America) dispersed into South America in the Late Cretaceous in order to give us these two ankylosaurs.
2.There are nodosaurids in the early-mid Cretaceous of Asia, but not necessarily the ones that have been proposed previously. Zhongyuansaurus, for example, was first described as a nodosaurid but is instead a junior synonym of the shamosaurine ankylosaurid Gobisaurus. However, a couple of taxa, like Taohelong, Sauroplites, and Dongyangopelta, are recovered as basal nodosaurids. At present, there doesn't seem to be much overlap between Asian nodosaurids and ankylosaurids, which is interesting! Why didn't nodosaurids hang on in Asia once ankylosaurids evolved, when the two groups seem to have coexisted pretty happily in North America later on?
3.The ankylosaurids from the Late Cretaceous of North America represent a dispersal of Asian ankylosaurines sometime during the early-mid Late Cretaceous. The earliest ankylosaurine is probably Crichtonpelta, from China, and North American ankylosaurines are a deeply nested clade within Ankylosaurinae. We propose the new name Ankylosaurini for the North American ankylosaurines (plus Talarurus, for now).
Here, have some frowny-faced rainbow ankylosaurs. Ankylosaurs are very serious dinosaurs.
4.Where do ankylosaurids first evolve? Unfortunately, that question isn't easy to answer right now: down at the base of Ankylosauridae, there's a mix of taxa from North America and Asia. The position of Gastonia as an ankylosaurid tips the scales slightly in favour of a North American origin for the clade, but some analyses recover this taxon as a nodosaurid, so I think we should be a little cautious about this result. One step up the tree, we've got a polytomy of Aletopelta and Cedarpelta (both from North America) and Liaoningosaurus and Chuanqilong (both from China). Does Ankylosauridae originate in North America with something like Cedarpelta, with a subsequent migration and diversification into Asia? Or does this group originate in Asia with something like Liaoningosaurus and Chuanqilong, and Cedarpelta represents an immigration into North America?
5.And finally, what's going on with ankylosaurids in the mid-Cretaceous of North America? Why don't we find any ankylosaurids between Cedarpelta and the later ankylosaurins? Did 'endemic' North American ankylosaurids go extinct during that time? And why does Aletopelta have such a weird basal phylogenetic position despite being from the Campanian? I don't really have answers for some of these questions, although if you come to the Society of Vertebrate Paleontology meeting in Dallas this October I'm going to try addressing some of them. For now, Aletopelta remains the biggest ankylosaurid enigma to me – it really shares very few things in common with the other Campanian ankylosaurids and I doubt it is an ankylosaurin from the Asian immigration into North America – could it represent a distinctive lineage of North American ankylosaurids stemming from things like Gastonia or Cedarpelta, for which we just don't have other representatives at the moment? Or, is it a nodosaurid masquerading as an ankylosaurid because I haven't sampled the right taxa or characters?
Darn you Aletopelta, why must you vex me so?
As usual, I wind up with more questions than answers every time I try to figure something out.
That wraps up the summaries for this paper, but stay tuned for some more cool research coming out in the next few weeks, and some summer fieldwork recaps!
Arbour VM, Currie PJ. In press. Systematics, phylogeny and palaeobiogeography of the ankylosaurid dinosaurs. Journal of Systematic Palaeontology.
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How the ankylosaur got its tail club
Ankylosaur tail clubs are odd structures, odder than they are usually given credit for. They represent substantial modifications to two different skeletal systems – the endoskeleton, in the form of the caudal vertebrae, and the dermal skeleton, in the form of the caudal osteoderms. The centra of the caudal vertebrae lengthen but stay robust, and the neural arches undergo huge changes, such that the prezygapophyses, postzygapophyses, and neural spine become a robust, V-shaped structure on the top of the centrum, and which creates a tightly interlocking vertebral series with almost no flexibility. We call this the handle of the tail club. The osteoderms at the tip of the tail smush together and two of them become huge: although the tail club knob is small in some species, there are colossal knobs exceeding 60 cm in width. The ankylosaur tail club represents one of the most extreme modifications to the tail in terrestrial tetrapods.
Look at that thing. That is a weird thing.
(This is UALVP 47273, a really nice club that I studied for my MSc work on tail club biomechanics.)
One of the questions I became interested in during my MSc research on ankylosaur tail club biomechanics was how the tail club evolved in the first place. Most ankylosaurs with tail clubs are known from a relatively narrow slice of time right at the end of the Cretaceous, but when and where did the tail club first evolve? Did the stiffening of the tail occur before the enlargement of the tail osteoderms, or vice versa? Or did both changes happen at about the same time? This was a fun question to address during my PhD research, once I had a fairly well resolved phylogeny of ankylosaurids, and once I had looked at tons of ankylosaurid fossils.
So, how did the ankylosaur get its tail club? Well, based on what we see in the fossil record, it looks like the changes to the vertebrae predate the changes to the osteoderms – in other words, the handle comes first and the knob comes later. There is at least one ankylosaur out there that seems to have a tail club handle but not a knob: Gobisaurus!
Hello Gobisaurus! Many many thanks to my friend and colleague Sydney Mohr for preparing this awesome illustration of Gobisaurus for me.
Gobisaurus, a shamosaurine ankylosaurid, has a really nice complete tail club handle that is indistinguishable from other ankylosaurid tail club handles, but does not have a knob. And it's not just because the knob is broken off – it seems as though the last vertebrae in the tail are preserved, because they look very similar to the terminal vertebrae in a CT scan of a tail club from the University of Alberta collections. It's likely that Gobisaurus had osteoderms along the sides of the tail like we see in most other ankylosaurs, but it doesn't appear that there were osteoderms tightly enveloping the tip of the tail.
An even earlier ankylosaur seems to show some changes towards acquiring a tail club handle, as well. Liaoningosaurus, a basal ankylosaurid known only from a very small juvenile, has distal caudal vertebrae where the prezyapophyses extend about 50% the length of the adjacent vertebra. This is what we see in ankylosaurid tail clubs, but not in more basal taxa like Mymoorapelta where the prezygapophyses are much shorter. Liaoningosaurus is missing the tip of the tail and also lacks osteoderms on most of its body because it's a juvenile, so it's harder to say whether or not it had a tail club knob based just on the fossil alone.
I also did a cool and relatively simple thing with my phylogenetic tree to see if I could better understand the likelihood that some ankylosaurs without preserved tail material had a tail club handle or full tail club with a knob. Unsurprisingly, all shamosaurine and ankylosaurine ankylosaurids probably had a tail club handle. Liaoningosaurus is part of a basal polytomy of ankylosaurids, and it was a bit more equivocal whether or not any of these taxa was likely to have a tail club handle or not, partly because another basal ankylosaurid in this region of the tree, Chuanqilong, does not have modified distal caudal vertebrae.
All ankylosaurine ankylosaurids more derived than Pinacosaurus (so including things like Tsagantegia, Saichania, Euoplocephalus, etc.) almost certainly had a tail club knob, and shamosaurine ankylosaurids probably did not. Crichtonpelta, the most basal ankylosaurine, may or may not have had a tail club – we'll need more data to know for sure. There is amounted skeleton of Crichtonpelta at the Sihetun visitor center in Liaoning, and it is shown with a tail club, but it isn't clear whether or not this is sculpted or original material belonging to this specimen, and a full description of this material is necessary.
Gobisaurus and Liaoningosaurus both lived much earlier than the more familiar tail-clubbed ankylosaurs: Gobisaurus is no younger than 92 million years old, and Liaoningosaurus is about 122 million years old. The earliest ankylosaurid with a tail club in the fossil record is Pinacosaurus(from the Campanian), although there is a caveat to this: Talarurus, which is a bit older than Pinacosaurus, should have a full tail club based on its position in the phylogenetic tree, and while a tail club handle is known for this taxon, we haven't found a tail club knob for Talarurus. Talarurus is in kind of a weird spot phylogenetically, since it's from Mongolia but comes out as closely related to North American ankylosaurines, so I think it's worth keeping an eye on this taxon in the future – perhaps Talarurus is another taxon with only a handle and not a knob, which would fit a bit better with its chronologic position if not its phylogenetic position.
Regardless, the changes to the vertebrae of ankylosaurs, starting with Liaoningosaurus at least 122 million years ago and continuing on towards Gobisaurus about 92 million years ago, seem to have occurred long before ankylosaurs evolved a huge osteodermal knob at the end of the tail. Was a stiff tail as good a weapon as a full tail club with a knob? What drove the evolution of the knob so long after the evolution of a stiff handle? And why did ankylosaurs even evolve a tail club at all? Now that I've had fun investigating how ankylosaurs might have used their tails, and how the tail club evolved, the next question feels like it should be 'why'....so stay tuned for more tail club fun over the next year or so as I make an attempt at that question!
Read it for yourself! Arbour VM, Currie PJ. In press. Ankylosaurid dinosaur tail clubs evolved through stepwise acquisition of key features. Journal of Anatomy.
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Snapshots from the Field Museum
Last week I got a chance to visit the Field Museum in Chicago for the first time! It's a great big museum with lots of cool stuff, so I figured I'd share a few impressions from my lunchtime jaunts through the exhibits. Let's get started with all the fossil exhibits outside of the main fossil hall (there are several, but some of them are kind of hidden away!).
SUE
Sue the Tyrannosaurusis most definitely not hidden away, and occupies a place of pride in the museum's main entrance hall. Sue is undeniably a great fossil, although I (and I suspect probably some other palaeontologists as well) have mixed feelings about this fossil: it's incredibly well preserved, but the intense backstory to Sue's acquisition is filled with several unpleasant twists and turns. I'm glad Sue found a home in a museum, but I wish it hadn't been placed up for auction - Sue's auctioning may not have directly led to the trend of putting dinosaurs up for auction for millions of dollars, but I feel like it set a bad precedent all the same.
One thing that's particularly enjoyable about this specific Tyrannosaurus skeleton are the abundant pathologies to be found. Sue has a busted/infected shin, holes in its jaw, and rough bumpy spots on its vertebrae. These vertebrae near the end of the tail have a big mass of crinkly bone around them. It's obvious Sue got up to some trouble during its life, and it's interesting to speculate on the causes of the various oddities in the skeleton (and indeed, others have!).
Extinct Madagascar
Sadly, this exhibit is tucked so far out of the way that basically nobody had wandered back there besides me (you need to go through the conservation gallery to reach it). It's also a little bit specimen-sparse, a trend I've noticed recently in many museums and which I find somewhat concerning. However, I feel like it makes up for the lack of 3D objects in its cool and unusual subject matter - the extinct fauna of Madagascar. The main point to the gallery was showcasing the social media response to new images of Madagascar's prehistory, and the scientific process that went into those images. It was an interesting way to approach the topic, but might have been more compelling with video, audio, or more fossils.
It was pretty cool to see an Aepyornis (elephant bird) egg and life-size silhouette. They really were terrifyingly large and strange birds.
A highlight for me was this Palaeopropithecus skeleton - a lemur that lived and looked like a sloth.
Tracking the Reptiles of Pangea
Tucked away in the African mammals area was a room devoted to palaeontological fieldwork in Tanzania, featuring the newly described silesaurid Asilisaurus! This isn't a skeleton you're going to see in most museums - I only wish more people had been stepping into this little exhibit room to check it out.
A nice touch was showing the original fossil material in its cabinet-ready storage foam. Those are some nice fossils.
And one last fossil....
Seriously, how were these machines not in constant use? They're in the hallway leading towards the bottom-floor cafeteria, and you can get yourself a freshly-made retro Triceratops, Brontosaurus, Tyrannosaurus, or Stegosaurus. I made a Brontosaurus and consider it $2 extremely well spent, especially since it meant I got rid of a bunch of dimes and nickels I didn't know what to do with:
Next time: Evolving Planet!
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Ankylosaurus is a genus of armored dinosaur. Its fossils have been found in geological formations dating to the very end of the Cretaceous Period, about 68–66 million years ago, in western North America, making it among the last of the non-avian dinosaurs. It was named by Barnum Brown in 1908; it is monotypic, containing only A. magniventris. The generic name means "fused" or "bent lizard", and the specific name means "great belly". A handful of specimens have been excavated to date, but a complete skeleton has not been discovered. Though other members of Ankylosauria are represented by more extensive fossil material, Ankylosaurus is often considered the archetypal member of its group, despite having some unusual features.
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Not to be confused with Ankylosuchus.
Ankylosaurus[nb 1] is a genus of armored dinosaur. Its fossils have been found in geological formations dating to the very end of the Cretaceous Period, about 68–66 million years ago, in western North America, making it among the last of the non-avian dinosaurs. It was named by Barnum Brown in 1908; it is monotypic, containing only A. magniventris. The generic name means "fused" or "bent lizard", and the specific name means "great belly". A handful of specimens have been excavated to date, but a complete skeleton has not been discovered. Though other members of Ankylosauria are represented by more extensive fossil material, Ankylosaurus is often considered the archetypal member of its group, despite having some unusual features.
Possibly the largest-known ankylosaurid, Ankylosaurus is estimated to have been between 6 and 8 meters (20 and 26 ft) long and to have weighed between 4.8 and 8 metric tons (5.3 and 8.8 short tons). It was quadrupedal, with a broad, robust body. It had a wide, low skull, with two horns pointing backward from the back of the head, and two horns below these that pointed backward and down. Unlike other ankylosaurs, its nostrils faced sideways rather than towards the front. The front part of the jaws was covered in a beak, with rows of small, leaf-shaped teeth farther behind it. It was covered in armor plates, or osteoderms, with bony half-rings covering the neck, and had a large club on the end of its tail. Bones in the skull and other parts of the body were fused, increasing their strength, and this feature is the source of the genus name.
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