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All modern amphibians are included in the subclass Lissamphibia, which is usually considered a clade, a group of species that have evolved from a common ancestor. The three modern orders are Anura (the frogs), Caudata (or Urodela, the salamanders), and Gymnophiona (or Apoda, the caecilians). It has been suggested that ...
Authorities disagree as to whether Salientia is a superorder that includes the order Anura, or whether Anura is a sub-order of the order Salientia. The Lissamphibia are traditionally divided into three orders, but an extinct salamander-like family, the Albanerpetontidae, is now considered part of Lissamphibia alongside...
Evolutionary history
The first major groups of amphibians ("stem") developed in the Devonian period, around 370 million years ago, from lobe-finned fish which were similar to the modern coelacanth and lungfish. These ancient lobe-finned fish had evolved multi-jointed leg-like fins with digits that enabled them to crawl along the sea bottom...
Many examples of species showing transitional features have been discovered. Ichthyostega was one of the first primitive amphibians, with nostrils and more efficient lungs. It had four sturdy limbs, a neck, a tail with fins and a skull very similar to that of the lobe-finned fish, Eusthenopteron. Amphibians evolved ada...
At the end of the Devonian period (360 million years ago), the seas, rivers and lakes were teeming with life while the land was the realm of early plants and devoid of vertebrates, though some, such as Ichthyostega, may have sometimes hauled themselves out of the water. It is thought they may have propelled themselves ...
After the Carboniferous rainforest collapse amphibian dominance gave way to reptiles, and amphibians were further devastated by the Permian–Triassic extinction event. During the Triassic Period (250 to 200 million years ago), the reptiles continued to out-compete the amphibians, leading to a reduction in both the amphi...
The origins and evolutionary relationships between the three main groups of amphibians is a matter of debate. A 2005 molecular phylogeny, based on rDNA analysis, suggests that salamanders and caecilians are more closely related to each other than they are to frogs. It also appears that the divergence of the three group...
As they evolved from lunged fish, amphibians had to make certain adaptations for living on land, including the need to develop new means of locomotion. In the water, the sideways thrusts of their tails had propelled them forward, but on land, quite different mechanisms were required. Their vertebral columns, limbs, lim...
Characteristics
The superclass Tetrapoda is divided into four classes of vertebrate animals with four limbs. Reptiles, birds and mammals are amniotes, the eggs of which are either laid or carried by the female and are surrounded by several membranes, some of which are impervious. Lacking these membranes, amphibians require water bodie...
Modern amphibians have a simplified anatomy compared to their ancestors due to paedomorphosis, caused by two evolutionary trends: miniaturization and an unusually large genome, which result in a slower growth and development rate compared to other vertebrates. Another reason for their size is associated with their rapi...
The smallest amphibian (and vertebrate) in the world is a microhylid frog from New Guinea (Paedophryne amauensis) first discovered in 2012. It has an average length of and is part of a genus that contains four of the world's ten smallest frog species. The largest living amphibian is the Chinese giant salamander (Andr...
Amphibians are ectothermic (cold-blooded) vertebrates that do not maintain their body temperature through internal physiological processes. Their metabolic rate is low and as a result, their food and energy requirements are limited. In the adult state, they have tear ducts and movable eyelids, and most species have ear...
Anura
The order Anura (from the Ancient Greek a(n)- meaning "without" and oura meaning "tail") comprises the frogs and toads. They usually have long hind limbs that fold underneath them, shorter forelimbs, webbed toes with no claws, no tails, large eyes and glandular moist skin. Members of this order with smooth skins are co...
Anura is divided into three suborders that are broadly accepted by the scientific community, but the relationships between some families remain unclear. Future molecular studies should provide further insights into their evolutionary relationships. The suborder Archaeobatrachia contains four families of primitive frogs...
Caudata
The order Caudata (from the Latin cauda meaning "tail") consists of the salamanders—elongated, low-slung animals that mostly resemble lizards in form. This is a symplesiomorphic trait and they are no more closely related to lizards than they are to mammals. Salamanders lack claws, have scale-free skins, either smooth o...
The suborder Cryptobranchoidea contains the primitive salamanders. A number of fossil cryptobranchids have been found, but there are only three living species, the Chinese giant salamander (Andrias davidianus), the Japanese giant salamander (Andrias japonicus) and the hellbender (Cryptobranchus alleganiensis) from Nort...
The suborder Salamandroidea contains the advanced salamanders. They differ from the cryptobranchids by having fused prearticular bones in the lower jaw, and by using internal fertilisation. In salamandrids, the male deposits a bundle of sperm, the spermatophore, and the female picks it up and inserts it into her cloaca...
The third suborder, Sirenoidea, contains the four species of sirens, which are in a single family, Sirenidae. Members of this order are eel-like aquatic salamanders with much reduced forelimbs and no hind limbs. Some of their features are primitive while others are derived. Fertilisation is likely to be external as sir...
Gymnophiona
The order Gymnophiona (from the Greek gymnos meaning "naked" and ophis meaning "serpent") or Apoda comprises the caecilians. These are long, cylindrical, limbless animals with a snake- or worm-like form. The adults vary in length from 8 to 75 centimetres (3 to 30 inches) with the exception of Thomson's caecilian (Caeci...
Anatomy and physiology
Skin
The integumentary structure contains some typical characteristics common to terrestrial vertebrates, such as the presence of highly cornified outer layers, renewed periodically through a moulting process controlled by the pituitary and thyroid glands. Local thickenings (often called warts) are common, such as those fou...
Amphibian skin is permeable to water. Gas exchange can take place through the skin (cutaneous respiration) and this allows adult amphibians to respire without rising to the surface of water and to hibernate at the bottom of ponds. To compensate for their thin and delicate skin, amphibians have evolved mucous glands, pr...
The skin colour of amphibians is produced by three layers of pigment cells called chromatophores. These three cell layers consist of the melanophores (occupying the deepest layer), the guanophores (forming an intermediate layer and containing many granules, producing a blue-green colour) and the lipophores (yellow, the...
Skeletal system and locomotion
Amphibians have a skeletal system that is structurally homologous to other tetrapods, though with a number of variations. They all have four limbs except for the legless caecilians and a few species of salamander with reduced or no limbs. The bones are hollow and lightweight. The musculoskeletal system is strong to ena...
In most amphibians, there are four digits on the fore foot and five on the hind foot, but no claws on either. Some salamanders have fewer digits and the amphiumas are eel-like in appearance with tiny, stubby legs. The sirens are aquatic salamanders with stumpy forelimbs and no hind limbs. The caecilians are limbless. T...
In frogs, the hind legs are larger than the fore legs, especially so in those species that principally move by jumping or swimming. In the walkers and runners the hind limbs are not so large, and the burrowers mostly have short limbs and broad bodies. The feet have adaptations for the way of life, with webbing between ...
Salamanders use their tails in defence and some are prepared to jettison them to save their lives in a process known as autotomy. Certain species in the Plethodontidae have a weak zone at the base of the tail and use this strategy readily. The tail often continues to twitch after separation which may distract the attac...
Circulatory system
Amphibians have a juvenile stage and an adult stage, and the circulatory systems of the two are distinct. In the juvenile (or tadpole) stage, the circulation is similar to that of a fish; the two-chambered heart pumps the blood through the gills where it is oxygenated, and is spread around the body and back to the hear...
Nervous and sensory systems
The nervous system is basically the same as in other vertebrates, with a central brain, a spinal cord, and nerves throughout the body. The amphibian brain is less well developed than that of reptiles, birds and mammals but is similar in morphology and function to that of a fish. It is believed amphibians are capable of...
Tadpoles retain the lateral line system of their ancestral fishes, but this is lost in terrestrial adult amphibians. Some caecilians possess electroreceptors that allow them to locate objects around them when submerged in water. The ears are well developed in frogs. There is no external ear, but the large circular eard...
The eyes of tadpoles lack lids, but at metamorphosis, the cornea becomes more dome-shaped, the lens becomes flatter, and eyelids and associated glands and ducts develop. The adult eyes are an improvement on invertebrate eyes and were a first step in the development of more advanced vertebrate eyes. They allow colour vi...
Digestive and excretory systems
Many amphibians catch their prey by flicking out an elongated tongue with a sticky tip and drawing it back into the mouth before seizing the item with their jaws. Some use inertial feeding to help them swallow the prey, repeatedly thrusting their head forward sharply causing the food to move backwards in their mouth by...
Amphibians possess a pancreas, liver and gall bladder. The liver is usually large with two lobes. Its size is determined by its function as a glycogen and fat storage unit, and may change with the seasons as these reserves are built or used up. Adipose tissue is another important means of storing energy and this occurs...
There are two kidneys located dorsally, near the roof of the body cavity. Their job is to filter the blood of metabolic waste and transport the urine via ureters to the urinary bladder where it is stored before being passed out periodically through the cloacal vent. Larvae and most aquatic adult amphibians excrete the ...
Respiratory system
The lungs in amphibians are primitive compared to those of amniotes, possessing few internal septa and large alveoli, and consequently having a comparatively slow diffusion rate for oxygen entering the blood. Ventilation is accomplished by buccal pumping. Most amphibians, however, are able to exchange gases with the wa...
Reproduction
For the purpose of reproduction most amphibians require fresh water although some lay their eggs on land and have developed various means of keeping them moist. A few (e.g. Fejervarya raja) can inhabit brackish water, but there are no true marine amphibians. There are reports, however, of particular amphibian populatio...
Several hundred frog species in adaptive radiations (e.g., Eleutherodactylus, the Pacific Platymantis, the Australo-Papuan microhylids, and many other tropical frogs), however, do not need any water for breeding in the wild. They reproduce via direct development, an ecological and evolutionary adaptation that has allow...
In the tropics, many amphibians breed continuously or at any time of year. In temperate regions, breeding is mostly seasonal, usually in the spring, and is triggered by increasing day length, rising temperatures or rainfall. Experiments have shown the importance of temperature, but the trigger event, especially in arid...
In caecilians, fertilisation is internal, the male extruding an intromittent organ, the , and inserting it into the female cloaca. The paired Müllerian glands inside the male cloaca secrete a fluid which resembles that produced by mammalian prostate glands and which may transport and nourish the sperm. Fertilisation pr...