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Update app.py
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app.py
CHANGED
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@@ -4,6 +4,85 @@ import streamlit as st
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summarizer = pipeline("summarization", model="facebook/bart-large-cnn")
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ARTICLE = """
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. Across the history of limnology, small and shallow waterbodies
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are widely referred to as ponds8–11, yet pond defnitions difer across the globe and are not based on scientifc
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evidence. Te lack of a universal, scientifcally-based pond defnition that diferentiates ponds from other lentic
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@@ -27,5 +106,515 @@ from scientifc literature and evaluated legislative defnitions of ponds, wetland
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a large dataset of pond characteristics and ecosystem function from a global literature survey and compared
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ecosystem structural and functional metrics among ponds, wetlands, and lakes. Finally, we propose an evidencebased pond defnition.
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Results and discussion
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| 30 |
"""
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-
st.write(summarizer(ARTICLE, max_length=
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summarizer = pipeline("summarization", model="facebook/bart-large-cnn")
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ARTICLE = """
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+
1
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+
Vol.:(0123456789)
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+
Scientifc Reports | (2022) 12:10472 | https://doi.org/10.1038/s41598-022-14569-0
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| 10 |
+
www.nature.com/scientificreports
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| 11 |
+
A functional defnition
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+
to distinguish ponds from lakes
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+
and wetlands
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+
David C. Richardson 1,19*, MeredithA. Holgerson 2,19, Matthew J. Farragher 3
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+
,
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+
Kathryn K. Hofman 4
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| 17 |
+
, Katelyn B. S. King 5
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+
, María B.Alfonso 6
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| 19 |
+
, Mikkel R.Andersen 7
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+
, Kendra Spence Cheruveil 8
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+
, KristenA. Coleman9
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| 22 |
+
, Mary Jade Farruggia 10,
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+
Rocio Luz Fernandez 11, Kelly L. Hondula 12, GregorioA. López Moreira Mazacotte 13, Katherine Paul1
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+
, Benjamin L. Peierls 14, Joseph S. Rabaey 15, Steven Sadro 10,
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+
María Laura Sánchez 16, Robyn L. Smyth 17 & Jon N. Sweetman 18
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+
Ponds are often identifed by their small size and shallow depths, but the lack of a universal evidencebased defnition hampers science and weakens legal protection. Here, we compile existing pond
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+
defnitions, compare ecosystem metrics (e.g., metabolism, nutrient concentrations, and gas fuxes)
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among ponds, wetlands, and lakes, and propose an evidence-based pond defnition. Compiled
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defnitions often mentioned surface area and depth, but were largely qualitative and variable.
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+
Government legislation rarely defned ponds, despite commonly using the term. Ponds, as defned in
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published studies, varied in origin and hydroperiod and were often distinct from lakes and wetlands
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+
in water chemistry. We also compared how ecosystem metrics related to three variables often
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seen in waterbody defnitions: waterbody size, maximum depth, and emergent vegetation cover.
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+
Most ecosystem metrics (e.g., water chemistry, gas fuxes, and metabolism) exhibited nonlinear
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relationships with these variables, with average threshold changes at 3.7± 1.8 ha (median: 1.5 ha)
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in surface area, 5.8 ± 2.5 m (median: 5.2 m) in depth, and 13.4 ± 6.3% (median: 8.2%) emergent
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vegetation cover. We use this evidence and prior defnitions to defne ponds as waterbodies that are
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small (< 5 ha), shallow (< 5 m), with< 30% emergent vegetation and we highlight areas for further
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+
study near these boundaries. This defnition will inform the science, policy, and management of
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+
globally abundant and ecologically signifcant pond ecosystems.
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+
Lentic (still) waterbodies have long been placed into categories to improve our understanding of aquatic ecosystems, aid science communication, and facilitate management decisions1,2
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+
. For instance, lentic ecosystems
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+
have been sorted into discrete categories by size or depth3,4
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+
, trophic status5
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+
, and mixing regime6,7
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+
. Ofen, lentic
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+
waterbodies are categorized by diferent ecosystem types, such as lakes, ponds, and wetlands (Fig. 1). Categorizing
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| 48 |
+
OPEN
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+
1
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| 50 |
+
Biology Department, State University of New York at New Paltz, New Paltz, NY, USA. 2
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| 51 |
+
Department of Ecology
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| 52 |
+
and Evolutionary Biology, Cornell University, Ithaca, NY, USA. 3
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| 53 |
+
School of Biology and Ecology, Climate Change
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| 54 |
+
Institute, University of Maine, Orono, ME, USA. 4
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+
Departments of Biology and Environmental Studies, St. Olaf
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+
College, Northfeld, MN, USA. 5
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| 57 |
+
Department of Fisheries and Wildlife, Michigan State University, East Lansing,
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| 58 |
+
MI, USA. 6
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| 59 |
+
Instituto Argentino de Oceanografía (IADO), Universidad Nacional del Sur (UNS)-CONICET, Florida
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| 60 |
+
8000, Complejo CCT CONICET Bahía Blanca, Edifcio E1, B8000BFW Bahía Blanca, Argentina. 7
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| 61 |
+
Centre for
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| 62 |
+
Freshwater and Environmental Studies, Dundalk Institute of Technology, Dundalk, Ireland. 8
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| 63 |
+
Department
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| 64 |
+
of Fisheries and Wildlife and the Lyman Briggs College, Michigan State University, East Lansing, MI,
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| 65 |
+
USA. 9
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+
Department of Geography, York University, Toronto, ON, Canada. 10Department of Environmental Science
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| 67 |
+
and Policy, University of California, Davis, Davis, CA, USA. 11National Scientifc and Technical Research Council
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| 68 |
+
(CONICET), Cordoba, Argentina. 12Battelle, National Ecological Observatory Network (NEON), Boulder, CO,
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| 69 |
+
USA. 13Department of Ecohydrology and Biogeochemistry, Leibniz-Institute of Freshwater Ecology and Inland
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| 70 |
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Fisheries (IGB), Müggelseedamm 310, 12587 Berlin, Germany. 14Lakes Environmental Association, Bridgton,
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+
ME, USA. 15Department of Ecology, Evolution, and Behavior, University of Minnesota-Twin Cities, St. Paul, MN,
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| 72 |
+
USA. 16CONICET-Universidad de Buenos Aires, Buenos Aires, Argentina. 17Environmental and Urban Studies, Bard
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| 73 |
+
College, Annandale‑on‑Hudson, NY, USA. 18Department of Ecosystem Science and Management, Penn State
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| 74 |
+
University, University College, PA, USA. 19These authors contributed equally: David C. Richardson and Meredith
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| 75 |
+
A. Holgerson. *email: richardsond@newpaltz.edu
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+
2
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| 77 |
+
Vol:.(1234567890)
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| 78 |
+
Scientifc Reports | (2022) 12:10472 | https://doi.org/10.1038/s41598-022-14569-0
|
| 79 |
+
www.nature.com/scientificreports/
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| 80 |
+
waterbodies using physical and biological characteristics facilitates generalizations and decision making, but
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| 81 |
+
categories may not always align with ecological inferences2
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+
.
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+
Categorizing small waterbodies is particularly challenging. Te majority of the world’s lentic waterbodies are
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small: over 95% are less than 10 ha (0.1 km2
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+
)3,8
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| 86 |
. Across the history of limnology, small and shallow waterbodies
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| 87 |
are widely referred to as ponds8–11, yet pond defnitions difer across the globe and are not based on scientifc
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| 88 |
evidence. Te lack of a universal, scientifcally-based pond defnition that diferentiates ponds from other lentic
|
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|
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| 106 |
a large dataset of pond characteristics and ecosystem function from a global literature survey and compared
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| 107 |
ecosystem structural and functional metrics among ponds, wetlands, and lakes. Finally, we propose an evidencebased pond defnition.
|
| 108 |
Results and discussion
|
| 109 |
+
Current scientifc defnitions of ponds. We compiled existing scientifc defnitions of ponds by conducting a backwards and forwards search of papers referenced in or subsequently referencing three seminal
|
| 110 |
+
pond papers8,17,18 (see “Methods”). We ultimately compiled 54 pond defnitions from scientifc literature (data
|
| 111 |
+
available19). Te variables most ofen included in defnitions were surface area (91% of defnitions), depth (48%),
|
| 112 |
+
permanence (48%), origin (i.e., natural or human-made; 33%), and standing water (33%; Fig. 2a). When surface
|
| 113 |
+
area or depth were included in defnitions, they were ofen mentioned qualitatively (e.g., “small” and “shallow”).
|
| 114 |
+
Of the 61% of defnitions that included a maximum pond surface area, the range was 0.1 to 100 ha, the median
|
| 115 |
+
was 2 ha, and all but two defnitions were≤10 ha (Fig. 2b). For depth, only 17% of studies provided a maximum
|
| 116 |
+
depth cutof, which ranged 2 to 8 m (Fig. 2c). Of the 26 defnitions mentioning permanence, 22 stated that ponds
|
| 117 |
+
could be temporary or permanent and only three indicated that ponds are exclusively permanent waterbodies.
|
| 118 |
+
Of the 18 defnitions mentioning origin, 17 mentioned that ponds could be natural or human-made with the
|
| 119 |
+
remaining study indicating ponds can have diverse origins.
|
| 120 |
+
Other important factors included in defnitions related to morphometry. For example, 30% of defnitions
|
| 121 |
+
mentioned the potential for plants to colonize the entire basin, which relates to high light penetration (mentioned
|
| 122 |
+
in 11% of defnitions) and/or shallow depths. For example, Wetzel11 defnes ponds as having enough light penetration that macrophyte photosynthesis can occur over the entire waterbody. As such, these conditions may be
|
| 123 |
+
Figure 1. We call lentic waterbodies by a variety of names in the English language including ponds, lakes,
|
| 124 |
+
wetlands, reservoirs, oxbows, prairie potholes, vernal pools, lagoons, dams, puddles, and shallow lakes. Tese
|
| 125 |
+
names may or may not correspond to ecological and systematic diferences. Generally, laypeople and experts, as
|
| 126 |
+
individuals, will quickly diferentiate among broad categories of ponds, lakes, and wetlands; however, individuals
|
| 127 |
+
may respond in diferent ways depending on their background and experiences. We present three diferent
|
| 128 |
+
images of waterbodies that could each be categorized as lake, pond, or wetland using objective (e.g., morphology
|
| 129 |
+
or vegetative cover) or more subjective criteria keeping cognizant of the complexity within and potential overlap
|
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+
among waterbody types.
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+
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+
comparable to the littoral region of lakes (11% of defnitions). Lastly, 7% of pond defnitions mentioned mixing
|
| 136 |
+
versus stratifcation, whereby ponds mix more than lakes20 yet less than shallow lakes due to a smaller fetch16.
|
| 137 |
+
To assess if there was agreement in pond defnitions among papers, we examined the number of times each
|
| 138 |
+
defnition was cited. Across the 54 defnitions, there were 89 citations of 48 unique papers. Ultimately, most
|
| 139 |
+
papers (75%) were only cited only once, indicating no consensus in pond defnition. Te most cited paper was
|
| 140 |
+
Biggs et al.21, which accounted for 15% of citations. Te next two most cited papers were Oertli et al.17 and Sondergaard et al.18, which were seminal papers included in our backwards-forwards search, and each comprised
|
| 141 |
+
8% of citations.
|
| 142 |
+
International defnitions. At an international level, there is no consensus on how to discriminate among
|
| 143 |
+
ponds, lakes, and wetlands. In North America, wetlands are generally considered to be shallow:<2 m in Canada22
|
| 144 |
+
and<2.5 m in the US23, which diferentiates them from lakes. Some nations, such as Australia, South Korea, and
|
| 145 |
+
Uganda, explicitly include ponds and lakes in federal wetland defnitions24 (see also22). Te inclusion of ponds
|
| 146 |
+
and some lakes within wetland defnitions ofen stems from the Ramsar Convention, an international body
|
| 147 |
+
interested in global wetland conservation that has been signed by 172 countries representing 6 continents as
|
| 148 |
+
of 202125. Te Ramsar Convention defned wetlands as “areas of marsh, fen, peatland, or water” across marine,
|
| 149 |
+
brackish, and freshwater with varying degrees of permanence and natural or artifcial states with a maximum
|
| 150 |
+
depth of 6 m26, which overlaps depths found in many defnitions of ponds and shallow lakes. In other countries,
|
| 151 |
+
ponds are included in lake defnitions under federal conservation laws. For example, in the Danish “nature
|
| 152 |
+
protection” law §3, lakes are defned as waterbodies with a surface area of>100 m2
|
| 153 |
+
. As 98% of Danish ‘lakes’ are
|
| 154 |
+
smaller than 1 ha27, this law protects many small waterbodies that may be considered ponds elsewhere. Still other
|
| 155 |
+
agencies have only qualitative pond defnitions: the European Commission simply defnes ponds as “relatively
|
| 156 |
+
shallow” and may also be called “pool, tarn, mere, or small lake,” a defnition also used by the International
|
| 157 |
+
Union for Conservation of Nature28,29. Tese examples underscore that waterbody defnitions vary globally, are
|
| 158 |
+
generally qualitative, and are rarely based on scientifc evidence relating to ecosystem structure or function. Te
|
| 159 |
+
defnitions possibly derive from diferent management, protection, and monitoring strategies; for instance, the
|
| 160 |
+
European Union’s Water Framework Directive excludes waterbodies<50 ha (0.5 km2
|
| 161 |
+
) in size from monitoring30.
|
| 162 |
+
Figure 2. Summary of “pond” defnitions from scientifc literature including (a) presence of various
|
| 163 |
+
morphological, biological, and physical characteristics in the defnition as blue bars (n=54 defnitions total).
|
| 164 |
+
Bold black lines indicate the number of defnitions with surface area and depth values. Histograms of the upper
|
| 165 |
+
limits from “pond” defnitions for (b) surface area and (c) maximum depth.
|
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+
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|
| 170 |
+
Current U.S. Federal and State defnitions. In the US, waterbody defnitions vary among federal agencies, with implications for both legal protection and monitoring. Te US Environmental Protection Agency
|
| 171 |
+
(EPA) and the US Army Corps of Engineers (ACE) defne wetlands based on saturated soils and hydrophytic
|
| 172 |
+
vegetation, which has the potential to include ponds within the category of wetlands. Conversely, the US Fish
|
| 173 |
+
and Wildlife Service (USFWS) distinguishes among wetlands and lakes based on surface area, depth, and emergent vegetation31. Lakes are≥8 ha or if smaller, they must be≥2.5 m in maximum depth. In contrast, wetlands
|
| 174 |
+
are are dominated by>30% emergent plant cover; if there is less, the site may still be a wetland if<2.5 m deep
|
| 175 |
+
and<8 ha in size. Terefore, ponds are ofen considered by USFWS to be wetlands, but this is not always the case:
|
| 176 |
+
ponds have been used as an example of a waterbody that can be classifed as lake, wetland, or both32. Te lack of
|
| 177 |
+
an explicit, unifed, and scientifcally based pond defnitions across three federal agencies (EPA, ACE, USFWS)
|
| 178 |
+
is confusing and contributes to ponds being underrepresented in US aquatic waterbody monitoring relative to
|
| 179 |
+
their numerical dominance on the landscape3,8
|
| 180 |
+
. For example, US EPA monitoring programs include ponds in
|
| 181 |
+
both the National Wetland Condition Assessment and the National Lake Assessment; however, “ponds” represent a small number of waterbodies in each of these surveys (<12% classifed qualitatively as “pond” in 2011
|
| 182 |
+
wetland survey; 13% of waterbodies were<5 ha in 2012 lake survey).
|
| 183 |
+
Refecting political and geographic variability at the national scale, most US states have their own waterbody
|
| 184 |
+
protections33. We surveyed US state agencies to examine state defnitions of ponds, lakes, and wetlands (see
|
| 185 |
+
“Methods”). Our survey responses included 42 of 50 (84%) states (Fig. 3). Only one state (Michigan) explicitly
|
| 186 |
+
defned ponds, 11 states defned lakes (26%), and 30 states defned wetlands (71%). While only one state defned
|
| 187 |
+
ponds, half of the surveyed states used the term “pond” in their legislation. Specifcally, ponds were referenced
|
| 188 |
+
as state waters (e.g., Vermont) or were included in state defnitions for lakes (e.g., Kansas) or wetlands (e.g.,
|
| 189 |
+
Rhode Island). It is unclear how these defnitions impact monitoring and protection or why the distinctions
|
| 190 |
+
were originally made. For instance, many states monitor lakes based on minimum size thresholds, which vary
|
| 191 |
+
widely from < 1 ha in Arizona and Alaska, 2–4 ha in many northeastern states, and up to 8 ha in Washington
|
| 192 |
+
and Nebraska. Te variety of defnitions and monitoring size cutofs do not appear to be scientifcally based,
|
| 193 |
+
but may stem from arbitrary decisions, historic references, mapping capabilities from decades ago, and resource
|
| 194 |
+
limitations for monitoring; the same rationale for defnitions likely apply to local, regional, and international
|
| 195 |
+
organizations around the globe.
|
| 196 |
+
Comparing lake, pond, and wetlands characteristics from literature. We compared biological,
|
| 197 |
+
physical, and chemical characteristics of waterbodies that scientists called lakes, ponds, or wetlands in published
|
| 198 |
+
studies. To obtain data for the pond characteristics, we used the same literature search summarized above for
|
| 199 |
+
pond defnitions (also, see “Methods”). From the 519 papers that we examined, we extracted data on sites the
|
| 200 |
+
authors called “ponds” and other variants (e.g., ‘small ponds’, ‘fsh ponds’, but NOT ‘lakes’). We fltered waterbodies that were≤20 ha surface area and≤9 m depth (global distribution; n=1327) to include waterbodies
|
| 201 |
+
slightly greater than the maximum depth and maximum surface area used to defne ponds in prior studies34,35.
|
| 202 |
+
To compare ponds to lakes and wetlands, we used existing lake (US and Europe; n=55,173) and wetland (US;
|
| 203 |
+
n=400) databases; waterbodies were classifed as lake or wetland by the scientists or managers who published
|
| 204 |
+
the database. Wetlands were classifed as<1 m in depth with no defned surface area and lakes were all>0.02 ha
|
| 205 |
+
with no defned depth (see “Methods’’ for details).
|
| 206 |
+
From the waterbodies that scientists called “ponds,” hydroperiod and origin varied over a large range of
|
| 207 |
+
characteristics. Of the 608 ponds with hydroperiod data, permanent ponds accounted for 74% (n =450) and
|
| 208 |
+
temporary ponds for 26% (n=158). Out of 648 ponds with known origins, 65% (n=418) were constructed or
|
| 209 |
+
manipulated and 35% (n=230) were natural. Terefore, scientists consider ponds to be inclusive of both permanent and temporary hydroperiod and have natural or human-made origins.
|
| 210 |
+
Figure 3. US state responses to surveys indicating if the state has a defnition of wetland, lake, or pond and if
|
| 211 |
+
the state used the term “pond” in their legislation. NR=no response.
|
| 212 |
+
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+
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|
| 216 |
+
When examining water chemistry, nutrients, and biotic data across diferent waterbody types, as defned by
|
| 217 |
+
publishing scientists and managers, we found that ponds were distinct from lakes and wetlands in two metrics
|
| 218 |
+
(TN, pH), similar to wetlands in one metric (TP), and similar to lakes in one metric (chl a; Fig. 4; Tables S1, S2).
|
| 219 |
+
For example, ponds had distinctly high TN concentrations, which were greater than either lakes or wetlands
|
| 220 |
+
(Fig. 4b; Table S2). Ponds and wetlands had similarly high TP concentrations, which were signifcantly greater
|
| 221 |
+
than lakes; ponds were also most variable in TP (Fig. 4a; Table S2). Lastly, ponds chlorophyll (chl) a concentrations were similar to lakes, with wetlands being most variable but lower, on average (Fig. 4d; Table S2).
|
| 222 |
+
Does ecosystem structure and function distinguish ponds from lakes and wetlands? We evaluated the relationship between key metrics of ecosystem structure or function with three quantitative variables
|
| 223 |
+
that ofen showed up in pond, lake, or wetland defnitions: surface area, maximum depth (hereafer depth), and
|
| 224 |
+
emergent vegetation cover. Our metrics of ecosystem structure or function include nutrients (total phosphorus
|
| 225 |
+
(TP), total nitrogen (TN)), water chemistry (pH), primary producer biomass (chl a), metabolism (gross primary
|
| 226 |
+
production—GPP, respiration—R, net ecosystem production—NEP), and heat and gas distributions and movement (diel temperature ranges—DTR, methane fuxes, gas transfer velocities). Te data was collated from global
|
| 227 |
+
surveys of literature and federal or international databases (see “Methods”) with ultimately ten comparisons for
|
| 228 |
+
surface area, six comparisons for depth, and four comparisons for emergent vegetation cover with a range of
|
| 229 |
+
sample sizes for each comparison (n=67 to 7931, see Tables S3, S5, S7). We assessed each relationship for four
|
| 230 |
+
Figure 4. Comparison of various chemical and biological parameters across wetlands, ponds, and lakes, with
|
| 231 |
+
waterbody category based on the term used by publishing scientists and managers (Table S2). Violin plots
|
| 232 |
+
indicate distributions of waterbody characteristics, the white box indicates 25th to 75th percentile with median
|
| 233 |
+
in the middle, whiskers indicate 1.5×interquartile range, and outliers are black closed circles. Letters inside the
|
| 234 |
+
plot indicate signifcant diferences in means (LSD, alpha=0.05). Note all x-axes have logarithmic scales.
|
| 235 |
+
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+
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|
| 239 |
+
diferent patterns in increasing order of complexity: null, linear, segmented (nonlinear), and logistic (nonlinear)
|
| 240 |
+
patterns and selected the best ft and most parsimonious relationship.
|
| 241 |
+
Ecosystem structure and function were mostly nonlinearly related to surface area (n=9/10 variables), depth
|
| 242 |
+
(n=5/6 variables), and emergent vegetation cover (n=3/4 variables) with both segmented and logistic relationships occurring (Figs. 5, 6, 7; Tables S3–S8). For surface area, six variables had logistic relationships: TP (Fig. 5b),
|
| 243 |
+
methane fuxes (Fig. 5d), chl a (Fig. 5f), diel temperature range (Fig. 5h), gas exchange rates (k600; Fig. 5i), and
|
| 244 |
+
pH (not pictured). Te infection occurred at 0.8 ha for TP, 1.1 ha for methane fuxes, 1.5 ha for chl a, 1.7 ha for
|
| 245 |
+
pH, 4.6 ha for diel temperature range, and 17.5 ha for gas exchange rates (Table S3). NEP (Fig. 5c), R (Fig. 5e),
|
| 246 |
+
and TN (Fig. 5g) all had segmented linear relationships where smaller systems had steeper slopes than larger
|
| 247 |
+
systems (Table S4). Te breakpoint in surface area was 1.0 ha for NEP, 1.2 ha for R, and 3.8 ha for TN (Table S3).
|
| 248 |
+
For depth, two variables had logistic relationships: diel temperature range (Fig. 6e) and chlorophyll a (Fig. 6f),
|
| 249 |
+
with the infection occurring at 5.9 m and 14.9 m, respectively (Table S5). pH (Fig. 6b), TP (Fig. 6c), and TN
|
| 250 |
+
(Fig. 6d) all had segmented linear relationships where smaller systems had steeper slopes than larger systems
|
| 251 |
+
(Table S6) with breakpoints occurring at 1.0, 2.1, and 5.2 m, respectively (Table S5). For emergent vegetation
|
| 252 |
+
cover, TN (Fig. 7b), TP (Fig. 7c), and pH (Fig. 7d) all had segmented linear relationships where systems with
|
| 253 |
+
more emergent vegetation had steeper slopes than more open systems (Table S8). Te breakpoint in emergent
|
| 254 |
+
vegetation cover was 6.0% for TN, 8.2% for TP, and 26.0% for pH (Table S7).
|
| 255 |
+
To summarize across all three metrics (surface area, depth, and emergent vegetation cover), we evaluated
|
| 256 |
+
where the boundaries of nonlinear relationships generally occurred, which informs boundaries between ponds,
|
| 257 |
+
lakes, and wetlands (Table 1). For surface area, the boundary was 3.7±1.8 ha (mean±standard error) and the
|
| 258 |
+
median was 1.5 ha, consistent with the median of 2 ha from scientifc defnitions (Fig. 2b). Te depth boundary
|
| 259 |
+
was 5.8 ± 2.5 m (mean ± standard error) and the median was 5.2 m, within the range of scientifc defnitions
|
| 260 |
+
(Fig. 2c). Te emergent vegetation cover boundary was 13.4±6.3% (mean±standard error) and the median was
|
| 261 |
+
8.2%, both of which were lower than the previously identifed wetland lower bound of 30%31.
|
| 262 |
+
Figure 5. Relationships between lentic waterbody size (excluding wetlands) and ecosystem structure and
|
| 263 |
+
function metrics: (a) gross primary production (GPP), (b) total phosphorus concentrations (TP), (c) net
|
| 264 |
+
ecosystem production (NEP), (d) methane fuxes (CH4 fux), (e) respiration (R), (f) chlorophyll a concentrations
|
| 265 |
+
(Chl a), (g) total nitrogen concentrations (TN), (h) diel temperature ranges (DTR), and (i) gas transfer piston
|
| 266 |
+
velocity (k600). Optimal model fts from null, linear, segmented, and logistic curves in bold foreground lines. For
|
| 267 |
+
nonlinear segmented and logistic models (b–i), plots are ordered by boundaries between ponds and lakes, as
|
| 268 |
+
defned by model breakpoints or infection points (vertical background lines).
|
| 269 |
+
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|
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+
Pond morphology (e.g., size and depth) creates fundamentally distinct conditions that govern ecosystem
|
| 274 |
+
structure and function. Specifcally, ponds experience less wind-driven turbulence than larger waterbodies due
|
| 275 |
+
to small fetch and sheltering from the landscape36. We found that gas exchange rates (k600) decreased at~18 ha,
|
| 276 |
+
presumably due to reduced wind shear (Fig. 5i; also supported by37) and altered thermal dynamics. For instance,
|
| 277 |
+
ponds and shallow lakes can warm dramatically during the day, inducing stratifcation, and cool of and mix completely overnight38. We found higher diel temperature ranges were more common in waterbodies<5 ha (Fig. 5h)
|
| 278 |
+
and<6 m (Fig. 6e; see also39). Such diferences in temperature and mixing can promote internal nutrient loading40
|
| 279 |
+
and ecosystem respiration41, which may explain the higher TN (Figs. 4b, 5g), TP (Figs. 4a, 5b) and ecosystem
|
| 280 |
+
respiration (Fig. 5e) found in ponds. Lastly, diferences in water column mixing, increased nutrients, and higher
|
| 281 |
+
respiration can all contribute to the higher greenhouse gas emissions found in ponds relative to lakes (Fig. 5d)4,42.
|
| 282 |
+
Metrics of phytoplankton biomass (chl a) and total ecosystem production in the water (GPP) exhibited weak
|
| 283 |
+
or inconsistent relationships with surface area and depth, likely due to diferences in the location and types of
|
| 284 |
+
primary production across waterbody types. While total primary production in deep lakes is ofen dominated
|
| 285 |
+
by phytoplankton43, shallow waterbodies can shif toward non-planktonic primary production like benthic algae
|
| 286 |
+
or foating, emergent, or submerged macrophytes44. Ponds have pelagic phytoplankton, benthic algae (i.e., periphyton), and sediment rooted-submerged or foating macrophytes. In contrast, wetland productivity ofen predominantly occurs above the air–water interface45. Where emergent vegetation dominates, they may limit light
|
| 287 |
+
and reduce water column nutrients, both of which are needed by phytoplankton and periphyton. Macrophytes
|
| 288 |
+
can also modify water column and sediment geochemistry by providing autotrophic organic carbon and oxygen
|
| 289 |
+
to rooting systems in the sediments46. Consequently, these opposing drivers can explain the high variability in
|
| 290 |
+
primary production we observed (Fig. 5f, Table S2). Distinguishing ponds from wetlands will ultimately be aided
|
| 291 |
+
by additional ecosystem measurements of metabolism, greenhouse gas production, and additional metrics (e.g.,
|
| 292 |
+
carbon burial) across shallow waterbodies with a range of emergent vegetation cover.
|
| 293 |
+
Figure 6. Relationships between lentic waterbody maximum depth (Max depth) and various ecosystem
|
| 294 |
+
structure and function metrics: (a) methane fuxes (CH4 fux), (b) pH, (c) total phosphorus concentrations (TP),
|
| 295 |
+
(d) total nitrogen concentrations (TN), (e) diel temperature ranges (DTR), and (f) chlorophyll a concentrations
|
| 296 |
+
(Chl a) from literature data extraction with optimal model fts from null, linear or null, segmented linear, and
|
| 297 |
+
logistic curves in bold foreground lines. For nonlinear segmented and logistic models (b–f), plots are ordered by
|
| 298 |
+
model breakpoints or infection points (vertical background lines), indicative of boundaries between ponds and
|
| 299 |
+
lakes.
|
| 300 |
+
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|
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+
A functional pond defnition. Our review of existing pond defnitions highlights that surface area and
|
| 305 |
+
depth are the most common variables used to defne ponds; yet how small and how shallow a waterbody must
|
| 306 |
+
be to classify as a pond is unclear, with defnitions ranging by orders of magnitude. Emergent vegetation is a
|
| 307 |
+
third variable useful in distinguishing wetlands from ponds, but the threshold value,>30% emergent vegetation
|
| 308 |
+
coverage for wetlands established at the US federal level, is not based on documented changes in ecosystem function. Comparing characteristics among waterbodies that scientists self-categorized into lakes, ponds, or wetlands, ponds were sometimes distinct from lakes and wetlands (pH, TN), sometimes similar to wetlands (TP),
|
| 309 |
+
and sometimes similar to lakes (chl a), suggesting ponds are an ecologically distinct type of ecosystem. Lastly, we
|
| 310 |
+
Figure 7. Relationships between lentic waterbody emergent vegetation cover (Emergent veg.) and various
|
| 311 |
+
ecosystem structure and function metrics: (a) chlorophyll a concentrations (Chl a), (b) total nitrogen
|
| 312 |
+
concentrations (TN), (c) total phosphorus concentrations (TP), (d) pH from literature data extraction with
|
| 313 |
+
optimal model fts from null, linear or null, segmented linear, and logistic curves in bold foreground lines. For
|
| 314 |
+
nonlinear segmented and logistic models (b–d), plots are ordered by model breakpoints or infection points
|
| 315 |
+
(vertical background lines), indicative of boundaries between ponds and wetlands.
|
| 316 |
+
Table 1. Nonlinear boundary values, parameter estimate±standard error (SE), from comparisons between
|
| 317 |
+
surface area, maximum (max.) depth, and emergent vegetation (veg.) cover and ecosystem structure/function
|
| 318 |
+
metrics including gross primary production (GPP), total phosphorus concentrations (TP), methane fuxes
|
| 319 |
+
(CH4 fux), respiration (R), net ecosystem production (NEP), chlorophyll a concentrations (Chl a), pH, total
|
| 320 |
+
nitrogen concentrations (TN), diel temperature ranges (DTR), and gas transfer piston velocity (k600). Boundary
|
| 321 |
+
estimates are included if the nonlinear models (segmented regression or logistic relationships) were selected as
|
| 322 |
+
optimal fts with standard error as determined when ftting the parameter. NA indicates a null or linear ft, –
|
| 323 |
+
indicates not enough data was available to perform the analysis.
|
| 324 |
+
Ecosystem metric
|
| 325 |
+
Surface area
|
| 326 |
+
Boundary est.±SE (ha)
|
| 327 |
+
Max. depth
|
| 328 |
+
Boundary est.±SE (m)
|
| 329 |
+
Emergent veg. cover
|
| 330 |
+
Boundary est.±SE (%)
|
| 331 |
+
GPP NA – –
|
| 332 |
+
TP 0.8±1.2 2.1±1.2 8.2±1.2
|
| 333 |
+
NEP 1.0±1.4 – –
|
| 334 |
+
CH4 fux 1.1±1.7 NA –
|
| 335 |
+
R 1.2±1.5 – –
|
| 336 |
+
Chl a 1.5±1.7 14.9±1.2 NA
|
| 337 |
+
pH 1.7±1.5 1.0±1.4 26.0±1.3
|
| 338 |
+
TN 3.8±1.4 5.2±1.4 6.0±1.3
|
| 339 |
+
DTR 4.6±1.3 5.9±1.3 –
|
| 340 |
+
k600 17.5±1.5 – –
|
| 341 |
+
Mean 3.7±1.8 5.8±2.5 13.4±6.3
|
| 342 |
+
Median 1.5 5.2 8.2
|
| 343 |
+
9
|
| 344 |
+
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| 345 |
+
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| 346 |
+
www.nature.com/scientificreports/
|
| 347 |
+
found clear nonlinear relationships when we examined relationships between ecosystem structure or function
|
| 348 |
+
and surface area, depth, and emergent vegetation cover; these boundaries help to quantitatively defne ponds.
|
| 349 |
+
Specifcally, we found that across available ecosystem metrics, ecosystems shif in structure and function at
|
| 350 |
+
average (±SE) values of 3.7 (±1.8) ha in size, 5.8 (±2.5) m in depth, and 13.4 (±6.3) % emergent vegetation cover
|
| 351 |
+
(Table 1). For surface area, all but one ecosystem metric (k600) was below 5 ha in surface area, which fts well
|
| 352 |
+
within the range of most existing defnitions (≤10 ha; Fig. 2), and we suggest may be used to distinguish ponds
|
| 353 |
+
from lakes. For maximum depth, all but one ecosystem metric (chl. a) was below a 6 m depth threshold, which
|
| 354 |
+
also fts well within the range of depths reported in pond defnitions (Fig. 2), and matches the published threshold of 5 m maximum depth for shallow lakes44. Our depth analysis was less robust than surface area because we
|
| 355 |
+
had less depth data, a common challenge in lentic studies47; we therefore advise further studies in waterbodies
|
| 356 |
+
to explicitly evaluate this threshold. Until further work is done, we recommend using 5 m as a maximum depth
|
| 357 |
+
threshold for ponds as it is close to both threshold shifs in ecosystem function and matches with the shallow
|
| 358 |
+
lake literature44,48. We had the fewest ecosystem metric comparisons for emergent vegetative cover, and observed
|
| 359 |
+
three nonlinear boundaries ranging from 6 to 26% cover. Te mean (13.4%), though smaller, is not statistically
|
| 360 |
+
diferent than the 30% emergent vegetation cover (one sample t-test, t2=− 2.6, p=0.12) proposed by Cowardin
|
| 361 |
+
et al.31 to separate wetlands from lakes. We recommend separating ponds and wetlands using the 30% coverage
|
| 362 |
+
in emergent vegetation threshold for now, but recognize that the Cowardin et al.31 metric is not data driven and
|
| 363 |
+
our analysis was limited by existing data. Future studies must examine how ecosystem structure and function
|
| 364 |
+
shifs across a gradient of emergent vegetation cover to better functionally distinguish wetlands from ponds and
|
| 365 |
+
could ultimately lower that boundary.
|
| 366 |
+
Our review of data from the literature showed scientists and managers view ponds as permanent or temporary
|
| 367 |
+
and natural or human made in origin. Terefore, we felt it necessary to provide the inclusivity of these concepts
|
| 368 |
+
in a pond defnition. Other defnitions also link depth to light availability, where light penetrates to the sediments
|
| 369 |
+
across the pond (e.g.,11). However, light availability is not only mediated by depth; even in the shallowest systems
|
| 370 |
+
light can be limiting due to turbidity, dissolved organic matter, and submerged or foating plants (e.g.,49,50). For
|
| 371 |
+
example, foating duckweed can cover most of a pond’s surface area and reduce light penetration to<1% relative
|
| 372 |
+
to the light above the water’s surface49, and dramatically change the ecology of shallow systems51.
|
| 373 |
+
As our analyses indicate that ponds are functionally distinct from lakes and wetlands, we propose the following scientifcally informed pond defnition (Fig. 8):
|
| 374 |
+
Ponds are small and shallow waterbodies with a maximum surface area of 5 ha, a maximum depth of 5 m,
|
| 375 |
+
and < 30% coverage of emergent vegetation. Ponds will have light penetration to the sediments if water
|
| 376 |
+
clarity permits and can be permanent or temporary and natural or human-made.
|
| 377 |
+
Our proposed defnition is based on the current state of the science; we anticipate that future research will
|
| 378 |
+
further resolve diferences among these fve categories. For example, we call for future research to examine how
|
| 379 |
+
ecosystem structure and function shif across our proposed boundaries, particularly for depth and emergent
|
| 380 |
+
Figure 8. Conceptual model to defne lentic waterbodies based on three diferent criteria (depth, surface area,
|
| 381 |
+
and emergent vegetation). Boundaries for all three axes come from our analysis and are informed by existing
|
| 382 |
+
pond, lake, and wetland defnitions. Figure by Visualizing Science.
|
| 383 |
+
10
|
| 384 |
+
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|
| 385 |
+
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|
| 386 |
+
www.nature.com/scientificreports/
|
| 387 |
+
vegetation, which had smaller sample sizes and fewer ecosystem metrics than surface area. Additional variables
|
| 388 |
+
such as basin geometry (e.g., area:volume ratios), sheltering from wind, water residence time, water clarity, and
|
| 389 |
+
geographic location, may also afect a waterbody’s ecosystem structure and function, creating some overlap
|
| 390 |
+
between classifcations especially along the upper and lower bounds of our pond defnition. For instance, a
|
| 391 |
+
landscape with little wind sheltering increases water column mixing that could cause a waterbody the size of a
|
| 392 |
+
pond to function more like a shallow lake. We therefore advocate for additional sampling of lentic waterbodies,
|
| 393 |
+
especially in locations where lentic waterbodies are understudied or being rapidly constructed like tropical and
|
| 394 |
+
subtropical regions52, to help resolve boundaries among waterbody types and further refne the pond defnition.
|
| 395 |
+
Conclusion
|
| 396 |
+
Scientists, policy makers, water resource managers and the public all use the word “pond” to describe small and
|
| 397 |
+
shallow waterbodies, which are globally abundant8
|
| 398 |
+
and hotspots for biogeochemistry4,8
|
| 399 |
+
and biodiversity53. Yet,
|
| 400 |
+
the lack of a universal pond defnition means that ponds can fall between lake and wetland jurisdictions and
|
| 401 |
+
categorizations22, thus potentially limiting their legal protections. Globally, the situation is similar to US policy
|
| 402 |
+
as some nations defne ponds as wetlands (e.g., the Ramsar Convention), some as lakes (e.g., Denmark), and
|
| 403 |
+
others specifcally defne ponds (e.g., United Kingdom). Te pond defnition presented here will favor more
|
| 404 |
+
frequent and consistent use of the term and ultimately improve the protection, monitoring, and scientifc study
|
| 405 |
+
of ponds, which are globally abundant and structurally and functionally distinct from other lentic waterbodies.
|
| 406 |
+
Methods
|
| 407 |
+
Literature survey. To compile biological, physical, and chemical characteristics of ponds, we conducted a
|
| 408 |
+
literature search based on three seminal papers establishing the ecological importance of ponds: Oertli et al.35,
|
| 409 |
+
Søndergaard et al.18, and Downing8
|
| 410 |
+
, each of which has>100 citations and is more than ten years old. We conducted a backwards and forwards search in April 2019 to compile all papers cited by these three papers, and
|
| 411 |
+
all papers that cited them, yielding 519 unique papers. We extracted physical, chemical, and biological data for
|
| 412 |
+
papers that reported data for individual waterbodies defned as ponds by the publishing scientists. To ensure
|
| 413 |
+
consideration of all potential ponds, we checked that waterbodies selected were small (≤20 ha in surface area)
|
| 414 |
+
and shallow (≤9 m in maximum or mean depth), boundaries that are slightly greater than the maximum depth
|
| 415 |
+
(8 m)35 and maximum surface area (10 ha)34 used to defne ponds in a few prior studies. We used the resulting
|
| 416 |
+
1327 waterbodies in our analysis, which had a global distribution (Fig. S1)19.
|
| 417 |
+
Scientifc defnitions. To investigate how scientifc researchers defned ponds, we reviewed all 519 papers
|
| 418 |
+
for pond defnitions. We included defnitions where the authors explicitly referred to their study waterbodies
|
| 419 |
+
as ponds (e.g., we excluded “shallow lakes” and “small lakes”), yielding 40 pond defnitions. Te defnitions
|
| 420 |
+
included 89 citations of 48 unique papers; we evaluated all cited papers that were not already in our compilation
|
| 421 |
+
for additional defnitions and citations. Tis process added 14 defnitions, plus an additional fve cited papers not
|
| 422 |
+
assessed due to our inability to access or translate them (data available19).
|
| 423 |
+
Federal and state defnitions. We examined policy defnitions using the United States (US) federal and
|
| 424 |
+
state legislation as an example because we posited diferences at this scale would refect the challenges faced by
|
| 425 |
+
governments from other countries in formulating a unifed pond defnition. At the federal level, we examined
|
| 426 |
+
three agencies with monitoring or regulatory responsibilities: US Environmental Protection Agency (EPA), US
|
| 427 |
+
Army Corps of Engineers (ACE), and US Fish and Wildlife Service (USFWS). Due to the difculty of fnding all
|
| 428 |
+
state policies, we sent electronic surveys to individuals working in state environmental agencies in all states. We
|
| 429 |
+
asked whether their state defned lakes, ponds, and wetlands, and requested the legislative sources. We received
|
| 430 |
+
responses from 42/50 states and evaluated all defnitions provided and their associated legislation.
|
| 431 |
+
Pond, lake, and wetland data. We compared chemical and biological characteristics among various
|
| 432 |
+
lentic waterbodies as defned by scientists as ponds, wetlands, or lakes. For ponds, we used data from the literature data extraction as described above (n=1327). Wetland data came from the US EPA’s 2011 National
|
| 433 |
+
Wetland Condition Assessment, which surveyed wetlands with standing water<1 m in depth and variable surface area54,55. We selected wetland sites that were freshwater and had water chemistry data (n=400). Lake data
|
| 434 |
+
was extracted from LAGOS-NE (lakes≥4 ha; n=51,101)56, EPA’s 2012 National Lake Assessment (lakes≥1 ha;
|
| 435 |
+
n=1130)57,58, and the European Environmental Agency’s Waterbase database (lakes>0.02 ha; n=2942)59. From
|
| 436 |
+
these sources, we compared nutrients (total phosphorus (TP), total nitrogen (TN)), water chemistry (pH), and
|
| 437 |
+
primary producer biomass (chl a) among waterbody types (ponds, wetlands, and lakes).
|
| 438 |
+
We also examined diferences in six additional metrics of ecosystem function across waterbodies using data
|
| 439 |
+
from a variety of sources and ranging in sample size from 67 to 198 global sites (gross primary production—GPP,
|
| 440 |
+
respiration—R, net ecosystem production—NEP, diel temperature ranges—DTR, methane fuxes, gas transfer
|
| 441 |
+
velocities). We extracted metabolism metrics (GPP and R) from an existing literature review60 and two published
|
| 442 |
+
studies of various sized lentic ecosystems41,61. DTR, calculated as the diel diference between the maximum and
|
| 443 |
+
minimum surface temperature for each waterbody, were extracted from multiple studies38,39. Areal methane
|
| 444 |
+
fuxes42 and gas transfer velocities (k600)37 were extracted from existing literature reviews.
|
| 445 |
+
Comparing lake, pond, and wetlands characteristics from literature. We evaluated whether there
|
| 446 |
+
were diferences among waterbody types as defned by scientists and managers. We determined signifcant differences in waterbody characteristics across waterbody types using ANOVA and post-hoc Least Signifcant
|
| 447 |
+
11
|
| 448 |
+
Vol.:(0123456789)
|
| 449 |
+
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|
| 450 |
+
www.nature.com/scientificreports/
|
| 451 |
+
Diference (LSD) analysis. We determined the variation within each freshwater type using the coefcient of
|
| 452 |
+
variation (cv) and tested for signifcant diferences using Levene’s test. We acknowledge that the confounding
|
| 453 |
+
defnitions of waterbodies resulted in overlapping size distributions; therefore any statistical diferences across
|
| 454 |
+
waterbody types will be conservative.
|
| 455 |
+
Does pond ecosystem structure and function distinguish ponds difer from lakes and wet- lands? We evaluated where the cutofs might exist along pond to lake and pond to wetland gradients. We
|
| 456 |
+
used the relationship between surface area, depth, or emergent vegetation cover represented by x below and
|
| 457 |
+
each ecosystem variable (n=10 variables for surface area; n=6 variables for depth, n=4 for emergent vegetation cover) represented by y below for four diferent patterns in increasing order of complexity: null, linear,
|
| 458 |
+
segmented (nonlinear), and logistic (nonlinear) patterns. We ft null models by taking the arithmetic mean
|
| 459 |
+
(Eq. 1), linear models using ordinary least-squares linear regression (Eq. 2), segmented using regressions with
|
| 460 |
+
one breakpoint (Eq. 3) via the “segmented” package62, and logistic using sigmoidal curves (Eq. 4) via the nls
|
| 461 |
+
function in R (Fig. S2). Parameters a – h and bp (breakpoint) were ft using the methods above.
|
| 462 |
+
We log-transformed surface area and depth to account for the several order of magnitude scale and nonnormality. Similarly, we transformed some of the ecosystem variables depending on normality and distributions.
|
| 463 |
+
To select among the four models for each relationship, we examined the AICc fts and selected the minimum
|
| 464 |
+
AICc as the optimal ft with consideration of other model fts within 11 units of the minimum AICc using root
|
| 465 |
+
mean squared error and parsimony63. If one of the nonlinear models was selected, we then objectively quantifed
|
| 466 |
+
the boundary among ecosystem types (i.e., pond vs. lake or pond vs. wetland) using either the breakpoint or the
|
| 467 |
+
infection point parameter from the segmented regression or sigmoid curve, respectively (Fig. S2c,d).
|
| 468 |
+
Data availability
|
| 469 |
+
All data used for this manuscript is available through an Environmental Data Initiative data publication (Richardson et al. 2022: https://doi.org/10.6073/pasta/ec507ac70846b17d0633d95aa3c680c6).
|
| 470 |
+
Received: 7 January 2022; Accepted: 8 June 2022
|
| 471 |
+
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|
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(1) y = a
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y = bx + c (2)
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y = f + (4) g − f
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1 + e(bp−x)/h
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Acknowledgements
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Tis work was supported by the Global Lake Ecological Observatory Network (GLEON). We thank participants
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in the GLEON 19 pond ad hoc meeting for discussions that formed the basis for this project. MRA was supported
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as part of the BEYOND 2020 project (grant-aid agreement no. PBA/FS/16/02) by the Marine Institute and funded
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under the Marine Research Programme by the Irish Government. Funding to KSC and KBSK was from the
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US National Science Foundation (EF-1638679 and EF-1638539). KKH was supported by the Adam S. Tomas
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Endowment and by the St. Olaf Collaborative Undergraduate Research and Inquiry Program. Tis material is
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13
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Vol.:(0123456789)
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Scientifc Reports | (2022) 12:10472 | https://doi.org/10.1038/s41598-022-14569-0
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www.nature.com/scientificreports/
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based upon work supported by the National Science Foundation Graduate Research Fellowship Program under
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| 595 |
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Grant No. 2036201 to MJF (Farruggia). Any opinions, fndings, and conclusions or recommendations expressed
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in this material are those of the authors and do not necessarily refect the views of the National Science Foundation. Credit for Fig. 8 to Fiona Martin at Visualizing Science (https://www.visualizingscience.com/).
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Author contributions
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D.C.R. and M.A.H. led the project. D.C.R., M.A.H., M.J.F., K.K.H., and K.B.S.K. organized the literature searching, survey, data analysis, fgure preparation, and wrote the manuscript. All authors completed data mining from
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literature survey and were substantively involved in two of the following three categories: (1) project conceptual development, (2) data extraction/analysis/interpretation, and (3) writing/revising manuscript. All authors
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approved the fnal version of this manuscript prior to submission.
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Competing interests
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Te authors declare no competing interests.
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Additional information
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+
Supplementary Information Te online version contains supplementary material available at https://doi.org/
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+
10.1038/s41598-022-14569-0.
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+
Correspondence and requests for materials should be addressed to D.C.R.
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+
Reprints and permissions information is available at www.nature.com/reprints.
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Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and
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institutional afliations.
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Open Access Tis article is licensed under a Creative Commons Attribution 4.0 International
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License, which permits use, sharing, adaptation, distribution and reproduction in any medium or
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format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the
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Creative Commons licence, and indicate if changes were made. Te images or other third party material in this
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article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the
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material. If material is not included in the article’s Creative Commons licence and your intended use is not
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permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from
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the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
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© Te Author(s) 2022
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"""
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