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==Evolution== The origins and evolutionary relationships between the three main groups of amphibians are hotly debated. A [[molecular phylogeny]] based on [[Ribosomal DNA|rDNA]] analysis dating from 2005 suggests that [[salamanders]] and [[caecilian]]s are more closely related to each other than they are to frogs and the [[Genetic divergence|divergence]] of the three groups took place in the [[Paleozoic]] or early [[Mesozoic]] before the break-up of the supercontinent [[Pangaea]] and soon after their divergence from the [[Sarcopterygii|lobe-finned fishes]]. This would help account for the relative scarcity of amphibian fossils from the period before the groups split.<ref>{{cite journal |author1=San Mauro, Diego |author2=Vences, Miguel |author3=Alcobendas, Marina |author4=Zardoya, Rafael |author5=Meyer, Axel |year=2005 |title=Initial Diversification of living amphibians predated the breakup of Pangaea |journal=The American Naturalist |volume=165 |issue=5 |pages=590–599 |jstor=429523 |doi=10.1086/429523 |pmid=15795855 |bibcode=2005ANat..165..590S |s2cid=17021360 |url=http://nbn-resolving.de/urn:nbn:de:bsz:352-opus-33053 }}{{subscription required}}</ref> Another molecular phylogenetic analysis conducted about the same time concluded that [[lissamphibian]]s first appeared about 330 million years ago and that the [[Temnospondyli|temnospondyl-origin]] hypothesis is more credible than other theories. The [[neobatrachia]]ns seemed to have originated in Africa/India, the salamanders in East Asia and the caecilians in tropical Pangaea.<ref>{{cite journal |author1=Zhang, Peng |author2=Zhou, Hui |author3=Chen, Yue-Qin |author4=Liu, Yi-Fei |author5=Qu, Liang-Hu |year=2005 |title=Mitogenomic perspectives on the origin and phylogeny of living amphibians |journal=Systematic Biology |volume=54 |issue=3 |pages=391–400 |doi=10.1080/10635150590945278 |pmid=16012106 |doi-access=free }}</ref> Other researchers, while agreeing with the main thrust of this study, questioned the choice of calibration points used to synchronise the data. They proposed that the date of lissamphibian diversification should be placed in the [[Permian]], rather less than 300 million years ago, a date in better agreement with the palaeontological data.<ref>{{cite journal |author1=Marjanović, David |author2=Laurin, Michel |year=2007 |title=Fossils, molecules, divergence times, and the origin of lissamphibians |journal=Systematic Biology |volume=56 |issue=3 |pages=369–388 |doi=10.1080/10635150701397635 |pmid=17520502|doi-access=free }}</ref> A further study in 2011 using both extinct and living taxa sampled for morphological, as well as molecular data, came to the conclusion that Lissamphibia is [[Monophyly|monophyletic]] and that it should be nested within [[Lepospondyli]] rather than within [[Temnospondyli]]. The study postulated that Lissamphibia originated no earlier than the late [[Carboniferous]], some 290 to 305 million years ago. The split between Anura and [[Caudata]] was estimated as taking place 292 million years ago, rather later than most molecular studies suggest, with the caecilians splitting off 239 million years ago.<ref>{{cite journal |author=Pyron, R. Alexander |year=2011 |title=Divergence time estimation using fossils as terminal taxa and the origins of Lissamphibia |journal=Systematic Biology |volume=60 |issue=4 |pages=466–481 |doi=10.1093/sysbio/syr047 |pmid=21540408 |url=https://zenodo.org/record/889970 |doi-access=free }}</ref> [[File:Fossilised frog.jpg|thumb|A fossilised frog from the [[Czech Republic]], possibly ''[[Palaeobatrachus gigas]]'']] In 2008, ''[[Gerobatrachus hottoni]]'', a [[Temnospondyli|temnospondyl]] with many frog- and salamander-like characteristics, was discovered in [[Texas]]. It dated back 290 million years and was hailed as a [[Transitional fossil|missing link]], a [[Stem group|stem]] batrachian close to the [[common ancestor]] of frogs and salamanders, consistent with the widely accepted hypothesis that frogs and salamanders are more closely related to each other (forming a [[clade]] called Batrachia) than they are to caecilians.<ref>{{cite news |title="Frog-amander" fossil may be amphibian missing link |author=Casselman, Anne |url=http://news.nationalgeographic.com/news/2008/05/080521-frog-fossil.html |archive-url=https://web.archive.org/web/20080525210713/http://news.nationalgeographic.com/news/2008/05/080521-frog-fossil.html |url-status=dead |archive-date=May 25, 2008 |newspaper=National Geographic News |date=May 21, 2008 |access-date=July 5, 2012}}</ref><ref>{{cite journal |author1=Anderson, Jason S. |author2=Reisz, Robert R. |author3=Scott, Diane |author4=Fröbisch, Nadia B. |author5=Sumida, Stuart S. |year=2008 |title=A stem batrachian from the Early Permian of Texas and the origin of frogs and salamanders |journal=Nature |volume=453 |pages=515–518 |doi=10.1038/nature06865 |issue=7194 |pmid=18497824 |bibcode=2008Natur.453..515A |s2cid=205212809 }}</ref> However, others have suggested that ''Gerobatrachus hottoni'' was only a [[Dissorophoidea|dissorophoid]] temnospondyl unrelated to extant amphibians.<ref name="M&L09">{{cite journal |last1=Marjanović |first1=D. |last2=Laurin |first2=M. |year=2009 |title=The origin(s) of modern amphibians: a commentary |journal=Evolutionary Biology |volume=36 |issue=3 |pages=336–338 |doi=10.1007/s11692-009-9065-8 |bibcode=2009EvBio..36..336M |s2cid=12023942 |url=https://hal.archives-ouvertes.fr/hal-00549002/file/MARJANOVIC_David.pdf }}</ref> [[Salientia]] (Latin ''salire'' (''salio''), "to jump") is the name of the total group that includes modern frogs in the order Anura as well as their close fossil relatives, the "proto-frogs" or "stem-frogs". The common features possessed by these proto-frogs include 14 [[Vertebral column|presacral vertebrae]] (modern frogs have eight or 9), a long and forward-sloping [[Ilium (bone)|ilium]] in the [[pelvis]], the presence of a [[Parietal bone|frontoparietal bone]], and a [[Mandible|lower jaw]] without teeth. The earliest known amphibians that were more closely related to frogs than to salamanders are ''[[Triadobatrachus massinoti]]'', from the early [[Triassic]] period of [[Madagascar]] (about 250 million years ago), and ''[[Czatkobatrachus polonicus]]'', from the Early Triassic of [[Poland]] (about the same age as ''Triadobatrachus'').<ref name=TOLweb>{{cite web |last=Cannatella |first=David |title=''Triadobatrachus massinoti'' |url=http://www.tolweb.org/Triadobatrachus_massinoti/16962 |work=Tree of Life |year=1995 |access-date=June 26, 2008 }}</ref> The skull of ''Triadobatrachus'' is frog-like, being broad with large eye sockets, but the fossil has features diverging from modern frogs. These include a longer body with more [[vertebrae]]. The tail has separate vertebrae unlike the fused urostyle or coccyx in modern frogs. The tibia and fibula bones are also separate, making it probable that ''Triadobatrachus'' was not an efficient leaper.<ref name=TOLweb/> A 2019 study has noted the presence of Salientia from the [[Chinle Formation]], and suggested that anurans might have first appeared during the [[Late Triassic]].<ref>{{cite journal |author1=Michelle R. Stocker |author2=Sterling J. Nesbitt |author3=Ben T. Kligman |author4=Daniel J. Paluh |author5=Adam D. Marsh |author6=David C. Blackburn |author7=William G. Parker |year=2019 |title=The earliest equatorial record of frogs from the Late Triassic of Arizona |journal=Biology Letters |volume=15 |issue=2 |pages=Article ID 20180922 |doi=10.1098/rsbl.2018.0922 |pmid=30958136 |pmc=6405462 |hdl=10919/87931 }}</ref> On the basis of fossil evidence, the earliest known "true frogs" that fall into the anuran lineage proper all lived in the early [[Jurassic]] period.<ref name=Salientia>{{cite web |url=http://tolweb.org/Salientia/ |title=Salientia |author=Cannatella, David C.|year=1997 |work=Tree of Life Web Project |access-date=August 7, 2012}}</ref><ref name="Heatwole00">{{cite book |title=Amphibian Biology: Paleontology: The Evolutionary History of Amphibians |editor1-last=Heatwole |editor1-first=H. |editor2-last=Carroll |editor2-first=R. L. |year=2000 |volume=4 |publisher=Surrey Beatty & Sons |isbn=978-0-949324-87-0 |chapter=14. Mesozoic Amphibians |last1=Roček |first1=Z. |pages=1295–1331 |chapter-url=http://rocek.gli.cas.cz/Reprints/AmphBiol3.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://rocek.gli.cas.cz/Reprints/AmphBiol3.pdf |archive-date=October 9, 2022 |url-status=live}}</ref> One such early frog species, ''[[Prosalirus bitis]]'', was discovered in [[1995 in paleontology|1995]] in the [[Kayenta Formation]] of [[Arizona]] and dates back to the [[Early Jurassic]] epoch (199.6 to 175 million years ago), making ''Prosalirus'' somewhat more recent than ''Triadobatrachus''.<ref name="jurassicdistribution">{{cite book |title=Dinosaur distribution (Early Jurassic, North America): The Dinosauria |editor1-last=Weishampel |editor1-first=D. B. |editor2-last=Dodson |editor2-first=P. |editor3-last=Osmólska |editor3-first=H. |year=2004 |publisher=University of California Press |isbn=978-0-520-24209-8 |pages=530–532 |edition=2nd }}</ref> Like the latter, ''Prosalirus'' did not have greatly enlarged legs, but had the typical three-pronged [[pelvic]] structure of modern frogs. Unlike ''Triadobatrachus'', ''Prosalirus'' had already lost nearly all of its tail<ref>{{cite journal |journal=Nature |volume=377 |pages=49–52 |year=1995 |doi=10.1038/377049a0 |title=An Early Jurassic jumping frog |author1=Shubin, N. H. |author2=Jenkins, F. A. Jr |issue=6544 |bibcode=1995Natur.377...49S |s2cid=4308225 }}</ref> and was well adapted for jumping.<ref name="foster-anura">{{cite book |last1=Foster |first1=J. |year=2007 |chapter=Anura (Frogs) |title=Jurassic west: the dinosaurs of the Morrison Formation and their world |publisher=Indiana University Press |pages=135–136 |isbn=978-0-253-34870-8 }}</ref> Another Early Jurassic frog is ''[[Vieraella herbsti]]'', which is known only from [[dorsum (biology)|dorsal]] and [[ventral]] impressions of a single animal and was estimated to be {{convert|33|mm|in|frac=8|abbr=on}} from snout to vent. ''[[Notobatrachus degiustoi]]'' from the [[middle Jurassic]] is slightly younger, about 155–170 million years old. The main evolutionary changes in this species involved the shortening of the body and the loss of the tail. Tadpoles of ''N. degiustoi'' constitute the oldest tadpoles found as of 2024, dating back to 168–161 million years ago. These tadpoles also showed adaptations for [[filter-feeding]], implying residence in temporary pools by filter-feeding larvae was already commonplace.<ref>{{Cite journal |last1=Chuliver |first1=Mariana |last2=Agnolín |first2=Federico L. |last3=Scanferla |first3=Agustín |last4=Aranciaga Rolando |first4=Mauro |last5=Ezcurra |first5=Martín D. |last6=Novas |first6=Fernando E. |last7=Xu |first7=Xing |date=2024-10-30 |title=The oldest tadpole reveals evolutionary stability of the anuran life cycle |url=https://www.nature.com/articles/s41586-024-08055-y |journal=Nature |volume=636 |issue=8041 |language=en |pages=138–142 |doi=10.1038/s41586-024-08055-y |pmid=39478214 |bibcode=2024Natur.636..138C |issn=1476-4687}}</ref> The evolution of modern Anura likely was complete by the Jurassic period. Since then, evolutionary changes in [[Ploidy|chromosome numbers]] have taken place about 20 times faster in mammals than in frogs, which means [[speciation]] is occurring more rapidly in mammals.<ref>{{cite journal |author1=Wilson, A. C. |author2=Sarich, V. M. |author3=Maxson, L. R. |year=1974 |title=The importance of gene rearrangement in evolution: evidence from studies on rates of chromosomal, protein, and anatomical evolution |journal=Proceedings of the National Academy of Sciences |volume=71 |issue=8 |pages=3028–3030 |doi=10.1073/pnas.71.8.3028 |pmid=4528784 |bibcode=1974PNAS...71.3028W |pmc=388613 |doi-access=free }}</ref> According to genetic studies, the families [[Hyloidea]], [[Microhylidae]], and the clade [[Natatanura]] (comprising about 88% of living frogs) diversified simultaneously some 66 million years ago, soon after the [[Cretaceous–Paleogene extinction event]] associated with the [[Chicxulub impactor]]. All origins of arboreality (e.g. in Hyloidea and Natatanura) follow from that time and the resurgence of forest that occurred afterwards.<ref>{{cite news |title=Frog evolution linked to dinosaur asteroid strike |url=https://www.bbc.com/news/science-environment-40482039 |newspaper=BBC News |date=July 3, 2017 |access-date=July 3, 2017}}</ref><ref>{{cite journal |periodical=Proceedings of the National Academy of Sciences of the United States of America |title=Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous–Paleogene boundary |author1=Feng, Yan-Jie |author2=Blackburn, David C. |author3=Liang, Dan |author4=Hillis, David M. |author5=Wake, David B. |author6=Cannatella, David C. |author7=Zhang, Peng |year=2017 |doi=10.1073/pnas.1704632114 |pmc=5530686 |pmid=28673970 |volume=114 |issue=29 |pages=E5864–E5870 |bibcode=2017PNAS..114E5864F |doi-access=free}}</ref> Frog fossils have been found on all of the Earth's continents.<ref>{{cite journal |last1=Evans |first1=S. E. |last2=Jones |first2=M. E. H. |last3=Krause |first3=D. W. |title=A giant frog with South American affinities from the Late Cretaceous of Madagascar |journal=Proceedings of the National Academy of Sciences |volume=105 |pages=2951–2956 |year=2008 |doi=10.1073/pnas.0707599105 |pmid=18287076 |issue=8 |pmc=2268566 |bibcode=2008PNAS..105.2951E |doi-access=free }}</ref><ref>{{cite journal |last1=Mörs |first1=Thomas |last2=Reguero |first2=Marcelo |last3=Vasilyan |first3=Davit |title=First fossil frog from Antarctica: implications for Eocene high latitude climate conditions and Gondwanan cosmopolitanism of Australobatrachia |journal=Scientific Reports |date=2020 |volume=10 |issue=1 |page=5051 |doi=10.1038/s41598-020-61973-5 |pmid=32327670 |pmc=7181706 |bibcode=2020NatSR..10.5051M |doi-access=free}}</ref> In 2020, it was announced that 40 million year old [[Calyptocephalellidae|helmeted frog]] fossils had been discovered by a team of vertebrate palaeontologists in [[Seymour Island]] on the [[Antarctic Peninsula]], indicating that this region was once home to frogs related to those now living in South American [[Valdivian temperate rain forest|''Nothofagus'' forest]].<ref>{{Cite news |last=Joel |first=Lucas |date=April 23, 2020 |title=Fossil Shows Cold-Blooded Frogs Lived on Warm Antarctica |language=en-US |work=The New York Times|url=https://www.nytimes.com/2020/04/23/science/frog-antarctica-fossil.html |archive-url=https://web.archive.org/web/20200423151008/https://www.nytimes.com/2020/04/23/science/frog-antarctica-fossil.html |archive-date=April 23, 2020 |url-access=subscription |url-status=live |access-date=May 13, 2020 |issn=0362-4331}}</ref> ===Phylogeny=== A [[cladogram]] showing the relationships of the different [[Family (biology)|families]] of frogs in the clade Anura can be seen in the table below. This diagram, in the form of a [[Phylogenetic tree|tree]], shows how each frog family is related to other families, with each node representing a point of common ancestry. It is based on Frost ''et al.'' (2006),<ref>{{Cite journal | last1 = Frost | first1 = D. R. | last2 = Grant | first2 = T. | last3 = Faivovich | first3 = J. N. | last4 = Bain | first4 = R. H. | last5 = Haas | first5 = A. | last6 = Haddad | first6 = C. L. F. B. | last7 = De Sá | first7 = R. O. | last8 = Channing | first8 = A. | last9 = Wilkinson | first9 = M. | last10 = Donnellan | first10 = S. C. | last11 = Raxworthy | first11 = C. J. | last12 = Campbell | first12 = J. A. | last13 = Blotto | first13 = B. L. | last14 = Moler | first14 = P. | last15 = Drewes | first15 = R. C. | last16 = Nussbaum | first16 = R. A. | last17 = Lynch | first17 = J. D. | last18 = Green | first18 = D. M. | last19 = Wheeler | first19 = W. C. | title = The Amphibian Tree of Life | journal = Bulletin of the American Museum of Natural History | volume = 297 | pages = 1–291| year = 2006 | doi = 10.1206/0003-0090(2006)297[0001:TATOL]2.0.CO;2 | hdl = 2246/5781| s2cid = 86140137 | doi-access = free }}</ref> Heinicke ''et al.'' (2009)<ref>{{cite journal |author1=Heinicke M. P. |author2=Duellman, W. E. |author3=Trueb, L. |author-link3=Linda Trueb |author4=Means, D. B. |author5=MacCulloch, R. D. |author6=Hedges, S. B. | year = 2009 | title = A new frog family (Anura: Terrarana) from South America and an expanded direct-developing clade revealed by molecular phylogeny | url = http://www.mapress.com/zootaxa/2009/f/z02211p035f.pdf | journal= Zootaxa | volume = 2211 | pages = 1–35 |doi=10.11646/zootaxa.2211.1.1 }}</ref> and Pyron and Wiens (2011).<ref>{{cite journal |author1=R. Alexander Pyron |author2=John J. Wiens | year = 2011 | title= A large-scale phylogeny of Amphibia including over 2800 species, and a revised classification of extant frogs, salamanders, and caecilians | journal = Molecular Phylogenetics and Evolution | volume = 61 | issue = 2 | pages = 543–583 | doi=10.1016/j.ympev.2011.06.012| pmid = 21723399 | doi-access = free |bibcode=2011MolPE..61..543A }}</ref> {{clade| style=font-size:80%;line-height:100% |label1='''Anura''' |1={{clade |label1= |1={{clade |1=[[Leiopelmatidae]] |2=[[Ascaphidae]] }} |label2=[[Bombianura]] |2={{clade |label1=[[Costata]] |1={{clade |1=[[Bombinatoridae]] |label2= |2={{clade |1=[[Alytidae]] |2=[[Discoglossidae]] }} }} |label2=[[Pipanura]] |2={{clade |label1=[[Xenoanura]] |1={{clade |1=[[Pipidae]] |2=[[Rhinophrynidae]] }} |label2=[[Acosmanura]] |2={{clade |label1=[[Anomocoela]] |1={{clade |1=[[Scaphiopodidae]] |2={{clade |1=[[Pelodytidae]] |label2=[[Pelobatoidea]] |2={{clade |1=[[Pelobatidae]] |2=[[Megophryidae]] }} }} }} |label2=[[Neobatrachia]] |2=NEOBATRACHIA }} }} }} }} <!----- The main frog cladogram has three subtrees: NEOBATRACHIA, HYLOIDEA, RANO-------------------> |targetA=NEOBATRACHIA |subcladeA={{clade |1=[[Heleophrynidae]] |label2=[[Phthanobatrachia]] |2={{clade |1={{clade <!--dummy clade for reduced horizontal spacing--> |label1=[[Sooglossoidea]] |1={{clade |1=[[Sooglossidae]] |2=[[Nasikabatrachidae]] }} }} |label2= |2={{clade |label1=[[Notogaeanura]] |1={{clade |label1=[[Australobatrachia]] |1={{clade |1=[[Calyptocephalellidae]] |label2= |2={{clade |1=[[Myobatrachidae]] |2=[[Limnodynastidae]] }} }} |label2=[[Nobleoanura]] |2=HYLOIDEA }} |label2=[[Ranoidea|Ranoides]] |2=RANOIDEA }} }} }} <!-- end NEOBATRACHIA subtree--> |targetB=HYLOIDEA |subcladeB={{clade |1={{clade <!--dummy clade for reduced horizontal spacing--> |label1=[[Brachycephaloidea]] |sublabel1=([[Terrarana]]) |1={{clade |1=[[Ceuthomantidae]] |label2= |2={{clade |1=[[Brachycephalidae]] |label2= |2={{clade |1=[[Eleutherodactylidae]] |2=[[Craugastoridae]] }} }} }} }} |2={{clade |1=[[Hemiphractidae]] |label2=[[Athesphatanura]] |2={{clade |1=[[Hylidae]] |label2=[[Leptodactyliformes]] |2={{clade |label1= |1={{clade |1={{clade <!--dummy clade for reduced horizontal spacing--> |label1=[[Agastorophrynia]] |1={{clade |1=[[Bufonidae]] |2={{clade |1=[[Aromobatidae]] |2=[[Dendrobatidae]] }} }} }} |2={{clade |1={{clade <!--dummy clade for reduced horizontal spacing--> |label1=[[Diphyabatracea]] |1={{clade |1=[[Leptodactylidae]] |2={{clade |1=[[Allophrynidae]] |2=[[Centrolenidae]] }} }} }} |2={{clade |1=[[Ceratophryidae]] |2={{clade |1=[[Odontophrynidae]] |2={{clade |1={{clade |1=[[Cycloramphidae]] |2={{clade |1=[[Alsodidae]] |2=[[Hylodidae]] }} }} |2={{clade |1=[[Telmatobiidae]] |2={{clade |1=[[Batrachylidae]] |2=[[Rhinodermatidae]] }} }} }} }} }} }} }} }} }} }} }} <!-- end HYLOIDEA subtree--> |targetC=RANOIDEA |subcladeC={{clade |label1=[[Allodapanura]] |1={{clade |1=[[Microhylidae]] |label2=Afrobatrachia |2={{clade |label1=[[Xenosyneunitanura]] |1={{clade |1=[[Brevicipitidae]] |2=[[Hemisotidae]] }} |label2=[[Laurentobatrachia]] |2={{clade |1=[[Hyperoliidae]] |2=[[Arthroleptidae]] }} }} }} |label2=[[Natatanura]] |2={{clade |1=[[Ptychadenidae]] |label2=[[Victoranura]] |2={{clade |1=[[Micrixalidae]] |2={{clade |1=[[Phrynobatrachidae]] |2={{clade |1=[[Conrauidae]] |2={{clade |1={{clade <!--dummy clade for reduced horizontal spacing--> |label1=[[Pyxicephaloidea]] |1={{clade |1=[[Petropedetidae]] |2=[[Pyxicephalidae]] }} }} |2={{clade |1={{clade |1=[[Nyctibatrachidae]] |2=[[Ceratobatrachidae]] }} |label2=[[Saukrobatrachia]] |2={{clade |1={{clade |1=[[Ranixalidae]] |2=[[Dicroglossidae]] }} |label2=[[Aglaioanura]] |2={{clade |1={{clade |1=[[Rhacophoridae]] |2=[[Mantellidae]] }} |2=[[Ranidae]] }} }} }} }} }} }} }} }} }} <!-- end RANOIDEA subtree--> }}
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