Gryposuchinae: Difference between revisions

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updated evolution info with alternative theory and alternative cladogram
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==Classification==
Gryposuchinae was named in 2007 as a subfamily of closely related gavialid crocodilians. It was [[cladistics|cladistically]] defined as a [[stem-based taxon]] including ''Gryposuchus jessei'' and all crocodilians more closely related to it than to ''Gavialis gangeticus'' (the [[gharial]]) or ''Tomistoma schlegelii'' (the [[False gharial]]).<ref name=V-JBS07/> The [[tomistominae|tomistomines]] (including the living [[false gharial]]) were long thought to be classified as [[crocodile]]s and not closely related to [[gavialoids]].<ref name=Brochu2000>{{cite journal |last=Brochu |first=C.A. |author2=Gingerich, P.D. |year=2000 |title=New tomistomine crocodylian from the Middle Eocene (Bartonian) of Wadi Hitan, Fayum Province, Egypt |journal=University of Michigan Contributions from the Museum of Paleontology |volume=30 |issue=10 |pages=251–268}}</ref> However, recent molecular studies using [[DNA sequencing]] have consistently indicated that the [[false gharial]] (''Tomistoma'') (and by inference other related extinct forms in [[Tomistominae]]) actually belong to [[Gavialoidea]] (and [[Gavialidae]]).<ref name="Harshman2003">{{cite journal |pmid=12775527 |year=2003 |last1=Harshman |first1=J. |title=True and false gharials: A nuclear gene phylogeny of crocodylia |journal=Systematic Biology |volume=52|issue=3|pages=386–402|last2=Huddleston|first2=C. J. |last3=Bollback |first3=J. P. |last4=Parsons|first4=T. J.|last5=Braun|first5=M. J.|doi=10.1080/10635150309323 |url=http://si-pddr.si.edu/bitstream/handle/10088/6275/2003C_Harshman_et_al.pdf|doi-access=free }}</ref><ref name="Gatesy2003">{{cite journal |last=Gatesy |first=Jorge |author2=Amato, G. |author3=Norell, M. |author4=DeSalle, R. |author5= Hayashi, C. |year=2003 |title=Combined support for wholesale taxic atavism in gavialine crocodylians |journal=Systematic Biology |volume=52 |issue=3 |pages=403–422 |doi= 10.1080/10635150309329|url=http://www.faculty.ucr.edu/~mmaduro/seminarpdf/GatesyetalSystBiol2003.pdf|doi-access=free }}</ref><ref name="Willis2007">{{Cite journal | last1 = Willis | first1 = R. E. | last2 = McAliley | first2 = L. R. | last3 = Neeley | first3 = E. D. | last4 = Densmore Ld | first4 = L. D. | title = Evidence for placing the false gharial (''Tomistoma schlegelii'') into the family Gavialidae: Inferences from nuclear gene sequences | doi = 10.1016/j.ympev.2007.02.005 | journal = Molecular Phylogenetics and Evolution | volume = 43 | issue = 3 | pages = 787–794 | date = June 2007 | pmid = 17433721}}</ref><ref name="Gatesy2008">{{cite journal|last1=Gatesy |first1=J. |last2=Amato |first2=G. |year=2008 |title=The rapid accumulation of consistent molecular support for intergeneric crocodylian relationships |journal=[[Molecular Phylogenetics and Evolution]]|volume=48 |issue=3 |pages=1232–1237 |doi=10.1016/j.ympev.2008.02.009|pmid=18372192}}</ref><ref name=bite>{{cite journal| author=Erickson, G. M.| author2=Gignac, P. M.| author3=Steppan, S. J.| author4=Lappin, A. K.| author5=Vliet, K. A.| author6=Brueggen, J. A.| author7=Inouye, B. D.| author8=Kledzik, D.| author9=Webb, G. J. W. | year=2012 | title=Insights into the ecology and evolutionary success of crocodilians revealed through bite-force and tooth-pressure experimentation | journal=PLOS ONE | volume=7 |issue=3 |page=e31781 |doi=10.1371/journal.pone.0031781|editor1-last=Claessens|editor1-first=Leon|bibcode = 2012PLoSO...731781E | pmid=22431965 | pmc=3303775}}</ref><ref name="LeeYates2018">{{cite journal | author=Michael S. Y. Lee |author2=Adam M. Yates |date=27 June 2018 |title=Tip-dating and homoplasy: reconciling the shallow molecular divergences of modern gharials with their long fossil |journal=[[Proceedings of the Royal Society B]] |volume=285 |issue=1881 |doi=10.1098/rspb.2018.1071 |doi-access=free}}</ref><ref name="Hekkala2021">{{Cite journal|last=Hekkala |first=E. |last2=Gatesy |first2=J. |last3=Narechania |first3=A. |last4=Meredith |first4=R. |last5=Russello |first5=M. |last6=Aardema |first6=M. L. |last7=Jensen |first7=E. |last8=Montanari |first8=S. |last9=Brochu |first9=C. |last10=Norell |first10=M. |last11=Amato |first11=G. |date=2021-04-27 |title=Paleogenomics illuminates the evolutionary history of the extinct Holocene “horned” crocodile of Madagascar, Voay robustus |url=https://www.nature.com/articles/s42003-021-02017-0 |journal=Communications Biology |language=en |volume=4 |issue=1 |pages=1–11 |doi=10.1038/s42003-021-02017-0 |issn=2399-3642 |doi-access=free}}</ref>
Gryposuchinae was named in 2007 as a subfamily of closely related gavialid crocodilians. It was defined as a [[stem-based taxon]] including ''Gryposuchus jessei'' and all crocodilians more closely related to it than to ''Gavialis gangeticus'' (the [[gharial]]) or ''Tomistoma schlegelii'' (the [[False gharial]]).<ref name=V-JBS07/> Other gavialoids from the Americas include thoracosaurs from the [[Late Cretaceous]] and [[Paleocene]] and long-snouted [[Tomistominae|tomistomines]] from the [[Eocene]], but neither of these groups are closely related to gryposuchines. The relationship of tomistomines in particular is unclear, as they have also been classified as [[crocodile]]s.<ref name=___>{{cite journal |last=Brochu |first=C.A. |author2=Gingerich, P.D. |year=2000 |title=New tomistomine crocodylian from the Middle Eocene (Bartonian) of Wadi Hitan, Fayum Province, Egypt |journal=University of Michigan Contributions from the Museum of Paleontology |volume=30 |issue=10 |pages=251–268}}</ref>
 
A [[phylogenetic]] analysis conducted in the 2007 study found Gryposuchinae to include the genera ''Aktiogavialis'', ''Gryposuchus'', ''[[Ikanogavialis]]'', ''[[Piscogavialis]]'', and ''[[Siquisiquesuchus]]''. Below is a [[cladogram]] from the 2007 analysis showing the phylogenetic relationships of gryposuchines among [[Gavialoidea|gavialoids]]:<ref name=V-JBS07/> ''Hesperogavialis'' was excluded due to a lack of skull material, and ''[[Dadagavialis]]'' due to its 2018 discovery.<ref>{{Cite journal|last1=Salas-Gismondi|first1=Rodolfo|last2=Moreno-Bernal|first2=Jorge W.|last3=Scheyer|first3=Torsten M.|last4=Sánchez-Villagra|first4=Marcelo R.|last5=Jaramillo|first5=Carlos|date=2019-06-18|title=New Miocene Caribbean gavialoids and patterns of longirostry in crocodylians|journal=Journal of Systematic Palaeontology|volume=17|issue=12|pages=1049–1075|doi=10.1080/14772019.2018.1495275|s2cid=91495532|issn=1477-2019}}</ref>
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|5=''[[Aktiogavialis]]''
}} }} }} }} }} }} }}
 
Alternatively, a 2018 [[tip dating]] study by Lee & Yates simultaneously using [[morphology (biology)|morphological]], molecular ([[DNA sequencing]]), and [[stratigraphic]] ([[fossil]] age) data indicated that the members of Gryposuchinae and the [[genus]] ''[[Gryposuchus]]'' may in fact be [[paraphyletic]] and rather an [[evolutionary grade]] towards ''[[Gavialis]]'' and the [[gharial]], as shown in the cladogram below:<ref name="LeeYates2018"/>
 
{{clade| style=font-size:85%;line-height:85%
|grouplabel1={{clade labels |width=10em; |label1='''Gryposuchinae''' |top1=40% |color1=green}}
|label1='''[[Gavialidae]]'''
|1={{clade
|1={{clade
|1={{clade
|1={{clade
|1={{clade
|1={{clade
|1={{clade
|1={{clade
|1=''Gavialis gangeticus'' '''[[Gharial]]'''
|2=''[[Gavialis bengawanicus]]''{{extinct}} }}
|2=''[[Gavialis browni]]''{{extinct}} }}
|2=''[[Gryposuchus colombianus]]''{{extinct}} |barbegin2=green }}
|2=''[[Ikanogavialis]]''{{extinct}} |bar2=green }}
|2={{clade
|1=''[[Gryposuchus pachakamue]]''{{extinct}} |bar1=green
|2=''[[Piscogavialis]]''{{extinct}} |barend2=green }} }}
|2=''[[Harpacochampsa]]''{{extinct}} }}
|2={{clade
|1={{clade
|1=''[[Toyotamaphimeia]]''{{extinct}}
|2=''[[Penghusuchus]]''{{extinct}} }}
|2=''[[Gavialosuchus]]''{{extinct}} }} }}
|2={{clade
|1=''[[Tomistoma lusitanica]]''{{extinct}}
|2=''Tomistoma schlegelii'' '''[[False gharial]]''' }} }} }}
 
==Paleobiology==
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{{Taxonbar|from=Q5612452}}
 
[[Category:Crocodilians]]
[[Category:Gavialidae]]
[[Category:Paleogene crocodylomorphs]]