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{{Short description|Superfamily of large reptiles}}
{{Short description|Superfamily of large reptiles}}
{{Automatic taxobox
{{Automatic taxobox
| fossil_range = [[Late Cretaceous]] - Recent, {{fossilrange|72.1|0}}
| fossil_range = [[Paleocene]] - [[Anthropocene|Recent]], {{fossilrange|61|0}}
| image = Gavialis gangeticus, ZOO Praha 045.jpg
| image = Gavialis gangeticus, ZOO Praha 045.jpg
| image_caption = Indian [[gharial]], ''Gavialis gangeticus''
| image_caption = Indian [[gharial]], ''Gavialis gangeticus''
Line 8: Line 8:
| subdivision_ranks = Subgroups
| subdivision_ranks = Subgroups
| subdivision =
| subdivision =
*[[Gavialidae]]
* {{extinct}} ''[[Argochampsa]]''
*{{extinct}}''[[Kentisuchus]]''
* {{extinct}} ''[[Dolichochampsa]]''<ref>{{cite journal | last1 = Jouve | first1 = S. | last2 = de Muizon | first2 = C. | last3 = Cespedes-Paz | first3 = R. | last4 = Sossa-Soruco | first4 = V. | last5 = Knoll | first5 = S. | year = 2020 | title = The longirostrine crocodyliforms from Bolivia and their evolution through the Cretaceous–Palaeogene boundary | journal = Zoological Journal of the Linnean Society | doi=10.1093/zoolinnean/zlaa081 }}</ref>
* {{extinct}} ''[[Eosuchus]]''
*{{extinct}}''[[Paratomistoma]]''
* {{extinct}} ''[[Eothoracosaurus]]''
*{{extinct}}''[[Maroccosuchus]]''
* {{extinct}} ''[[Ocepesuchus]]''
*{{extinct}}''[[Dollosuchoides]]''
*{{extinct}}''[[Dolichochampsa]]''?<ref>{{cite journal | last1 = Jouve | first1 = S. | last2 = de Muizon | first2 = C. | last3 = Cespedes-Paz | first3 = R. | last4 = Sossa-Soruco | first4 = V. | last5 = Knoll | first5 = S. | year = 2020 | title = The longirostrine crocodyliforms from Bolivia and their evolution through the Cretaceous–Palaeogene boundary | journal = Zoological Journal of the Linnean Society | doi=10.1093/zoolinnean/zlaa081 }}</ref>
* {{extinct}} ''[[Thoracosaurus]]''
*{{extinct}}''[[Ocepesuchus]]''?
* [[Gavialidae]]}}
}}


'''Gavialoidea''' is one of three [[superfamily (zoology)|superfamilies]] of [[crocodylian]]s, the other two being [[Alligatoroidea]] and [[Crocodyloidea]]. Although many extinct species are known, only the [[gharial]] ''Gavialis gangeticus'' and the [[false gharial]] ''Tomistoma schlegelii'' are alive today.
'''Gavialoidea''' is one of three [[superfamily (zoology)|superfamilies]] of [[crocodylian]]s, the other two being [[Alligatoroidea]] and [[Crocodyloidea]]. Although many extinct species are known, only the [[gharial]] ''Gavialis gangeticus'' and the [[false gharial]] ''Tomistoma schlegelii'' are alive today.


Extinct South American gavialoids likely [[Dispersal (ecology)|dispersed]] in the mid [[Tertiary]] from Africa and Asia.<ref>{{cite journal |last=Brochu |first=C. A. |year=2003 |title=Phylogenetic approaches toward crocodylian history |journal=Annual Review of Earth and Planetary Sciences |volume=31 |issue=31 |pages=357–397 |doi=10.1146/annurev.earth.31.100901.141308 |bibcode=2003AREPS..31..357B |url=https://www.researchgate.net/publication/253324276}}</ref>
Extinct South American gavialoids likely [[Dispersal (ecology)|dispersed]] in the mid [[Tertiary]] from Africa and Asia.<ref name=Brochu2003>{{cite journal |last=Brochu |first=C. A. |year=2003 |title=Phylogenetic approaches toward crocodylian history |journal=Annual Review of Earth and Planetary Sciences |volume=31 |issue=31 |pages=357–397 |doi=10.1146/annurev.earth.31.100901.141308 |bibcode=2003AREPS..31..357B |url=https://www.researchgate.net/publication/253324276}}</ref>
[[Fossil]] remains of the Puerto Rican gavialoid ''[[Aktiogavialis]] puertorisensis'' were discovered in a cave located in [[San Sebastián, Puerto Rico]] and dated to the [[Oligocene]]. This individual is thought to have crossed the [[Atlantic]] coming from Africa, indicating that this species was able to withstand saltwater.<ref name=Velez2007>{{cite journal |last=Vélez-Juarbe |first=J. |author2=Brochu, C. A.|author3=Santos, H. |name-list-style=amp |title=A gharial from the Oligocene of Puerto Rico: transoceanic dispersal in the history of a non-marine reptile |journal=Proceedings of the Royal Society |volume=274 |issue=1615|pages=1245–1254 |year=2007 |pmid=17341454 |pmc=2176176 |doi=10.1098/rspb.2006.0455}}</ref>
[[Fossil]] remains of the Puerto Rican gavialoid ''[[Aktiogavialis]] puertorisensis'' were discovered in a cave located in [[San Sebastián, Puerto Rico]] and dated to the [[Oligocene]]. This individual is thought to have crossed the [[Atlantic]] coming from Africa, indicating that this species was able to withstand saltwater.<ref name=Velez2007>{{cite journal |last=Vélez-Juarbe |first=J. |author2=Brochu, C. A.|author3=Santos, H. |name-list-style=amp |title=A gharial from the Oligocene of Puerto Rico: transoceanic dispersal in the history of a non-marine reptile |journal=Proceedings of the Royal Society |volume=274 |issue=1615|pages=1245–1254 |year=2007 |pmid=17341454 |pmc=2176176 |doi=10.1098/rspb.2006.0455}}</ref>


==Classification==
==Classification==
Gavialoidea is [[Cladistics|cladistically]] defined as ''Gavialis gangeticus'' (the [[gharial]]) and all [[crocodylia]]ns closer to it than to ''Alligator mississippiensis'' (the [[American alligator]]) or ''Crocodylus niloticus'' (the [[Nile crocodile]]).<ref name=Brochu2003/><ref name=Velez2007/> This is a [[stem-based taxon|stem-based]] definition for gavialoids, and is more inclusive than the [[crown group]] [[Gavialidae]].<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> As a crown group, Gavialidae only includes the [[last common ancestor]] of all [[extant taxon|extant]] (living) gavialids and their descendants (living or [[extinct]]), whereas Gavialoidea, as a stem group, also includes more [[basal (phylogenetics)|basal]] extinct gavialid ancestors that are more closely related to living gavialids than to [[crocodiles]] or [[alligators]]. When considering only living taxons ([[neontology]]), this makes Gavialoidea and Gavialidae [[synonym (taxonomy)|synonymous]], and only Gavialidae is used. Thus, Gavialoidea is only used in the context of [[paleontology]].
Gavialoidea contains the family [[Gavialidae]] and several more [[basal (phylogenetics)|basal]] extinct forms such as ''[[Thoracosaurus]]'' and ''[[Eosuchus]]''. Within Gavialidae are two subfamilies: [[Gavialinae]], which includes the living gharial, and [[Gryposuchinae]], which includes several extinct forms such as ''[[Gryposuchus]]'' and ''[[Aktiogavialis]]''.<ref name=Velez2007/>


Traditionally, crocodiles and alligators were considered more closely related and grouped together in the clade [[Brevirostres]], to the exclusion of the [[gharial]]s. This classification was based on [[morphology (biology)|morphological]] studies primarily focused on analyzing skeletal traits of living and extinct fossil species.<ref>{{cite journal | author=Holliday, Casey M. | author2=Gardner, Nicholas M. | year=2012 | title=A new eusuchian crocodyliform with novel cranial integument and its significance for the origin and evolution of Crocodylia | journal=PLOS ONE | volume=7 |issue=1 | pages=e30471 |doi=10.1371/journal.pone.0030471 |editor1-last=Farke |editor1-first=Andrew A |pmid=22303441 |pmc=3269432 |bibcode = 2012PLoSO...730471H }}</ref> However, recent molecular studies using [[DNA sequencing]] have rejected Brevirostres upon finding the crocodiles and gavialids to be more closely related than the alligators.<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="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"/><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> The new clade [[Longirostres]] was named by Harshman ''et al.'' in 2003.<ref name="Harshman2003"/>
In addition to these groups, recent molecular studies consistently indicate that the false gharial (and by inference other related extinct forms) traditionally viewed as belonging to the crocodylian subfamily [[Tomistominae]] actually belong to Gavialoidea.<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> As its name suggests, the false gharial was once thought to be only distantly related to the gharial despite its similar appearance. The false gharial and other tomistomines were traditionally classified within the superfamily [[Crocodyloidea]] as close relatives of [[crocodile]]s. This classification is based on morphological evidence, which, when incorporated into phylogenetic analyses, often place the group within Crocodyloidea.<ref name=Getal03>{{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>


In addition, these recent molecular DNA studies consistently indicate that the [[false gharial]] (''Tomistoma'') (and by inference other related extinct forms) traditionally viewed as belonging to the crocodylian subfamily [[Tomistominae]] actually belong to Gavialoidea (and Gavialidae).<ref name="Harshman2003"/><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"/><ref name=bite/><ref name="LeeYates2018"/><ref name="Hekkala2021"/> As its name suggests, the false gharial was once thought to be only distantly related to the gharial despite its similar appearance. The false gharial and other tomistomines were traditionally classified within the superfamily [[Crocodyloidea]] as close relatives of [[crocodile]]s, based solely on [[morphology (biology)|morphological]] evidence.<ref name="Gatesy2003"/>
Below is a cladogram from Vélez-Juarbe ''et al.'' (2007) showing the phylogenetic relationships of members of Gavialoidea, excluding tomistomines.<ref name=V-JBS07>{{cite journal |last=Vélez-Juarbe |first=Jorge |author2=Brochu, C.A. |author3= Santos, H. |year=2007 |title=A gharial from the Oligocene of Puerto Rico: transoceanic dispersal in the history of a non-marine reptile |journal=Proceedings of the Royal Society B |volume=274 |issue=1615 |pages=1245–1254 |doi=10.1098/rspb.2006.0455 |pmid=17341454 |pmc=2176176}}</ref>


A 2018 [[tip dating]] study by Lee & Yates simultaneously using [[morphology (biology)|morphological]], molecular ([[DNA sequencing]]), and [[stratigraphic]] ([[fossil]] age) data established the inter-relationships within [[Crocodilia]],<ref name="LeeYates2018"/> which was expanded upon in 2021 by Hekkala ''et al.'' using [[paleogenomics]] by extracting DNA from the extinct ''[[Voay]]''.<ref name="Hekkala2021"/> The tip dating analysis resolved the extinct ''[[Thoracosaurus]]'' and similar extinct close relatives as outside of Gavialoidea.
{{clade| style=font-size:100%;line-height:85%

|label1='''Gavialoidea'''&nbsp;
The below [[cladogram]] shows the results of the latest study, and Gavialoidea's relationships within [[Crocodylia]]:

{{clade| style=font-size:85%;line-height:85%
|label1='''[[Crocodylia]]'''
|sublabel1=([[crown group]])
|1={{clade
|1={{clade
|label1=[[Alligatoroidea]]
|1=''†[[Eothoracosaurus]]''
|sublabel1=([[stem-based taxon|stem group]])
|1={{clade
|1=extinct [[basal (phylogenetics)|basal]] Alligatoroids{{extinct}}
|label2=[[Alligatoridae]]
|sublabel2=([[crown group]])
|2={{clade
|label1=[[Caimaninae]]
|1={{clade
|1={{clade
|1=''[[Caiman (genus)|Caiman]]'' [[File:Caiman crocodilus llanos white background.JPG|90 px]]
|2=''[[Melanosuchus]]'' [[File:Melanosuchus niger white background.jpg|120 px]] }}
|2=''[[Paleosuchus]]'' [[File:Dwarf Caiman white background.jpg|90 px]] }}
|label2=[[Alligatorinae]]
|2=''[[Alligator]]'' [[File:Alligator white background.jpg|100 px]] }} }}
|label2=[[Longirostres]]
|sublabel2=([[crown group]])
|2={{clade
|label1=[[Crocodyloidea]]
|sublabel1=([[stem-based taxon|stem group]])
|1={{clade
|1=''[["Crocodylus" megarhinus]]''{{extinct}}
|label2=[[Crocodylidae]]
|sublabel2=([[crown group]])
|2={{clade
|1=''[[Crocodylus]]'' [[File:Siamese Crocodile white background.jpg|90 px]]
|2={{clade
|1=''[[Mecistops]]'' [[File:Crocodylus cataphractus faux-gavial d'Afrique2 white background.JPG|90 px]]
|2=''[[Osteolaemus]]'' [[File:Bristol.zoo.westafrican.dwarf.croc.arp. white background.jpg|90 px]] }} }} }}
|label2=[[Gavialoidea]]
|sublabel2=([[stem-based taxon|stem group]])
|2={{clade
|1=''[[Kentisuchus]]''{{extinct}}
|2={{clade
|2={{clade
|1=''†[[Thoracosaurus]]''
|1={{clade
|1=''[[Maroccosuchus]]''{{extinct}}
|2=''[[Paratomistoma]]''{{extinct}} }}
|2={{clade
|1=''[[Dollosuchoides]]''{{extinct}}
|2={{clade
|1=''[[Tomistoma cairense]]''{{extinct}}
|label2=[[Gavialidae]]
|sublabel2=([[crown group]])
|2={{clade
|2={{clade
|1=''[[Gavialis]]'' [[File:Gavialis gangeticus (Gharial, Gavial) white background.jpg|110 px]]
|1=''†[[Eosuchus]]''
|2=''[[Tomistoma]]'' [[File:Tomistoma schlegelii. white background.JPG|90 px]]
|2=''†[[Argochampsa]]''
}} }} }} }} }} }} }} }}
|label3=[[Gavialidae]]&nbsp;
|3={{clade
|1=''†[[Eogavialis]]''
|2={{clade
|1=''[[Gavialis]]''
|label2=†[[Gryposuchinae]]&nbsp;
|2={{clade
|1=''†[[Ikanogavialis]]''
|2=''†[[Piscogavialis]]''
|3=''†[[Siquisiquesuchus]]''
|4=''†[[Gryposuchus]]''
|5=''†[[Aktiogavialis]]''
}} }} }} }} }} }} }}


==References==
==References==

Revision as of 20:05, 25 June 2021

Gavialoidea
Temporal range: Paleocene - Recent, 61–0 Ma
Indian gharial, Gavialis gangeticus
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauromorpha
Clade: Archosauriformes
Order: Crocodilia
Clade: Longirostres
Superfamily: Gavialoidea
Hay, 1930
Subgroups

Gavialoidea is one of three superfamilies of crocodylians, the other two being Alligatoroidea and Crocodyloidea. Although many extinct species are known, only the gharial Gavialis gangeticus and the false gharial Tomistoma schlegelii are alive today.

Extinct South American gavialoids likely dispersed in the mid Tertiary from Africa and Asia.[2] Fossil remains of the Puerto Rican gavialoid Aktiogavialis puertorisensis were discovered in a cave located in San Sebastián, Puerto Rico and dated to the Oligocene. This individual is thought to have crossed the Atlantic coming from Africa, indicating that this species was able to withstand saltwater.[3]

Classification

Gavialoidea is cladistically defined as Gavialis gangeticus (the gharial) and all crocodylians closer to it than to Alligator mississippiensis (the American alligator) or Crocodylus niloticus (the Nile crocodile).[2][3] This is a stem-based definition for gavialoids, and is more inclusive than the crown group Gavialidae.[4] As a crown group, Gavialidae only includes the last common ancestor of all extant (living) gavialids and their descendants (living or extinct), whereas Gavialoidea, as a stem group, also includes more basal extinct gavialid ancestors that are more closely related to living gavialids than to crocodiles or alligators. When considering only living taxons (neontology), this makes Gavialoidea and Gavialidae synonymous, and only Gavialidae is used. Thus, Gavialoidea is only used in the context of paleontology.

Traditionally, crocodiles and alligators were considered more closely related and grouped together in the clade Brevirostres, to the exclusion of the gharials. This classification was based on morphological studies primarily focused on analyzing skeletal traits of living and extinct fossil species.[5] However, recent molecular studies using DNA sequencing have rejected Brevirostres upon finding the crocodiles and gavialids to be more closely related than the alligators.[6][7][8][4][9] The new clade Longirostres was named by Harshman et al. in 2003.[6]

In addition, these recent molecular DNA studies consistently indicate that the false gharial (Tomistoma) (and by inference other related extinct forms) traditionally viewed as belonging to the crocodylian subfamily Tomistominae actually belong to Gavialoidea (and Gavialidae).[6][10][11][7][8][4][9] As its name suggests, the false gharial was once thought to be only distantly related to the gharial despite its similar appearance. The false gharial and other tomistomines were traditionally classified within the superfamily Crocodyloidea as close relatives of crocodiles, based solely on morphological evidence.[10]

A 2018 tip dating study by Lee & Yates simultaneously using morphological, molecular (DNA sequencing), and stratigraphic (fossil age) data established the inter-relationships within Crocodilia,[4] which was expanded upon in 2021 by Hekkala et al. using paleogenomics by extracting DNA from the extinct Voay.[9] The tip dating analysis resolved the extinct Thoracosaurus and similar extinct close relatives as outside of Gavialoidea.

The below cladogram shows the results of the latest study, and Gavialoidea's relationships within Crocodylia:

Crocodylia
(crown group)

References

  1. ^ Jouve, S.; de Muizon, C.; Cespedes-Paz, R.; Sossa-Soruco, V.; Knoll, S. (2020). "The longirostrine crocodyliforms from Bolivia and their evolution through the Cretaceous–Palaeogene boundary". Zoological Journal of the Linnean Society. doi:10.1093/zoolinnean/zlaa081.
  2. ^ a b Brochu, C. A. (2003). "Phylogenetic approaches toward crocodylian history". Annual Review of Earth and Planetary Sciences. 31 (31): 357–397. Bibcode:2003AREPS..31..357B. doi:10.1146/annurev.earth.31.100901.141308.
  3. ^ a b Vélez-Juarbe, J.; Brochu, C. A. & Santos, H. (2007). "A gharial from the Oligocene of Puerto Rico: transoceanic dispersal in the history of a non-marine reptile". Proceedings of the Royal Society. 274 (1615): 1245–1254. doi:10.1098/rspb.2006.0455. PMC 2176176. PMID 17341454.
  4. ^ a b c d Michael S. Y. Lee; Adam M. Yates (27 June 2018). "Tip-dating and homoplasy: reconciling the shallow molecular divergences of modern gharials with their long fossil". Proceedings of the Royal Society B. 285 (1881). doi:10.1098/rspb.2018.1071.
  5. ^ Holliday, Casey M.; Gardner, Nicholas M. (2012). Farke, Andrew A (ed.). "A new eusuchian crocodyliform with novel cranial integument and its significance for the origin and evolution of Crocodylia". PLOS ONE. 7 (1): e30471. Bibcode:2012PLoSO...730471H. doi:10.1371/journal.pone.0030471. PMC 3269432. PMID 22303441.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  6. ^ a b c Harshman, J.; Huddleston, C. J.; Bollback, J. P.; Parsons, T. J.; Braun, M. J. (2003). "True and false gharials: A nuclear gene phylogeny of crocodylia" (PDF). Systematic Biology. 52 (3): 386–402. doi:10.1080/10635150309323. PMID 12775527.
  7. ^ a b Gatesy, J.; Amato, G. (2008). "The rapid accumulation of consistent molecular support for intergeneric crocodylian relationships". Molecular Phylogenetics and Evolution. 48 (3): 1232–1237. doi:10.1016/j.ympev.2008.02.009. PMID 18372192.
  8. ^ a b Erickson, G. M.; Gignac, P. M.; Steppan, S. J.; Lappin, A. K.; Vliet, K. A.; Brueggen, J. A.; Inouye, B. D.; Kledzik, D.; Webb, G. J. W. (2012). Claessens, Leon (ed.). "Insights into the ecology and evolutionary success of crocodilians revealed through bite-force and tooth-pressure experimentation". PLOS ONE. 7 (3): e31781. Bibcode:2012PLoSO...731781E. doi:10.1371/journal.pone.0031781. PMC 3303775. PMID 22431965.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  9. ^ a b c Hekkala, E.; Gatesy, J.; Narechania, A.; Meredith, R.; Russello, M.; Aardema, M. L.; Jensen, E.; Montanari, S.; Brochu, C.; Norell, M.; Amato, G. (2021-04-27). "Paleogenomics illuminates the evolutionary history of the extinct Holocene "horned" crocodile of Madagascar, Voay robustus". Communications Biology. 4 (1): 1–11. doi:10.1038/s42003-021-02017-0. ISSN 2399-3642.
  10. ^ a b Gatesy, Jorge; Amato, G.; Norell, M.; DeSalle, R.; Hayashi, C. (2003). "Combined support for wholesale taxic atavism in gavialine crocodylians" (PDF). Systematic Biology. 52 (3): 403–422. doi:10.1080/10635150309329.
  11. ^ Willis, R. E.; McAliley, L. R.; Neeley, E. D.; Densmore Ld, L. D. (June 2007). "Evidence for placing the false gharial (Tomistoma schlegelii) into the family Gavialidae: Inferences from nuclear gene sequences". Molecular Phylogenetics and Evolution. 43 (3): 787–794. doi:10.1016/j.ympev.2007.02.005. PMID 17433721.