Crocodyloidea

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Crocodyloidea
Temporal range: Late Cretaceous - Recent, 70–0  Ma
Nilecroc100.jpg
Crocodylus niloticus (Nile crocodile)
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauromorpha
Clade: Archosauriformes
Order: Crocodilia
Clade: Longirostres
Superfamily: Crocodyloidea
Fitzinger, 1826
Subgroups

Crocodyloidea is one of three superfamilies of crocodilians, the other two being Alligatoroidea and Gavialoidea, and it includes the crocodiles. Crocodyloidea may also include the extinct Mekosuchinae, [1] [2] native to Australasia from the Eocene to the Holocene, although this is disputed. [3] [4]

Classification

Cladistically, it is defined as Crocodylus niloticus (the Nile crocodile) and all crocodylians more closely related to C. niloticus than to either Alligator mississippiensis (the American alligator) or Gavialis gangeticus (the gharial). [5] This is a stem-based definition for crocodiles, and is more inclusive than the crown group Crocodylidae. [3] As a crown group, Crocodylidae only includes the last common ancestor of all extant (living) crocodiles and their descendants (living or extinct), whereas Crocodyloidea, as a stem group, also includes more basal extinct crocodile ancestors that are more closely related to living crocodiles than to alligators or gavialids. When considering only living taxa (neontology), this makes Crocodyloidea and Crocodylidae synonymous, and only Crocodylidae is used. Thus, Crocodyloidea 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. [6] However, recent molecular studies using DNA sequencing have rejected Brevirostres upon finding the crocodiles and gavialids to be more closely related than the alligators. [7] [8] [9] [3] [4] The new clade Longirostres was named by Harshman et al. in 2003. [7]

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, [3] which was expanded upon in 2021 by Hekkala et al. using paleogenomics by extracting DNA from the extinct Voay . [4]

The below cladogram shows the results of the latest study, and how Crocodyloidea may only contain one additional taxon beyond Crocodylidae:

Crocodylia
Alligatoroidea

extinct basal Alligatoroids

Alligatoridae
Caimaninae

Caiman Caiman crocodilus llanos white background.JPG

Melanosuchus Melanosuchus niger white background.jpg

Paleosuchus Dwarf Caiman white background.jpg

Alligatorinae

Alligator Alligator white background.jpg

(crown group)
(stem-based group)

extinct basal Crocodilians (including Mekosuchinae )

Longirostres
Crocodyloidea

"Crocodylus" megarhinus

Crocodylidae

Crocodylus Siamese Crocodile white background.jpg

Mecistops Crocodylus cataphractus faux-gavial d'Afrique2 white background.JPG

Osteolaemus Bristol.zoo.westafrican.dwarf.croc.arp. white background.jpg

(crown group)
(stem-based group)
Gavialoidea

extinct basal Gavialoids

Gavialidae

Gavialis Gavialis gangeticus (Gharial, Gavial) white background.jpg

Tomistoma Tomistoma schlegelii. white background.JPG

(crown group)
(stem-based group)
(crown group)
(crown group)

Related Research Articles

<span class="mw-page-title-main">Crocodylinae</span> Subfamily of crocodiles

Crocodylinae is a subfamily of true crocodiles within the family Crocodylidae, and is the sister taxon to Osteolaeminae.

<span class="mw-page-title-main">Gavialinae</span> Subfamily of gharial crocodylians

Gavialinae is a subfamily of large semiaquatic crocodilian reptiles, resembling crocodiles, but with much thinner snouts. Gavialinae is one of the two major subfamilies within the family Gavialidae - the other being the subfamily Tomistominae, which contains the false gharial and extinct relatives.

<span class="mw-page-title-main">Gavialidae</span> Family of gharial crocodylians

Gavialidae is a family of large semiaquatic crocodilians with elongated, narrow snouts. Gavialidae consists of two living species, the gharial and the false gharial, both occurring in Asia. Many extinct members are known from a broader range, including the recently extinct Hanyusuchus. Gavialids are generally regarded as lacking the jaw strength to capture the large mammalian prey favoured by crocodiles and alligators of similar size so their thin snout is best used to catch fish, however the false gharial has been found to have a generalist diet with mature adults preying upon larger vertebrates, such as ungulates.

<i>Gavialis</i> Genus of reptiles

Gavialis is a genus of crocodylians that includes the living gharial Gavialis gangeticus and one known extinct species, Gavialis bengawanicus.G. gangeticus comes from the Indian Subcontinent, while G. bengawanicus is known from Java. Gavialis likely first appeared in the Indian Subcontinent in the Pliocene and dispersed into the Malay Archipelago through a path called the Siva–Malayan route in the Quaternary. Remains attributed to Gavialis have also been found on Sulawesi and Woodlark Island east of the Wallace Line, suggesting a prehistoric lineage of Gavialis was able to traverse marine environments and reach places possibly as far as western Oceania.

<span class="mw-page-title-main">Mekosuchinae</span> Extinct subfamily of reptiles

Mekosuchinae is an extinct clade of crocodilians from the Cenozoic of Australasia. They represented the dominant group of crocodilians in the region during most of the Cenozoic. They first appear in the fossil record in the Eocene in Australia, and survived until the arrival of humans: in the Late Pleistocene in Australia and within the Holocene in the Pacific islands of Fiji, New Caledonia and Vanuatu.

<i>Tomistoma</i> Genus of crocodilians

Tomistoma is a genus of gavialid crocodilians. They are noted for their long narrow snouts used to catch fish, similar to the gharial. Tomistoma contains one extant (living) member, the false gharial, as well as potentially several extinct species: T. cairense, T. lusitanicumT. coppensi, and T. dowsoni. However, these species may need to be reclassified to different genera as studies have shown them to be paraphyletic, for example: previously assigned species T. taiwanicus from Taiwan, is reclassified to the genus Toyotamaphimeia, and T. dowsoni should be excluded from Tomistoma based on phylogenetic analysis.

<i>Boverisuchus</i> Extinct genus of reptiles

Boverisuchus is an extinct genus of planocraniid crocodyliforms known from the middle Eocene of Germany and western North America. It was a relatively small crocodyliform with an estimated total length of approximately 2.2–3.6 metres (7.2–11.8 ft).

<i>Gavialosuchus</i> Extinct genus of reptiles

Gavialosuchus is an extinct genus of gavialoid crocodylian from the early Miocene of Europe. Currently only one species is recognized, as a few other species of Gavialosuchus have since been reclassified to other genera.

<span class="mw-page-title-main">Alligatoroidea</span> Superfamily of reptiles

Alligatoroidea is one of three superfamilies of crocodylians, the other two being Crocodyloidea and Gavialoidea. Alligatoroidea evolved in the Late Cretaceous period, and consists of the alligators and caimans, as well as extinct members more closely related to the alligators than the two other groups.

Dollosuchoides, colloquially known as the Crocodile of Maransart, is an extinct monospecific genus of gavialoid crocodilian, traditionally regarded as a member of the subfamily Tomistominae. Fossils have been found in the Brussel Formation of Maransart, Belgium and date back to the middle Eocene.

Maroccosuchus zennaroi is an extinct gavialoid crocodylian from the Early Eocene of Morocco, traditionally regarded as a member of the subfamily Tomistominae.

<i>Planocrania</i> Extinct genus of reptiles

Planocrania is an extinct genus of eusuchian crocodyliforms from what is now China. Two species are currently known to belong to the genus.

<span class="mw-page-title-main">Gavialoidea</span> Superfamily of large reptiles

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, with Hanyusuchus having become extinct in the last few centuries.

<span class="mw-page-title-main">Tomistominae</span> Subfamily of reptiles

Tomistominae is a subfamily of crocodylians that includes one living species, the false gharial. Many more extinct species are known, extending the range of the subfamily back to the Eocene epoch. In contrast to the false gharial, which is a freshwater species that lives only in southeast Asia, extinct tomistomines had a global distribution and lived in estuaries and along coastlines.

<span class="mw-page-title-main">Brevirostres</span> Taxon of reptiles

Brevirostres is a paraphyletic group of crocodilians that included alligatoroids and crocodyloids. Brevirostres are crocodilians with small snouts, and are distinguished from the long-snouted gharials. It is defined phylogenetically as the last common ancestor of Alligator mississippiensis and Crocodylus niloticus and all of its descendants. This classification was based on morphological studies primarily focused on analyzing skeletal traits of living and extinct fossil species, and placed the gharials outside the group due to their unique skull structure, and can be shown in the simplified cladogram below:

<i>"Crocodylus" affinis</i> Species of reptile (fossil)

"Crocodylus" affinis is an extinct species of crocodyloid from the Eocene of Wyoming. Fossils were first described from the Bridger Formation by American paleontologist Othniel Charles Marsh in 1871. Marsh described the species, along with every other species of crocodyloid in the Bridger Formation, under the genus Crocodylus. The known specimen of "Crocodylus" affinis is a skull found at Grizzly Buttes, Wyoming, measuring 13 inches in length on the upper surface. Recent phylogenetic studies of crocodyloids show that "C." affinis is not a species of Crocodylus, but a genus has not yet been erected to include the species. Other Bridger species such as Crocodylus clavis and Brachyuranochampsa zangerli have been synonymized with "C." affinis.

"Crocodylus" megarhinus is an extinct species of crocodile from the Eocene of Egypt. A partial skull was found by British paleontologist Charles William Andrews in the Fayum Depression. Andrews named Crocodylus megarhinus in 1905 on the basis of the holotype skull. A complete skull was also uncovered from Egypt in 1907 but was not recognized as "C." megarhinus until 1927.

<span class="mw-page-title-main">Planocraniidae</span> Extinct family of reptiles

Planocraniidae is an extinct family of eusuchian crocodyliforms known from the Paleogene of Asia, Europe and North America. The family was coined by Li in 1976, and contains three genera, Boverisuchus, Duerosuchus and Planocrania. Planocraniids were highly specialized crocodyliforms that were adapted to living on land. They had extensive body armor, long legs, and blunt claws resembling hooves, and are sometimes informally called "hoofed crocodiles".

Tomistoma cairense is an extinct species of gavialoid crocodilian from the Lutetian stage of the Eocene era. It lived in North East Africa, especially Egypt. Remains of T. cairense have been found in the Mokattam Formation, in Mokattam, Egypt. Tomistoma cairense did not have a Maxilla process within their lacrimal gland, whereas all extant (living) crocodilians do.

<span class="mw-page-title-main">Longirostres</span> Clade of crocodilians

Longirostres is a clade of crocodilians that includes the crocodiles and the gavialids, to the exclusion of the alligatoroids. Defined in 2003 by Harshman et al., Longirostres is a crown group defined phylogenetically as including the last common ancestor of Crocodylus niloticus and Gavialis gangeticus and all of its descendants.

References

  1. Stein, Michael D.; Yates, Adam; Hand, Suzanne J.; Archer, Michael (2017). "Variation in the pelvic and pectoral girdles of Australian Oligo–Miocene mekosuchine crocodiles with implications for locomotion and habitus". PeerJ. 5: e3501. doi: 10.7717/peerj.3501 . PMC   5494174 . PMID   28674657.
  2. Iijima, M.; Kobayashi, Y. (2019). "Mosaic nature in the skeleton of East Asian crocodylians fills the morphological gap between "Tomistominae" and Gavialinae". Cladistics. 35 (6): 623–632. doi: 10.1111/cla.12372 . PMID   34618925. S2CID   91400957.
  3. 1 2 3 4 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 . PMC   6030529 . PMID   30051855.
  4. 1 2 3 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): 505. doi: 10.1038/s42003-021-02017-0 . ISSN   2399-3642. PMC   8079395 . PMID   33907305.
  5. Brochu, Christopher A. (May 2003). "Phylogenetic approaches toward crocodylian history". Annual Review of Earth and Planetary Sciences. 31 (31): 360. Bibcode:2003AREPS..31..357B. doi:10.1146/annurev.earth.31.100901.141308.
  6. 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.
  7. 1 2 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. Archived from the original (PDF) on 2022-10-09. Retrieved 2021-06-25.
  8. 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.
  9. 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.