Crocodyloidea Temporal range: | |
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Crocodylus niloticus (Nile crocodile) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauromorpha |
Clade: | Archosauriformes |
Order: | Crocodilia |
Clade: | Longirostres |
Superfamily: | Crocodyloidea Fitzinger, 1826 |
Subgroups | |
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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]
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 |
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(crown group) |
Crocodylinae is a subfamily of true crocodiles within the family Crocodylidae, and is the sister taxon to Osteolaeminae.
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.
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.
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.
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.
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.
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).
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.
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.
Maroccosuchus zennaroi is an extinct gavialoid crocodylian from the Early Eocene of Morocco, traditionally regarded as a member of the subfamily Tomistominae.
Paratomistoma is an extinct monospecific genus of gavialoid crocodylian. It is based on the holotype specimen CGM 42188, a partial posterior skull and lower jaw discovered at Wadi Hitan, Egypt, in Middle Eocene-age rocks of the Gehannam Formation. The skull is unfused but considered morphologically mature. Paratomistoma was named in 2000 by Christopher Brochu and Philip Gingerich; the type species is P. courti in honor of Nicholas Court, who found CGM 42188. They performed a phylogenetic analysis and found Paratomistoma to be a derived member of Tomistominae, related to the false gharial. It may have been a marine or coastal crocodilian.
Planocrania is an extinct genus of eusuchian crocodyliforms from what is now China. Two species are currently known to belong to the genus.
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.
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.
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:
"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.
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.
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.