"Crocodylus" megarhinus Temporal range: Early Oligocene, | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauromorpha |
Clade: | Archosauriformes |
Order: | Crocodilia |
Superfamily: | Crocodyloidea |
Species: | † "Crocodylus" megarhinus Andrews, 1905 |
Binomial name | |
"Crocodylus" megarhinus | |
Synonyms | |
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"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. [2]
"C." megarhinus shares many features with living crocodiles like the Nile crocodile (C. niloticus), including a robust triangular skull that is shorter than most other crocodiles. Similarities are also seen in the teeth of the two species. Like living crocodiles, "C." megarhinus has several constricted areas along the upper jaw that provide spaces for the teeth of the lower jaw when the mouth is closed. The proportions of "C." megarhinus and C. niloticus are so similar that American paleontologist Charles C. Mook considered it "very probable that C. megarhinus is a direct ancestor of C. niloticus." [2]
A second Fayum crocodilian, "Crocodylus" articeps, was named alongside "C." megarhinus. Andrews distinguished "C." articeps from "C." megarhinus on the basis of its narrower snout, which is more similar to the slender-snouted crocodile than the Nile crocodile. "C." articeps has recently been synonymized with "C." megarhinus, and may represent a less mature form in the species' population. [3]
Although it has traditionally been described as a species of Crocodylus, "C." megarhinus has been placed outside the genus in many recent phylogenetic analyses. A new genus has not yet been erected for the species. "C." megarhinus is usually found to be a basal crocodyline outside the genus but still more closely related to it than the false gharial or mekosuchines. Below is a cladogram modified from Puértolas et al. (2011) showing its phylogenetic placement among crocodiles: [4]
A 2017 paper by Wu et al. (2017) recovers "Crocodylus" megarhinus as a relative of mekosuchines. [5]
A 2018 tip dating study by Lee & Yates simultaneously using morphological, molecular (DNA sequencing), and stratigraphic (fossil age) data established the inter-relationships within Crocodylia, [6] which was expanded upon in 2021 by Hekkala et al. using paleogenomics by extracting DNA from the extinct Voay . [7]
The below cladogram shows the results of the latest study, and how "C." megarhinus may be outside of Crocodylidae, as a basal member of Crocodyloidea (and perhaps the sole 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.
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.
Crocodylus is a genus of true crocodiles in the family Crocodylidae.
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.
Asiatosuchus is an extinct genus of crocodyloid crocodilians that lived in Eurasia during the Paleogene. Many Paleogene crocodilians from Europe and Asia have been attributed to Asiatosuchus since the genus was named in 1940. These species have a generalized crocodilian morphology typified by flat, triangular skulls. The feature that traditionally united these species under the genus Asiatosuchus is a broad connection or symphysis between the two halves of the lower jaw. Recent studies of the evolutionary relationships of early crocodilians along with closer examinations of the morphology of fossil specimens suggest that only the first named species of Asiatosuchus, A. grangeri from the Eocene of Mongolia, belongs in the genus. Most species are now regarded as nomina dubia or "dubious names", meaning that their type specimens lack the unique anatomical features necessary to justify their classification as distinct species. Other species such as "A." germanicus and "A." depressifrons are still considered valid species, but they do not form an evolutionary grouping with A. grangeri that would warrant them being placed together in the genus Asiatosuchus.
Voay is an extinct genus of crocodile from Madagascar that lived during the Late Pleistocene to Holocene, containing only one species, V. robustus. Numerous subfossils have been found, including complete skulls, noted for their distinctive pair of horns on the posterior, as well as vertebrae and osteoderms from such places as Ambolisatra and Antsirabe. The genus is thought to have become extinct relatively recently. It has been suggested to have disappeared in the extinction event that wiped out much of the endemic megafauna on Madagascar, such as the elephant bird and Malagasy hippo, following the arrival of humans to Madagascar around 2000 years ago. Its name comes from the Malagasy word for crocodile.
Kentisuchus is an extinct genus of gavialoid crocodylian, traditionally regarded as a member of the subfamily Tomistominae. Fossils have been found from England and France that date back to the early Eocene. The genus has also been recorded from Ukraine, but it unclear whether specimens from Ukraine are referable to Kentisuchus.
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.
Prodiplocynodon is an extinct genus of basal crocodyloid crocodylian. It is one of the only crocodyloids known from the Cretaceous and existed during the Maastrichtian stage. The only species of Prodiplocynodon is the type species P. langi from the Lance Formation of Wyoming, known only from a single holotype skull lacking the lower jaw.
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, native to Australasia from the Eocene to the Holocene, although this is disputed.
Rimasuchus is an extinct genus of crocodile from the Miocene of Egypt and possibly Libya. Only one species - Rimasuchus lloydi - is currently known. It was previously thought to be a species of Crocodylus, but is now thought to be more closely related to the modern African dwarf crocodiles (Osteolaemus).
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.
Osteolaeminae is a subfamily of true crocodiles within the family Crocodylidae containing the dwarf crocodiles and slender-snouted crocodiles, and is the sister taxon to Crocodylinae.
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" acer is an extinct species of crocodyloid from the Eocene of Utah. A single well preserved skull was described by paleontologist Edward Drinker Cope in 1882 and remains the only known fossil of the species. It was found from the Wasatchian-age Green River Formation. "C." acer had a long, narrow snout and a low, flattened skull.
Crocodylus palaeindicus is an extinct species of crocodile from southern Asia. C. palaeindicus lived from the Miocene to the Pliocene. It may be an ancestor of the living Mugger crocodile.
Crocodylus checchiai is an extinct species of crocodile from the Miocene to Pliocene of Libya and Kenya. C. checchiai was named in 1947 based on a skull from the Sahabi Formation. Remains from the lower Nawata Formation in the Turkana Basin of Kenya that were first attributed to the Nile crocodile have now been reassigned to C. checchiai, extending its geographic range. The morphology of the species, in particular the pronounced rostral boss, indicates that it may be the connecting link between African and American species of the genus Crocodylus.
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.