Voay Temporal range: Late Pleistocene, Divergence estimates suggest Oligocene origin | |
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Skull, American Museum of Natural History | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauromorpha |
Clade: | Archosauriformes |
Order: | Crocodilia |
Family: | Crocodylidae |
Subfamily: | Crocodylinae |
Genus: | † Voay Brochu, 2007 |
Type species | |
†Voay robustus (Grandidier & Vaillant, 1872) | |
Synonyms | |
Crocodylus robustusGrandidier & Vaillant, 1872 |
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. [2] Its name comes from the Malagasy word for crocodile.
One unusual feature of V. robustus that distinguishes it from other crocodilians is the presence of prominent "horns" extending from the posterior portion of the skull. They are actually the posterolaterally extended corners of the squamosal bone. Other related crocodilians such as Aldabrachampsus also had similar bony projections, although in Aldabrachampsus these projections were more like crests than horns. [3] Another diagnostic characteristic is the near-exclusion of the nasals from the external naris. It had a shorter and deeper snout than the extant Crocodylus niloticus, as well as relatively robust limbs. The osteoderms had tall keels and were dorsally symmetrical with curved lateral margins, running the entire length of the postcranial body. [4]
V. robustus would have measured around 3.5–4 m (11–13 ft) long and weighed about 170 kg (370 lb). [5] [6] These estimates suggest that V. robustus was the largest predator to have existed in Madagascar in recent times. Its size, stature, and presumed behavior is similar to the modern Nile crocodile (Crocodylus niloticus). Because V. robustus shared so many similarities with the Nile crocodile there must have been a great deal of interspecies competition for resources between the two crocodile genera if they were to have coexisted with one another. It has recently been proposed that the Nile crocodile only migrated to the island from mainland Africa after V. robustus had become extinct in Madagascar. [5] However, this was subsequently disproved after some Crocodylus specimens from Madagascar were found to be at least 7,500 years old and contemporaneous with Voay. [7]
When V. robustus was first described in 1872, it was originally assigned to the genus Crocodylus . [8] It was later found to morphologically have had more in common with the extant Osteolaemus, or dwarf crocodile, than Crocodylus. Some features it shared with Osteolaemus include a depressed pterygoid surface that forms a choanal "neck" on the palate. Because it was not close enough to be placed in the same genus as the dwarf crocodile, it was assigned to the new genus in 2007. Before this reassignment, the species was considered by some to be synonymous with Crocodylus niloticus. However, this was most likely due to a misinterpretation of remains from the living C. niloticus with V. robustus and the poor description of the original material from which the species was described. [9] [10] In contrast to the morphological similarities with Osteolaemus, a 2021 study using paleogenomics found Voay to be a sister group to Crocodylus, with both genera diverging in the mid-late Oligocene; this indicates that the apparent similarities with Osteolaemus are likely due to convergent evolution. [11]
The below cladogram shows the results of the latest study: [11]
Crocodylidae |
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(crown group) |
Crocodylinae is a subfamily of true crocodiles within the family Crocodylidae, and is the sister taxon to Osteolaeminae.
Crocodylus is a genus of true crocodiles in the family Crocodylidae.
The dwarf crocodile, also known as the African dwarf crocodile, broad-snouted crocodile or bony crocodile, is an African crocodile that is also the smallest extant (living) species of crocodile.
Boverisuchus is an extinct genus of planocraniid crocodyliforms known from the early to 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).
Aldabrachampsus is an extinct genus of small horned crocodile known from fragmentary remains. It lived during the Pleistocene on Aldabra Atoll, Seychelles in the western Indian Ocean. The name Aldabrachampsus dilophus means "Two-crested crocodile from Aldabra". It was a small animal, reaching a length of 2–2.5 m, comparable in size to the smallest extant crocodilians.
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.
Planocrania is an extinct genus of eusuchian crocodyliforms from what is now China. Two species are currently known to belong to the genus.
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" 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.
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.
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".
Portugalosuchus is an extinct genus of eusuchian crocodyliform that was possibly a basal crocodylian – if so then it would be the oldest known crocodylian to date. The type species is P. azenhae, described in 2018, and it is known from the Late Cretaceous (Cenomanian)-aged Tentugal Formation in Portugal. A 2021 morphological study recovered Portugalosuchus within Crocodylia as a member of Gavialidae closely related to similar "thoracosaurs", while also noting that it might also possibly be outside of Crocodylia completely. A 2022 tip dating analysis incorporating both morphological and DNA data placed Portugalosuchus outside of Crocodylia, as the sister taxon of the family Allodaposuchidae. A cladogram simplified after that analysis is shown below:
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.