Asiatosuchus | |
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A. nanlingensis specimens | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauromorpha |
Clade: | Archosauriformes |
Order: | Crocodilia |
Superfamily: | Crocodyloidea |
Genus: | † Asiatosuchus Mook, 1940 |
Type species | |
†Asiatosuchus grangeri Mook, 1940 | |
Species | |
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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. [2]
Like most other Paleogene crocodyloids, Asiatosuchus has a generalized crocodilian skull that is triangular in shape when viewed from above. Asiatosuchus species have teeth in the upper jaw that completely overlap the teeth in the lower jaw, giving them overbites. An overbite is a primitive feature among crocodyloids because modern crocodiles have teeth in the upper and lower jaws that interlock with each other with little overlap. Asiatosuchus can be distinguished from other early crocodyloids by its extended mandibular symphysis, the region where the two halves of the lower jaws connect. In many crocodyloids this joint is formed from two pairs of bones, the dentary bones and the splenial bones, but in Asiatosuchus it is only formed by the dentary bones. [2] Based on largely complete skeletons of "A." germanicus and "A." depressifrons, Asiatosuchus may have grown up to 4 metres (13 ft) long. [3]
A. grangeri, the type species of Asiatosuchus, was named by paleontologist Charles Mook in 1940. It was named on the basis of a lower jaw and pieces of a skull from the Irdin Manha Formation of Inner Mongolia, China, which dates back to the Middle Eocene. These fossils were discovered by the American Museum of Natural History's Central Asiatic Expedition of 1930 near Erenhot. Mook named Asiatosuchus grangeri after Walter W. Granger, a vertebrate paleontologist with the American Museum of Natural History and a member of the expedition. [4] Mook thought that Asiatosuchus grangeri was closely related to species of Crocodylus (modern crocodiles) but different in having 17 teeth in each half of the lower jaw and a splenial bone that does not form part of the mandibular symphysis. [2]
Well-preserved remains of a crocodyloid were first described from Germany and France in 1966 and placed in a new species of Asiatosuchus, A. germanicus. [2] The German remains came from the Messel Pit quarry, a fossil site that has preserved many forms of life that inhabited a series of anoxic lakes and surrounding subtropical forests during the Eocene. Of all the species that have been assigned to Asiatosuchus, "A." germanicus is known from the most complete material. [5]
"Asiatosuchus" depressifrons was first named in 1855 as Crocodilus depressifrons. The naming of this new species was based on a skull found in the Sables du Castrais Formation, France that dates back to the Early Eocene. The skull was illustrated in the 1855 paper but it was not thoroughly described. The fossil has since become heavily pyritized, losing much of its original anatomical detail. After its naming, several other crocodilian fossils in European museum collections were labeled as C. depressifrons. The species name depressifrons refers to the flattened shape of the frontal bone in the skull, a feature that is shared by all fossils attributed to the species. The fossils are also similar in having 6 pairs of teeth lining the symphysis at the tip of the lower jaw. As was the case for many other Paleogene crocodyloids, "A." depressifrons was originally placed in the still-living genus Crocodylus because the overall shape of its skull is similar to those of living crocodiles. Soon after Mook named Asiatosuchus grangeri, C. depressifrons was reassigned to Asiatosuchus. [2]
Many new and much more complete fossils of "A." depressifrons have been found from Early Eocene deposits in Belgium. Together these specimens provide details on most of the skeleton. "A." depressifrons can be distinguished from all other species of Asiatosuchus by a combination of several characteristics including a large hole and a depressed area on the jugal bone of the skull, a frontal bone that does not touch the supratemporal fenestrae (two holes at the top of the skull behind the eye sockets), and a postorbital bone behind the eye socket that is visible when the skull is viewed from the side. Another distinguishing feature of "A." depressifrons is its lack of an overbite. [2]
In 1964 Chinese paleontologist Yang Zhongjian named a new species of Asiatosuchus, "A." nanlingensis, based on fragmentary material from the Shanghu Formation in Nanxiong, Guangdong Province, China. [6] Small coprolites (fossilized feces) were found alongside the type specimen of A. nanlingensis. [7] A. nanlingensis was discovered concurrently with the similar Eoalligator chunyii , and a 2016 study proposed that they are synonymous. [8] A subsequent paper argued otherwise, [9] but other researchers agree with the initial assessment synonymizing "A." nanlingensis and "E." chunyii. [10] Since then, different analyses have recovered the species as a member of Mekosuchinae, [1] or as basal to a clade including mekosuchines and Crocodylidae. [10]
Two species of Asiatosuchus were named from Russia, A. zajsanicus in 1982 and A. volgensis in 1993. A. volgensis and A. zajsanicus were regarded as nomina dubia by Angielczyk and Gingerich (1998) because they are based on fossil specimens that preserve very little anatomical detail. A. zajsanicus was later reassigned to Dollosuchus , a genus of tomistomine crocodilians, by Efimov (1988), but Brochu (2007) treated Dollosuchus as dubious. [2]
Several crocodilian fossils from the Paleogene of North America have also been proposed to belong to Asiatosuchus. In comparison to A. grangeri, "Crocodylus" affinis from the Bridger Formation in Wyoming has a similarly shaped splenial bone in the lower jaw and frontal bone in the skull. Although "C." affinis is known from a complete skull, the skull material of A. grangeri is too fragmentary to support "C." affinis being classified within Asiatosuchus. [2]
The crocodyloid species "Crocodylus" monsvialensis was named from Early Oligocene deposits in Monteviale, Italy in 1914 and reassigned to Asiatosuchus in 1993, although subsequent authors questioned this referral and considered it synonymous with Diplocynodon ratelii . [2] "Crocodylus" vicetinus from the Middle Eocene locality of Monte Bolca, was assigned to Asiatosuchus sp. by Kotsakis et al. (2004) pending revision of the Mount Bolca crocodilian material. [11]
A partial skeleton of a crocodyloid from the Sulaiman Mountains of Pakistan was tentatively attributed to Asiatosuchus. The fossil was found in the Middle Eocene Drazinda Formation, a marine deposit which has also preserved the remains of archaeocete whales. The presence of a possible specimen of Asiatosuchus in marine deposits suggests that these crocodilians could have tolerated prolonged periods of time in the ocean, an ability that would have aided in the dispersal of early crocodyloids across Europe and Asia. [5]
Phylogenetic analyses of the evolutionary relationships of crocodilians place Asiatosuchus as a member of a clade or evolutionary grouping called Crocodyloidea, which includes living crocodiles and their extinct relatives. Recent phylogenetic analyses place Asiatosuchus as a basal ("primitive") member of this clade, close to the split between Crocodyloidea and Alligatoroidea, the group that includes living alligators, caimans, and their extinct relatives. Many of the species that are most closely related to species of Asiatosuchus were originally classified in the genus Crocodilus because they superficially resemble modern crocodiles. However, the majority of early crocodilians, even some early alligatoroids, resembled modern crocodiles because a triangular, crocodile-shaped head is a primitive condition for crocodilians.
Some phylogenetic analyses have placed "Asiatosuchus" germanicus as the sister taxon or closest relative of a group called Mekosuchinae. Mekosuchines are a group of crocodyloids from Australia and the South Pacific that are unusual in that they were highly specialized for life on land. If "A." germanicus is the sister taxon of Mekosuchinae, it may have been close to the ancestry of the group. The earliest known and most basal mekosuchine, Kambara , lived during the same time as Asiatosuchus, suggesting that Asiatosuchus or an Asiatosuchus-like crocodyloid could have dispersed into Australia as the ancestor of mekosuchines. Despite the results of the phylogenetic analysis, "A." germanicus is an unlikely candidate for the ancestor of mekosuchines because it lived very far from Australia and the likelihood that it could have reached Australia from Europe is very low. [12]
Most phylogenetic analyses do not support the idea that all species of Asiatosuchus belong to their own clade. Instead they find that Asiatosuchus species form a paraphyletic grouping, meaning that Asiatosuchus represents an evolutionary grade of successively more derived crocodyloids rather than its own separate lineage. Since a genus name is normally only applied to a monophyletic grouping by researchers who study prehistoric crocodilians, the type species A. grangeri is now considered the only valid species within Asiatosuchus. The species "A." germanicus and "A. depressifrons are written in quotes because they do not belong to Asiatosuchus and have not yet been given different genus names.The relationships of other putative Asiatosuchus species are uncertain because only A. grangeri, "A." germanicus, and "A." depressifrons have enough distinguishing features to be included in phylogenetic analyses. [2]
Below is a cladogram from Delfino and Smith (2009) showing that Asiatosuchus species represent a non-monophyletic grouping. Delfino and Smith considered these relationships to have very weak support because only a few characteristics entered into the data matrix differed between Asiatosuchus species, and none differed between A. grangeri and "C." affinis. [2]
Crocodylia |
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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, [13] which was expanded upon in 2021 by Hekkala et al. using paleogenomics by extracting DNA from the extinct Voay . [14]
The below cladogram shows the results of the latest studies, which placed Asiatosuchus outside of Crocodyloidea, as more basal than Longirostres (the combined group of crocodiles and gavialids). [13]
Crocodylia |
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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, first appearing in the fossil record in the Eocene of Australia, and surviving until the arrival of humans: the Late Pleistocene on the Australian continent and during the Holocene in the Pacific islands of Fiji, New Caledonia and Vanuatu.
Mekosuchus is a genus of extinct Australasian mekosuchine crocodilian. Species of Mekosuchus were generally small-sized, terrestrial animals with short, blunt-snouted heads and strong limbs. Four species are currently recognized, M. inexpectatus, M. whitehunterensis, M. sanderi and M. kalpokasi, all known primarily from fragmentary remains.
Quinkana is an extinct genus of mekosuchine crocodylians that lived in Australia from about 25 million to about 10,000 years ago, with the majority of fossils having been found in Queensland. Four species are currently recognized, all of which have been named between 1981 and 1997. The two best understood species are Q. fortirostrum, the type species, and Q. timara, a more gracile form from the Miocene. The other two species, Q. babarra and Q. meboldi, from the Pliocene and Oligocene respectively, are only known from a few poorly preserved bone fragments. The name Quinkana comes from the "Quinkans", a legendary folk spirit from Gugu-Yalanji mythology.
Australosuchus is an extinct monospecific genus of crocodylian belonging to the subfamily Mekosuchinae. The type and only known species Australosuchus clarkae lived during the Late Oligocene and the Early Miocene in the Lake Eyre Basin of South Australia. It was described in 1991 by Paul Willis and Ralph Molnar from fossil material discovered at Lake Palankarinna.
Kambara is an extinct genus of mekosuchine crocodylian that lived during the Eocene epoch in Australia. It is generally thought to have been a semi-aquatic generalist, living a lifestyle similar to many of today's crocodiles. Four species are currently recognised, the sympatric Kambara murgonensis and Kambara implexidens from sediments near Murgon, the poorly preserved Kambara molnari from the Rundle Formation and the youngest of the four, Kambara taraina, also from the Rundle Formation. Kambara were medium-sized crocodilians, with mature specimens generally reaching lengths from 3–4 m (9.8–13.1 ft).
Eoalligator is an extinct genus of alligatoroid crocodilian from Paleocene deposits in China.
Harpacochampsa is a poorly known Early Miocene crocodilian from the Bullock Creek lagerstätte of the Northern Territory, Australia. The current specimen consists of a partial skull and fragments of a long, slender snout reminiscent of that of a false gharial, demonstrating that it was a piscivore in life.
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.
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.
Arenysuchus is an extinct monospecific genus of allodaposuchid eusuchian crocodylomorph from Late Cretaceous deposits of north Spain. It is known from the holotype MPZ ELI-1, a partial skull from Elías site, and from the referred material MPZ2010/948, MPZ2010/949, MPZ2010/950 and MPZ2010/951, four teeth from Blasi 2 site. It was found by the researchers José Manuel Gasca and Ainara Badiola from the Tremp Formation, in Arén of Huesca, Spain. It was first named by Eduardo Puértolas, José I. Canudo and Penélope Cruzado-Caballero in 2011 and the type species is Arenysuchus gascabadiolorum.
"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.
Astorgosuchus is an extinct monospecific genus of crocodilian, closely related to true crocodiles, that lived in Pakistan during the late Oligocene period. This crocodile may have reached lengths of up to 7–8 m (23–26 ft) and is known to have preyed on many of the large mammals found in its environment. Bite marks of a large crocodile have been found on the bones of juvenile Paraceratherium, however if these were left by Astorgosuchus cannot be said with certainty. The genus contains a single species, Astorgosuchus bugtiensis, which was originally named as a species of Crocodylus in 1908 and was moved to its own genus in 2019.
Protoalligator is an extinct genus of alligatoroid crocodilian found in the Anhui province of China and lived during the Paleocene.
Ultrastenos is an extinct genus of Australian mekosuchine crocodilian that lived during the Late Oligocene in northwestern Queensland, Australia. Following its discovery, it was speculated that Ultrastenos was a slender-snouted animal similar to modern gharials or freshwater crocodiles due to the seemingly abruptly narrowing mandible. However, a later study found that this was a missinterpretation of the fossil specimen and that Ultrastenos instead had a more generalized lower jaw. The same publication also provided evidence that the fossils of Ultrastenos belonged to the same animal previously named "Baru" huberi, adding further evidence to the idea that the animal was short snouted, contrary to the initial hypothesis. Given that "Baru" huberi was named first, the type species of Ultrastenos changed from U. willisi to U. huberi in accordance with the rules of the ICZN. Ultrastenos was a small mekosuchine, measuring upwards of 1.5 m long.
Kalthifrons is an extinct monospecific genus of mekosuchine crocodylian known from the Pliocene Tirari Formation of Australia. More specifically, Kalthifrons was recovered from the Mampuwordu Sand Member, which underlies the younger sediments of the Pompapillina Member. This is significant, as the latter preserves some of the earliest records of the genus Crocodylus in Australia, which would eventually go on to replace mekosuchines. It is currently unclear whether or not the Tirari Crocodylus directly outcompeted Kalthifrons or simply moved into the region after the niche was left empty by the extinction of the local mekosuchines. Should the later be the case, then Kalthifrons may have simply been the victim of global cooling and aridification. A point in favour of the competition hypothesis is that both Kalthifrons and the Tirari Crocodylus have broadly similar skull forms, with both being interpreted as generalist semi-aquatic predators much like many of today's crocodiles. Though far from large, Kalthifrons was nonetheless bigger than many other mekosuchines such as Trilophosuchus and Mekosuchus. The genus is monotypic, meaning it contains only a single species, Kalthifrons aurivellensis.
Qianshanosuchus is a genus of basal crocodyloid from the Paleocene of the Qianshan Basin, China. The fossil material, which includes an incomplete skull and parts of the lower jaw, show various features usually associated with juvenile crocodiles alongside various unique traits that were used to erect a new genus. It is the first and only basal crocodyloid currently known from the Paleocene of China, which had previously only yielded alligatoroids and planocraniids. Its presence in this part of the world and its basal position to species of the genus Asiatosuchus supports the idea that crocodyloids dispersed from Asia into Europe. Qianshanosuchus only includes a single species, Qianshanosuchus youngi.
Antecrocodylus is a genus of crocodilian from the middle Miocene of Chiang Muan and Mae Moh, Thailand. The holotype consists of the back of the skull and an associated mandible. While far from complete and heavily damaged, the material highlights how little is known about the crocodylid fauna of Miocene Asia. Furthermore, Antecrocodylus was recovered as the basalmost member of Crocodylidae, having diverged before osteolaemines and crocodylines split from one another. This may suggest that it could provide significant information regarding the origins and evolution of modern crocodiles.
Ahdeskatanka is an extinct genus of alligator from the Early Eocene Golden Valley Formation of North Dakota, USA. Ahdeskatanka had a short, rounded snout with globular teeth that are well-suited for crushing hard-shelled prey, though its exact ecology is not known. Ahdeskatanka inhabited the vast wetlands that covered much of western North Dakota during the Early Eocene Climatic Optimum, an environment it shared with at least three other crocodilians. These include the large caiman Chrysochampsa and at least two unnamed forms, one a large crocodyloid and one more similar to Ahdeskatanka. Phylogenetic analysis suggests that it was an early diverging member of the Alligatorinae, possibly related to Allognathosuchus, though its position is not very stable. Only a single species, Ahdeskatanka russlanddeutsche, is placed in this genus.