Asiatosuchus

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. 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. Based on largely complete skeletons of 'A.' germanicus and 'A. depressifrons, Asiatosuchus may have grown up to 4 metres (13 ft) long. 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. 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 form part of the mandibular symphysis. 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. 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. 'Asiatosuchus' depressifrons was first named in 1855 as Crocodilus depressifrons. The naming of this new species was based on a skull found in 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. 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. In 1964 Chinese paleontologist Yang Zhongjian named a new species of Asiatosuchus, A. nanlingensis, from the Shanghu Formation in Nanxiong, Guangdong Province, China. Small coprolites (fossilized feces) were found alongside the type specimen of A. nanlingensis. Recent studies have found A. nanlingensis to be valid, although not clearly referable to Asiatosuchus. 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. 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.