Xilousuchus
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''Xilousuchus'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
poposauroid Poposauroidea is a clade of advanced pseudosuchians (archosaurs closer to crocodilians than to dinosaurs). It includes poposaurids, shuvosaurids, ctenosauriscids, and other unusual pseudosuchians such as ''Qianosuchus'' and ''Lotosaurus''. How ...
from lower
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
(
Olenekian In the geologic timescale, the Olenekian is an age in the Early Triassic epoch; in chronostratigraphy, it is a stage in the Lower Triassic series. It spans the time between Ma and Ma (million years ago). The Olenekian is sometimes divided into ...
stage) deposits of Fugu County of northeastern
Shanxi Province Shanxi (; ; Chinese postal romanization, formerly romanised as Shansi) is a landlocked Provinces of China, province of the China, People's Republic of China and is part of the North China region. The capital and largest city of the province is ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
. It is known from the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
, IVPP V 6026, a single well-preserved partial
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
including the
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
. It was found from the Heshanggou Formation of the Ordos Basin, Hazhen commune. It was first named by Xiao-Chun Wu in
1981 Events January * January 1 ** Greece enters the European Economic Community, predecessor of the European Union. ** Palau becomes a self-governing territory. * January 10 – Salvadoran Civil War: The FMLN launches its first major offensiv ...
and the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
is ''Xilousuchus sapingensis''. Wu (1981) referred ''Xilousuchus'' to the
Proterosuchia Proterosuchia is one of the suborders of the paraphyletic group Thecodontia; containing the most primitive and ancestral forms. These were primitive, vaguely crocodile-like, archosauriforms that mostly lived during the Early Triassic epoch. The ...
. Gower and Sennikov (1996) found it to be an erythrosuchian based strictly on the
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
. A more detailed re-description of the genus was provided by Nesbitt ''et al.'' (2010) and found poposauroid affinities. In his massive revision of archosaurs which included a large
cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
analysis, Sterling J. Nesbitt (2011) found ''Xilousuchus'' to be a poposauroid which is most closely related to ''
Arizonasaurus ''Arizonasaurus'' was a ctenosauriscid archosaur from the Middle Triassic (243 million years ago). ''Arizonasaurus'' is found in the Middle Triassic Moenkopi Formation of northern Arizona. A fairly complete skeleton was found in 2002 by Sterli ...
''. ''Xilousuchus'' is the oldest
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avian d ...
to date, although ''
Ctenosauriscus ''Ctenosauriscus'' is an extinct genus of sail-backed poposauroid archosaur from Early Triassic deposits of Lower Saxony in northern Germany. It gives its name to the family Ctenosauriscidae, which includes other sail-backed poposauroids such ...
'' and ''
Vytshegdosuchus ''Vytshegdosuchus'' is an extinct genus of paracrocodylomorph archosaur known from the Early Triassic (latest Olenekian stage) Yarenskian Gorizont of the Komi Republic of the European section of Russia. It contains a single species, ''Vytsheg ...
'' might be even older by less than one million year. Since ''Xilousuchus'' is a
suchia Suchia is a clade of archosaurs containing the majority of pseudosuchians (crocodilians and their extinct relatives). It was defined as the least inclusive clade containing '' Aetosaurus ferratus'', '' Rauisuchus tiradentes'', '' Prestosuchus ch ...
n archosaur, its early age suggests that most of the major groups of archosaurs (
ornithodira Avemetatarsalia (meaning "bird metatarsals") is a clade of diapsid reptiles containing all archosaurs more closely related to birds than to crocodilians. The two most successful groups of avemetatarsalians were the dinosaurs and pterosaurs. Dinos ...
ns,
ornithosuchid Ornithosuchidae is an extinct family of pseudosuchian archosaurs (distant relatives of modern crocodilians) from the Triassic period. Ornithosuchids were quadrupedal and facultatively bipedal (e.g. like chimpanzees), meaning that they had the a ...
s,
aetosaur Aetosaurs () are heavily armored reptiles belonging to the extinct order Aetosauria (; from Greek, (aetos, "eagle") and (, "lizard")). They were medium- to large-sized omnivorous or herbivorous pseudosuchians, part of the branch of archosaurs ...
s, and
paracrocodylomorphs Paracrocodylomorpha is a clade of pseudosuchian archosaurs. The clade includes the diverse and unusual group Poposauroidea as well as the generally carnivorous and quadrupedal members of Loricata, including modern crocodylians. Paracrocodylomorpha ...
) developed by the Early Triassic, soon after the appearance of the first archosaur.


Features

''Xilousuchus'' was probably quite a large animal when alive, possibly between three and four metres long. It had a small head, and a
sail A sail is a tensile structure—which is made from fabric or other membrane materials—that uses wind power to propel sailing craft, including sailing ships, sailboats, windsurfers, ice boats, and even sail-powered land vehicles. Sails may ...
along its neck and back. It probably had long and quite powerful legs, and would have been an active hunter. Only the partial
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
,
cervical vertebrae In tetrapods, cervical vertebrae (singular: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In ...
, and a few other fragments are preserved. ''Xilousuchus'' has a relatively small head and long neck, with a skull length of approximately 25 cm and neck of 45 cm. The skull is fragmentary, but much of the snout and
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
are present. Only one maxillary tooth has been fossilised. The maxilla has a partially-developed palatal process, and the angle of the dorsal process indicates that ''Xilousuchus'' had a large
antorbital fenestra An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with archosauriforms, first appearing during the Triassic Period. Among extant archosaurs, birds ...
. The single preserved maxillary tooth is large, almost 15 mm long, and in other
dental alveoli Dental alveoli (singular ''alveolus'') are sockets in the jaws in which the roots of teeth are held in the alveolar process with the periodontal ligament. The lay term for dental alveoli is tooth sockets. A joint that connects the roots of the t ...
where the teeth are missing replacement teeth are sometimes visible, indicating that it was a
polyphyodont A polyphyodont is any animal whose teeth are continually replaced. In contrast, diphyodonts are characterized by having only two successive sets of teeth. Polyphyodonts include most toothed fishes, many reptiles such as crocodiles and geckos, and ...
. The tooth has small
serrations Serration is a saw-like appearance or a row of sharp or tooth-like projections. A serrated cutting edge has many small points of contact with the material being cut. By having less contact area than a smooth blade or other edge, the applied p ...
, is curved backwards, and is laterally compressed, indicating that it could have punctured flesh or possibly sliced off chunks. The lacrimal and nasal bones are large but quite thin and fragile. The skull has a reasonably well-preserved
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
with thick walls and a small volume, and this shows that the brain was not large. It closely resembles that of ''Arizonasaurus,'' indicating that the two species are close relatives. The
dentary bone In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
is partially preserved for both left and right, and would have held thirteen teeth on each side, although only the
nutrient foramina The nutrient artery (arteria nutricia, or medullary), usually accompanied by one or two veins, enters the bone through the nutrient foramen, runs obliquely through the cortex, sends branches upward and downward to the bone marrow, which ramify in ...
remain. Like that of most archosaurs from the Triassic, the dentary is unspecialised. The cervical vertebrae have large, flattened
neural spines The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
which make up most of their height and would have formed a sail-like structure in life, similar to other
ctenosauriscids Ctenosauriscidae is an extinct family of pseudosuchian archosaurs within the clade Poposauroidea. Ctenosauriscids existed in Africa, Asia, Europe and North America during the Early Triassic to the Middle Triassic period (latest Olenekian to An ...
. Unusually, however, they curve slightly forwards. There is little space between the neural spines, indicating that ''Xilousuchus'' would probably not have had a very flexible neck. The spines gradually increase in height as they move backwards - that on the axis vertebra is 42 mm high, whereas that on the tenth cervical vertebra is 92 mm high. The height of the sail along the back is not known, as none of the dorsal vertebrae have been preserved, but it was probably similar to that in ''Arizonasaurus'' or ''
Hypselorhachis ''Hypselorhachis'' is a genus of extinct reptile, possibly a ctenosauriscid archosaur related to ''Ctenosauriscus''. It lived during the Triassic Period. It is currently known only from a single vertebra found from the Middle Triassic Manda Bed ...
.'' One
sacral vertebra The sacrum (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part ...
is present, and it shows a clear suture where it was joined to the sacral ribs and pelvis. The orientation of the sacral rib suggests that the ilium was downturned, although this is not certain. Two
caudal vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
are present as well, and these are poorly preserved with missing neural spines, but show facets where chevrons would have been attached. The proximal ends of two cervical ribs have also been fossilised, and these have two heads - a tuberculum and capitulum. They are slender, unlike those of
phytosaurs Phytosaurs (Φυτόσαυροι in greek) are an extinct group of large, mostly semiaquatic Late Triassic archosauriform reptiles. Phytosaurs belong to the order Phytosauria. Phytosauria and Phytosauridae are often considered to be equivalen ...
and
crocodylomorphs Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. During Mesozoic and early Cenozoic times, cr ...
. One
clavicle The clavicle, or collarbone, is a slender, S-shaped long bone approximately 6 inches (15 cm) long that serves as a strut between the shoulder blade and the sternum (breastbone). There are two clavicles, one on the left and one on the rig ...
and one
ungual An ungual (from Latin ''unguis'', i.e. ''nail'') is a highly modified distal toe bone which ends in a hoof, claw, or nail. Elephants and ungulates have ungual phalanges, as did the sauropods and horned dinosaurs. A claw is a highly modified ungual ...
are also known, but it is not known where the ungual is from. They have few distinguishing features and resemble those of other ctenosauriscids very closely.


Phylogeny

Cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
after Nesbitt, 2010:


References

{{Taxonbar, from=Q948654 Early Triassic reptiles of Asia Ctenosauriscids Fossil taxa described in 1981 Prehistoric pseudosuchian genera