Eurycephalosuchus
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''Eurycephalosuchus'' is an extinct genus of orientalosuchine
alligatoroid 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 c ...
from the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
Jiangxi Province Jiangxi (; ; formerly romanized as Kiangsi or Chianghsi) is a landlocked province in the east of the People's Republic of China. Its major cities include Nanchang and Jiujiang. Spanning from the banks of the Yangtze river in the north into hi ...
of
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 ...
. Known from a well preserved skull and mandible alongside various postcranial remains, ''Eurycephalosuchus'' possessed a short and broad skull with a very short skulltable. ''Eurycephalosuchus'' lived with at least one other crocodilian, an indetermined member of the clade
Brevirostres 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 ...
. The genus is
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
, containing only the species ''Eurycephalosuchus gannanensis''.


History and naming

''Eurycephalosuchus'' was discovered encased in a blox of matrix in the Municipality of Ganzhou City in 2021, located approximately northeast of
Ganzhou railway station Ganzhou railway station is a railway station located in Zhanggong District, Ganzhou, Jiangxi, China. It is one of two passenger railway stations in Ganzhou, the other being Ganzhou West railway station which is served by high-speed trains. It ha ...
and northeast of the Nankang District. The sediments the fossils were found in belong to the redbeds, which in this area are a part of the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
Hekou Formation, possibly correlating with the
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from ...
age. The block contained a nearly complete skull with the attached mandible, 14 vertebrae after the axis, 15 ribs, a partial front limb and several osteoderms, all of which together are designated as specimen IVPP V 31110, which forms 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 ...
of the genus. There is some compression that affected the skull during preservation, pushing parts of the frontal and prefrontal bones over the rear edge of the nasal bone. The same locality also yielded a shoulder blade articulated with a coracoid which are thought to belong to another type of crocodile, only described as being a member of
Brevirostres 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 ...
and notably bigger than ''Eurycephalosuchus''. The name ''Eurycephalosuchus'' is a composite word of the Greek "eurys" meaning wide or broad, "kephalos" meaning head and -suchus, derived from the Egyptian crocodile god Sobek. The species name on the other hand specifically points to Gannan, another name for Ganzhou, where the material was collected.


Description

''Eurycephalosuchus'' was a small to medium sized alligatoroid with an incredibly broad and short head, almost as wide as it is long. The fossil skull itself measures in dorsal cranial length, meaning measured from the tip of the snout to the rear end of the skulltable. Accounting for the slight compression the fossil underwent, a fully intact skull would likely measure , which is approximately the same as the maximum with across the quadratojugals. Measuring from the tip to the snout to the end of the quadrate condyle would render a skull only slightly longer than wide. Other parts of the skull likewise show bizarre proportions. The snout itself is short, only long and thus making up less than half of the total skull length. The skulltable, which is typically shorter than it is wide in crocodilians, takes this condition to an extreme. The length of this element is two thirds smaller than its width. The external nares are damaged towards the front, meaning that it is uncertain whether or not they form a singular confluent opening or not. Assuming a condition similar to '' Orientalosuchus'', a confluent opening would be wider than long. The
eyesocket In anatomy, the orbit is the cavity or socket of the skull in which the eye and its appendages are situated. "Orbit" can refer to the bony socket, or it can also be used to imply the contents. In the adult human, the volume of the orbit is , of ...
s were heavily distorted, but are clearly bigger than the triangular
supratemporal fenestra 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 ...
. Like the overall proportions of the skull, the supratemporals too are wider than long. The bones that form the edge of the fenestra, sans the
frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.''Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, par ...
, overhang the opening which differs from what is observed in any other member of Orientalosuchina that preserves this region of the skull. As in many crocodilians, ''Eurycephalosuchus'' has a notable notch located at the border of the
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
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 ...
that receives the enlarged fourth dentary tooth. This notch is moderately deep similar to ''Orientalosuchus'' and '' Leidyosuchus'', differing from the shallow notch of '' Jiangxisuchus'' and '' Dongnanosuchus''. The
nasal bones The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Eac ...
are paired and unfused and form a narrow point that extends into the external nares. The posterior end of these bones may have tapered following contact with the lacrimals, but the overlap caused by taphonomy makes it impossible to tell based on the fossil. The
maxillae 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 broad and long, ending in a wedge shaped process towards the front of the skull. When viewed from above the outer edges of the maxilla are sinuous, constricting at the contact with the premaxilla, broadening around the large 5th maxillary tooth before constricting and broadening once again shortly behind it. This is also reflected in sideview, with the tooth row showing two clear festoons, which peak with the 5th and 12th maxillary teeth respectively. The lacrimal bone is pointed towards the front, wedging itself into the maxilla, and broad towards the back, where the bone forms a process that contributes significantly towards the anterior margin of the orbits. The
prefrontal bone The prefrontal bone is a bone separating the lacrimal and frontal bones in many tetrapod skulls. It first evolved in the sarcopterygian clade Rhipidistia, which includes lungfish and the Tetrapodomorpha. The prefrontal is found in most modern and ...
s are triangular based on the better preserved left side, the
frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.''Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, par ...
contributes little to the orbital margins, but forms a moderately broad band between the eyes, broader than between the supratemporal fenestrae. The postorbital is almost rectangular with a slight point towards the front. The single parietal has almost straight contacts with the squamosals and a concave suture with the supraoccipital, which is exposed in dorsal view. The
jugal bone The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
is triradiate with a broad but short anterior ramus and a thinner but longer posterior ramus. The jugal contributes to the margins of the orbits and makes up the majority of the margin of the
infratemporal fenestra An infratemporal fenestra, also called the lateral temporal fenestra or simply temporal fenestra, is an opening in the skull behind the orbit in some animals. It is ventrally bordered by a zygomatic arch. An opening in front of the eye sockets, ...
. The quadratojugal bears a short spine that runs almost parallel to the rear edge of the infratemporal, resembling other alligatoroids like ''
Brachychampsa ''Brachychampsa'' is an extinct genus of alligatoroid, possibly a basal caiman. Specimens have been reported from New Mexico, Colorado, Wyoming, Montana, North and South Dakota, New Jersey, and Saskatchewan, though only those from Montana, Utah, ...
'' but differing from all other orientalosuchines. The
mandible 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 tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
shows a great increase in depth towards the back of the skull, starting off relatively shallow with a depth of only at the position of the fourth dentary tooth before increasing to a depth of around the
mandibular fenestra The skull is a bone protective Cranial cavity, 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 ...
. This means that the mandible increases in depth by over three times the number measured at the shallowest point. 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 ...
reaches its greatest breadth at the position of the 6th dentary tooth and the symphysis extends up to the 6th. The surangular bone interlocks with the dentary towards, forming two processes of nearly equal length. The surrangular thickens towards the back before tapering again as it contributes to the retroarticular process, almost reaching its end. The
angular bone The angular is a large bone in the lower jaw (mandible) of amphibians and reptiles (birds included), which is connected to all other lower jaw bones: the dentary (which is the entire lower jaw in mammals), the splenial, the suprangular, and the art ...
also reaches its greatest depth around the mandibular fenestra, tapering and underlying the dentary towards the front and tapering into the retroarticular process towards the back. When viewed from above the retroarticular process is roughly triangular with a ball-shaped end. Its surface is concave and bears a broad prominence that is however weakly developed. The premaxilla bears five teeth, the 3rd and 4th of which notably larger than the rest. Fourteen teeth sit in each maxilla, starting with a minute first tooth and rapidly increasing in length, reaching the maximum with a large
caniniform In mammalian oral anatomy, the canine teeth, also called cuspids, dog teeth, or (in the context of the upper jaw) fangs, eye teeth, vampire teeth, or vampire fangs, are the relatively long, pointed teeth. They can appear more flattened howeve ...
5th maxillary tooth. The following three teeth are all smaller than this caniniform and starting with the 8th these teeth are necked. They again show an increase in size, though not as rapidly and with as extreme results as the size increase leading up to the 5th tooth. These later teeth have bulb-shaped crowns with almost circular crosssections. The dentary toothrow is largely obscured by the teeth of the upper jaw, meaning the total number of teeth is unknown. As common in crocodilians, there is an enlarged 4th dentary tooth that slips into the notch present between the premaxilla and maxilla. Another large tooth is present further back, identified as the 12th dentary tooth based on ''Jiangxisuchus''. It slides in-between the 7th and 8th maxillary tooth. At least five more teeth sit behind this tooth, their bases visible beneath the overlying maxillary teeth. They seem to resemble their opposing teeth in size and shape. Based on skull length alone, ''Eurycephalosuchus'' may have been the smallest orientalosuchine, slightly shorter than ''
Krabisuchus ''Krabisuchus'' is an extinct genus of alligatoroid crocodylian that existed in what is now Thailand during the Eocene. It was first named by paleontologists Jeremy A. Martin and Komsorn Lauprasert in 2010, and the type species is ''K. siamogall ...
'' and with a narrower
occiput The occipital bone () is a cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone overlies the occipital lobes of the cereb ...
than ''
Eoalligator ''Eoalligator'' is an extinct genus of alligatoroid crocodilian from Paleocene deposits in China. Systematics ''Eoalligator'' was originally classified as an extinct relative of alligators. The holotype of ''E. chunyii'', IVPP V2716, is a partia ...
''.


Phylogeny

For the phylogenetic analysis the data matrix of Shan et al. (2021) was employed, specifically the first of the two employed by Shan et al. due to the inclusion of more terminal taxa. The strict consensus tree recovered agreed with the prior conclusion that Orientalosuchina had a basal position within
Globidonta Globidonta is a clade of alligatoroids that includes alligators, caimans, and closely related extinct forms. It is defined as a stem-based clade including ''Alligator mississippiensis'' (the American Alligator) and all forms more closely relate ...
. Even with ''Eurycephalosuchus'' included, Orientalosuchina was recovered as a
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
group. The genus clades with three other orientalosuchines from China and one from Vietnam, taking a sister relationship to the clade containing '' Jiangxisuchus'' and ''Eoalligator''. This relationship seems to match the distribution of the three taxa, all of them being known from the Late Cretaceous of China.


Paleobiology

The red beds where ''Eurycephalosuchus'' was found also preserve the fossils of a great variety of other reptiles including lizards, turtles and dinosaurs. A large bodied crocodilian only assigned to the clade Brevirostres was found and described alongside ''Eurycephalosuchus''. Given the proximaty between the two and similar preservation, they may have coexisted during the earliest Maastrichtian. This would not be out of the usual, as different crocodilian species are known to coexist both in the fossil record and in the modern day. Behavior, ecology, habitat preferences and tolerances may have all been factors that allowed the two to co-occur. Although the absence of skull material for the larger brevirostrine makes it impossible to determine its ecology, the greater size might indicate that it was the more dominant of the two species.


References

{{Taxonbar, from=Q115792592 Cretaceous crocodylomorphs Cretaceous reptiles of Asia Fossils of China Prehistoric pseudosuchian genera Fossil taxa described in 2022