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''Syntomiprosopus'' (meaning "short face") 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
archosauriform Archosauriformes (Greek for 'ruling lizards', and Latin for 'form') is a clade of diapsid reptiles that developed from archosauromorph ancestors some time in the Latest Permian (roughly 252 million years ago). It was defined by Jacques Gauthier ...
, possibly a
crocodylomorph 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, cro ...
from the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch ...
period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
of
Arizona Arizona ( ; nv, Hoozdo Hahoodzo ; ood, Alĭ ṣonak ) is a state in the Southwestern United States. It is the 6th largest and the 14th most populous of the 50 states. Its capital and largest city is Phoenix. Arizona is part of the Fou ...
. The type and only known species is ''S. sucherorum''. ''Syntomiprosopus'' was unusually short-snouted, comparable to 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 ...
notosuchian Notosuchia is a suborder of primarily Gondwanan mesoeucrocodylian crocodylomorphs that lived during the Jurassic and Cretaceous. Some phylogenies recover Sebecosuchia as a clade within Notosuchia, others as a sister group (see below); if Sebecosu ...
''
Simosuchus ''Simosuchus'' (meaning "pug-nosed crocodile" in Greek, referring to the animal's blunt snout) is an extinct genus of notosuchian crocodylomorphs from the Late Cretaceous of Madagascar. It is named for its unusually short skull. Fully grown indi ...
'', and is regarded as an example of convergent evolution between Triassic stem-
archosaurs 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-avi ...
and Cretaceous archosaurs.


Discovery and naming

Fossils of ''Syntomiprosopus'' were discovered at a locality in the Downs' Quarry located in eastern Arizona, just above the "''
Placerias ''Placerias'' (meaning 'broad body') is an extinct genus of dicynodonts that lived during the Carnian to the Norian age of the Triassic Period (230–220 million years ago). ''Placerias'' belongs to a group of dicynodonts called Kannemeyeriiforme ...
'' Quarry" and just below the principal horizon of the Downs' Quarry. The fossils were recovered during joint excavations by the
North Carolina Museum of Natural Sciences The North Carolina Museum of Natural Sciences (NCMNS) is the largest museum of its kind in the Southeastern United States. It is the oldest established museum in North Carolina, located in Raleigh. In 2013, it had about 1.2 million visitors, and i ...
(NCSM) and
Appalachian State University Appalachian State University (; Appalachian, App State, App, or ASU) is a public university in Boone, North Carolina. It was founded as a teachers college in 1899 by brothers B. B. and D. D. Dougherty and the latter's wife, Lillie Shull Dough ...
(ASU) between 2013 and 2015 as part of a series of excavations that had begun in 2010. The quarry is part of the expansive Late Triassic
Chinle Formation The Chinle Formation is an Upper Triassic continental geological formation of fluvial, lacustrine, and palustrine to eolian deposits spread across the U.S. states of Nevada, Utah, northern Arizona, western New Mexico, and western Colorado. In Ne ...
, and has been
radiometrically dated Radiometric dating, radioactive dating or radioisotope dating is a technique which is used to date materials such as rocks or carbon, in which trace radioactive impurities were selectively incorporated when they were formed. The method compares t ...
to have been deposited at a maximum age of approximately 219.39 ± 0.16 million years old. The
holotype specimen 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 ...
(NCSM 29059–29060) consists of two associated pieces of a complete right
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 ...
, along with the
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). Of ...
s NCSM 26729 (an
articular The articular bone is part of the lower jaw of most vertebrates, including most jawed fish, amphibians, birds and various kinds of reptiles, as well as ancestral mammals. Anatomy In most vertebrates, the articular bone is connected to two oth ...
complex from the rear of the left jaw), NCSM 26730 (the front of a left mandible), NCSM 27677 (the middle portion of a left mandible), NCSM 27678 (another left articular complex) and NCSM 29061 (a right articular complex). These belong to at least two, but possibly up to four individuals. Part of the back of a skull and braincase (NCSM 27679) found associated with the jaw material may also belong to ''Syntomiprosopus'', as well as some
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 ...
e (NCSM 27991 and NCSM 27992), but they cannot be definitively referred to ''Syntomiprosopus'' on anatomical grounds. However, because there are no other similarly sized vertebrates in the fossil bed that it can be referred to, has identical preservation, and that it does not match any previously known animal from the Downs' Quarry, Heckert and colleagues were confident in referring the skull to ''Syntomiprosopus''. Heckert and colleagues named ''Syntomiprosopus'' from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
prefix ''syntomi-'' ("short") and ''prosopus'' ("face") to refer to its characteristically short jaws. The specific name ''sucherorum'' is in honour of Scott ("Major") and Karen Sucher, who had voluntarily spent 22 years supporting and aiding Heckert and his colleagues, including participating in the excavations at Downs' Quarry that discovered the remains of ''Syntomiprosopus''.


Description

''Syntomiprosopus'' is only known by bones of the lower jaw and potentially part of 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 ...
. However, the lower jaw is distinct amongst Late Triassic reptiles, being short and almost as broad as they are long, meeting with a U-shaped curve at their tips. The lower jaw is robust, approximately 2.5x as long as high, and is twice as deep at its end as the tips. They have a coarse and roughly textured surface with irregularly placed pits and grooves without any clear pattern. ''Syntomiprosopus'' has very few teeth, with the smallest specimen having only eight active tooth positions and the largest specimen only four. The teeth are divided into a set 2—4 front teeth that are procumbent (angled forwards), a large canine-like (caniniform) tooth, followed by up to 3 small postcanine teeth, the smallest set in the toothrow. In larger specimens, the tooth sockets for these postcanines are resorbed into the jaw bone, and the teeth are lost. The teeth themselves are asymmetrical, with a flat outer surface and bulbous inner surface, and appear to have large denticles (serrations) along their edges.


Classification

''Syntomiprosopus'' is evidenced to be an archosauriform, based on the presence of the mandibular fenestra and serrated teeth. However, no characteristics are identifiable in the lower jaw to identify its affinities beyond Archosauriformes. Notably, the rugose mandibular symphysis, large caniniform tooth, short prearticular, and the angular that is well exposed on the internal side of the jaw are all characteristics variably found in crocodylomorphs, although the first two traits are also found elsewhere in archosauriforms. The braincase and back of the skull, if correctly referred to ''Syntomiprosopus'', may further indicate affinities to early crocodylomorphs. Crocodylomorph characteristics of this skull include a sagittal crest along the midline (formed by fused, raised
parietal bone The parietal bones () are two bones in the Human skull, skull which, when joined at a fibrous joint, form the sides and roof of the Human skull, cranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four borders, an ...
s), the rear edge of the parietals diverge at almost 90° from the midline of the skull, a contact between the laterosphenoid and
parabasisphenoid The parasphenoid is a bone which can be found in the cranium of many vertebrates. It is an unpaired dermal bone which lies at the midline of the roof of the mouth. In many reptiles (including birds), it fuses to the endochondral (cartilage-derived ...
of the braincase, and a possible contact between the prootic bone of the
middle ear The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear). The mammalian middle ear contains three ossicles, which transfer the vibrations of the eardrum into waves in the ...
and
quadrate bone The quadrate bone is a skull bone in most tetrapods, including amphibians, sauropsids (reptiles, birds), and early synapsids. In most tetrapods, the quadrate bone connects to the quadratojugal and squamosal bones in the skull, and forms upper ...
on the external surface of the skull. However, the partial skull also lacks characteristics found in more derived crocodylomorphs (such as deep hollow on the underside of the
basioccipital The basilar part of the occipital bone (also basioccipital) extends forward and upward from the foramen magnum, and presents in front an area more or less quadrilateral in outline. In the young skull this area is rough and uneven, and is joined t ...
, or on the
parabasisphenoid The parasphenoid is a bone which can be found in the cranium of many vertebrates. It is an unpaired dermal bone which lies at the midline of the roof of the mouth. In many reptiles (including birds), it fuses to the endochondral (cartilage-derived ...
)) and preserves features lost in more derived crocodylomorphs (such as the entrances for the
internal carotid arteries The internal carotid artery (Latin: arteria carotis interna) is an artery in the neck which supplies the anterior circulation of the brain. In human anatomy, the internal and external carotids arise from the common carotid arteries, where these b ...
being placed high on the skull and modifications to the middle ear). This combination of traits indicate that if ''Syntomiprosopus'' is indeed a crocodylomorph, it must be a relatively basal early-diverging species, less derived than ''
Sphenosuchus ''Sphenosuchus'' ("wedge crocodile") is an extinct genus of crocodylomorph from the Early Jurassic Elliot Formation of South Africa, discovered and described early in the 20th century. The skull is preserved very well but other than elements of ...
''.


Palaeobiology

The smaller specimen of ''Syntomiprosopus'' preserves more open tooth sockets, four in front, a canine-like tooth, and three postcanine teeth behind. The larger specimen only preserves the three front teeth and the caniniform, with the remaining tooth sockets behind being either fully or partially resorbed into the jaw bone. This suggests that ''Syntomiprosopus'' lost these teeth as it matured, losing close to half of its dentition. Some archosaurs are known to decrease their tooth counts as they mature (e.g.
tyrannosaurids Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family (biology), family of coelurosaurian Theropoda, theropod dinosaurs that comprises two subfamilies containing up to thirteen genus, genera, including the eponymous ''Tyrannos ...
and possibly modern
crocodylians Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period (Cenomanian stage) and are the closest living ...
), however, only ''
Limusaurus ''Limusaurus'' is a genus of theropod dinosaur that lived in what is now China during the Late Jurassic, around 161 to 157 million years ago. The type and only species ''Limusaurus inextricabilis'' was described in 2009 from specimens ...
'' shows a similarly drastic loss in teeth between juveniles and adults (whereby juveniles have sharp, pointed teeth that are lost for a beak in adults).


Palaeoecology

The only other fossils recovered from the same layers as ''Syntomiprosopus'' are of indeterminate
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 equivalent g ...
. However, the layers immediately below and above contain fossils of the herbivorous armoured
aetosaurs 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 ...
''
Desmatosuchus ''Desmatosuchus'' (, from Greek δεσμός ''desmos'' 'link' + σοῦχος ''soûkhos'' 'crocodile') is an extinct genus of archosaur belonging to the Order Aetosauria. It lived during the Late Triassic. Description ''Desmatosuchus'' was a ...
'', ''
Calyptosuchus ''Calyptosuchus'' (meaning "covered crocodile") is an extinct genus of aetosaur from the Late Triassic of North America. Like other aetosaurs, it was heavily armored and had a pig-like snout used to uproot plants. Description ''Calyptosuchus'' ...
'' and ''
Tecovasuchus ''Tecovasuchus'' is an extinct genus of aetosaur. It is known primarily from osteoderms found from the Tecovas Formation in Texas, which is Late Triassic in age, dating back to the lower Norian. Material is also known from several other localitie ...
'', as well as a
dicynodont Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivorous animals with a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, typicall ...
(likely ''Placerias'') and an unidentified
allokotosaur Allokotosauria is a clade of early archosauromorph reptiles from the Middle Triassic, Middle to Late Triassic known from Asia, Africa, North America and Europe. Allokotosauria was first described and named when a new monophyletic grouping of spec ...
.


References

{{Archosauromorpha, B., state=autocollapse Prehistoric archosauriforms Prehistoric crocodylomorphs Prehistoric reptile genera Norian genera Late Triassic reptiles of North America Fossil taxa described in 2021 Taxa named by Sterling Nesbitt