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''Calsoyasuchus'' (meaning " r. KyrilCalsoyas'
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large, semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term "crocodile" is sometimes used more loosely to include ...
") is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
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. Extinct crocodylomorphs were considerably mor ...
that lived in the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic� ...
. Its fossilized remains were found in the
Sinemurian In the geologic timescale, the Sinemurian is an age (geology), age and stage (stratigraphy), stage in the Early Jurassic, Early or Lower Jurassic epoch (geology), Epoch or series (stratigraphy), Series. It spans the time between 199.5 ±0.3 annu ...
-
Pliensbachian The Pliensbachian is an age of the geologic timescale and stage in the stratigraphic column. It is part of the Early or Lower Jurassic Epoch or Series and spans the time between 192.9 ±0.3 Ma and 184.2 ±0.3 Ma (million years ago). The Plie ...
-age
Kayenta Formation The Kayenta Formation is a geological formation in the Glen Canyon Group that is spread across the Colorado Plateau area of the United States, including northern Arizona, northwest Colorado, Nevada, and Utah. Originally suggested as being Sinemur ...
on
Navajo Nation The Navajo Nation (), also known as Navajoland, is an Indian reservation of Navajo people in the United States. It occupies portions of northeastern Arizona, northwestern New Mexico, and southeastern Utah. The seat of government is located in ...
land in
Coconino County Coconino County is a County (United States), county in the North Central Arizona, North-Central part of the U.S. state of Arizona. Its population was 145,101 at the 2020 United States census, 2020 census. The county seat is Flagstaff, Arizon ...
,
Arizona Arizona is a U.S. state, state in the Southwestern United States, Southwestern region of the United States, sharing the Four Corners region of the western United States with Colorado, New Mexico, and Utah. It also borders Nevada to the nort ...
,
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
. Formally described as ''C. valliceps'', it is known from a single incomplete skull which is unusually derived for such an early crocodile relative. This genus was described in 2002 by Ronald Tykoski and colleagues; the specific name means "valley head" and refers to a deep groove along the midline of the
nasal bone 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 ...
s and
frontal bone In the human skull, the frontal bone or sincipital bone is an unpaired bone which consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bo ...
s. It has often been interpreted as the earliest diverging member of Goniopholididae, but other studies have recovered it in various other positions.


Description

The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
skull ( TMM 43631-1) that would be named ''Calsoyasuchus'' was discovered in 1997 by members of an expedition composed of crews from
Texas Memorial Museum Texas Science & Natural History Museum is located on the campus of The University of Texas at Austin in Austin, Texas, U.S. It opened as Texas Memorial Museum during preparations for the Texas Centennial Exposition held in 1936. The museum's foc ...
of the
University of Texas at Austin The University of Texas at Austin (UT Austin, UT, or Texas) is a public university, public research university in Austin, Texas, United States. Founded in 1883, it is the flagship institution of the University of Texas System. With 53,082 stud ...
, the
Museum of Comparative Zoology The Museum of Comparative Zoology (formally the Agassiz Museum of Comparative Zoology and often abbreviated to MCZ) is a zoology museum located on the grounds of Harvard University in Cambridge, Massachusetts. It is one of three natural-history r ...
at
Harvard University Harvard University is a Private university, private Ivy League research university in Cambridge, Massachusetts, United States. Founded in 1636 and named for its first benefactor, the History of the Puritans in North America, Puritan clergyma ...
, and the Seba Dalkai Navajo Nation School. It was found in the middle third of the
silt Silt is granular material of a size between sand and clay and composed mostly of broken grains of quartz. Silt may occur as a soil (often mixed with sand or clay) or as sediment mixed in suspension (chemistry), suspension with water. Silt usually ...
y
facies In geology, a facies ( , ; same pronunciation and spelling in the plural) is a body of rock with distinctive characteristics. The characteristics can be any observable attribute of rocks (such as their overall appearance, composition, or con ...
of the Kayenta Formation, near the Adeii Eechii Cliffs. The skull is missing the lower jaws, part of the
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sep ...
, most of the suspensorium (the bones that make up the region where the upper and lower jaws articulate), and the
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 lies over the occipital lobes of the ...
and
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, brain-pan, or brainbox, 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 calv ...
. Sutures between the skull bones are mostly fused. As preserved, it is about 38.0 centimeters (15.0 in) long, making its owner a moderately sized animal. The skull was long, low, and curved so that both extremities were higher than the middle. 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 mammals h ...
bones that formed the end of the snout were enlarged to form a wide tip; there were at least four teeth in the right premaxilla and five in the left. The left
maxilla In vertebrates, the maxilla (: maxillae ) 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 two maxil ...
(main tooth-bearing bone of the upper jaw) is more complete than the right, and had at least 29 teeth. There was a deep groove along the midline of the nasals and the frontals; the frontals were fused into a single bone, as is seen in other adult mesoeucrocodylians. Unlike derived neosuchians, it had external
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 Archosauriformes, archosauriforms, first appearing during the Triassic Period. Among Extant ...
e. Tykoski and colleagues subjected the skull to
CT scanning A computed tomography scan (CT scan), formerly called computed axial tomography scan (CAT scan), is a medical imaging technique used to obtain detailed internal images of the body. The personnel that perform CT scans are called radiographers or ...
, which revealed internal cavities and air passages, and showed that it had a double-walled secondary palate similar to that of true crocodylians, and similar pneumatic cavities as well.


Phylogeny

Tykoski and colleagues performed a cladistic phylogenetic analysis with their new
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
, and found that it grouped with ''
Goniopholis ''Goniopholis'' (meaning "angled scale") is an extinct genus of goniopholidid crocodyliform that lived in Europe and North America during the Late Jurassic and Early Cretaceous. Like other goniopholidids, it resembled living crocodilians, and pro ...
'', '' Sunosuchus'', and, most closely, with '' Eutretauranosuchus'' in a weakly supported
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
, Goniopholididae. They noted that the skull of ''Calsoyasuchus'' is very similar to some goniopholid skulls from the younger,
Upper Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
Morrison Formation The Morrison Formation is a distinctive sequence of Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandstone, siltston ...
. ''Calsoyasuchus'' pushes back the earliest occurrence of goniopholids from the Late Jurassic into the Early Jurassic, and not only helps to bridge a temporal gap between groups of crocodyliforms, but also a morphological gap. It also implies that some groups of crocodyliforms have long undiscovered histories. In 2011, ''Calsoyasuchus'' was recovered as the basalmost goniopholidid and the sister taxon to all other goniopholidids. Its placement as a goniopholid is not uncontested. A cladistic analysis of Crocodylomorpha published by Wilberg et al. (2019) in their discussion of crocodylomorph paleobiology recovers ''Calsoyasuchus'' as a close relative of the basal mesoeucrocodyliform '' Hsisosuchus''. A 2022 analysis found it to be the sister group of Thalattosuchia.


Paleoecology

During the
Sinemurian In the geologic timescale, the Sinemurian is an age (geology), age and stage (stratigraphy), stage in the Early Jurassic, Early or Lower Jurassic epoch (geology), Epoch or series (stratigraphy), Series. It spans the time between 199.5 ±0.3 annu ...
and
Pliensbachian The Pliensbachian is an age of the geologic timescale and stage in the stratigraphic column. It is part of the Early or Lower Jurassic Epoch or Series and spans the time between 192.9 ±0.3 Ma and 184.2 ±0.3 Ma (million years ago). The Plie ...
ages of the Early Jurassic, the Kayenta Formation had a diverse
fauna Fauna (: faunae or faunas) is all of the animal life present in a particular region or time. The corresponding terms for plants and fungi are ''flora'' and '' funga'', respectively. Flora, fauna, funga and other forms of life are collectively ...
, with the remains of
caecilian Caecilians (; ) are a group of limbless, vermiform (worm-shaped) or serpentine (snake-shaped) amphibians with small or sometimes nonexistent eyes. They mostly live hidden in soil or in streambeds, and this cryptic lifestyle renders caecilians ...
s,
frog A frog is any member of a diverse and largely semiaquatic group of short-bodied, tailless amphibian vertebrates composing the order (biology), order Anura (coming from the Ancient Greek , literally 'without tail'). Frog species with rough ski ...
s,
turtle Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
s, at least five other taxa of crocodylomorphs,
pterosaur Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
s,
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
,
sauropodomorph Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lo ...
, and
ornithischia Ornithischia () is an extinct clade of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek ...
n
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s, and early relatives of mammals ( tritylodontids and
morganucodon ''Morganucodon'' ("Glamorgan tooth") is an early mammaliaform genus that lived from the Late Triassic to the Middle Jurassic. It first appeared about 205 million years ago. Unlike many other early mammaliaforms, ''Morganucodon'' is well represent ...
tids).


References


External links


Digimorph page
featuring various views and movies of the CT scans of the skull {{Taxonbar, from=Q2908523 Crocodyliformes Prehistoric pseudosuchian genera Early Jurassic reptiles of North America Early Jurassic crocodylomorphs Kayenta Formation Fossil taxa described in 2002