Hesperosuchus
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''Hesperosuchus'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
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 ...
reptile that contains a single species, ''Hesperosuchus agilis''. Remains of this
pseudosuchian Pseudosuchia, from Ancient Greek ψεύδος (''pseúdos)'', meaning "false", and σούχος (''soúkhos''), meaning "crocodile" is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely relat ...
have been found in
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 a ...
(
Carnian The Carnian (less commonly, Karnian) is the lowermost stage (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227.3 ...
) strata from
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 ...
and
New Mexico New Mexico is a state in the Southwestern United States, Southwestern region of the United States. It is one of the Mountain States of the southern Rocky Mountains, sharing the Four Corners region with Utah, Colorado, and Arizona. It also ...
.Colbert, E. H. 1952. A pseudosuchian reptile from Arizona. Bulletin of the American Museum of Natural History 99:561–592. Because of similarities in skull and neck anatomy and the presence of hollow bones ''Hesperosuchus'' was formerly thought to be an ancestor of later
carnosauria Carnosauria is an extinct group of carnivorous theropod dinosaurs that lived during the Jurassic and Cretaceous periods. While Carnosauria was historically considered largely synonymous with Allosauroidea, some recent studies have revived Ca ...
n dinosaurs, but based on more recent findings and research it is now known to be more closely related to
crocodilia Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
ns rather than dinosaurs.Brinkman, D. 1981. The origin of the crocodiloid tarsi and the interrelationships of thecodontian archosaurs. Breviora 464:1–23.Benton, M. J. and J. M. Clark. 1988. Archosaur phylogeny and the relationships of the Crocodylia. pp. 295–338. In M. J. Benton (ed.). The Phylogeny and Classification of the Tetrapods Vol. 1.Amphibians, Reptiles, Birds. Clarendon Press, Oxford.


Description


Stance and limbs

At only about in length, ''Hesperosuchus'' is a relatively small and lightly built pseudosuchian. It is very closely comparable to the different genera, ''
Ornithosuchus woodwardi ''Ornithosuchus'' (from , "bird" and , "crocodile") is an extinct genus of pseudosuchians from the Late Triassic (Carnian) Lossiemouth Sandstone of Scotland. It was originally thought to be the ancestor to the carnosaurian dinosaurs (such as ''Al ...
'' and '' Saltoposuchus longipes'',Baczko, M. B. von & Ezcurra, M. D. 2013. Ornithosuchidae: a group of Triassic archosaurs with a unique ankle joint. Geological Society, London, Special Publications 379(1), 187–202. which are both pseudosuchians as well. The hind limbs of ''Hesperosuchus'' are large and strong, while the forelimbs were smaller and much more slender. This observation lead to the hypothesis that ''Hesperosuchus'' was a
bipedal Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' ...
animal. Comparing the hind limbs of ''Saltoposuchus'' with those of ''Hesperosuchus'', they are evenly as large and strong. The length of the extended hind limb in both genera are approximately equal to the length of the presacral vertebrae. ''Saltoposuchus'' was described by von Huene, and portrayed as a facultative
quadruped Quadrupedalism is a form of locomotion in which animals have four legs that are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four legs is said to be a quadruped (fr ...
. It is believed that ''Hesperosuchus'' also practiced both bipedialism and quadrupedalism. Though it is believed to be more often on two feet as the long slender hands looked as if it was adapted for grasping, which may have been useful for food gathering, digging or defense. Five digits were found on both the hind limbs as well as the forelimbs. In order to counterbalance the weight of its body, ''Hesperosuchus'' is inferred to have had a relatively long tail. Since the caudal vertebrae aren't completely restored, it is inferred based on similar archosaurians, that the tail contained somewhere around 45 caudal vertebrae. The strong hind limbs and overall light weight made ''Hesperosuchus'' very quick and able to move rapidly. This advantage of speed allowed for it to catch small prey and escape from larger predators.


Skull

The skull of ''Hesperosuchus'' was only partially preserved and is missing many segments. The
mandible In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
and the skull of the specimen found was very poorly preserved, but there is just enough bone present to provide for an indication of what the basic structure of the jaw and skull may have looked like. It was found that the skull very closely resembled that of ''Ornithosuchus''. In the fronto-parietal regions of the skull, along with a flat cranial roof, marked depressions were found in the frontal and post orbital bones, in front of and lateral to the
supratemporal fenestrae Temporal fenestrae are openings in the Temple (anatomy), temporal region of the skull of some Amniote, amniotes, behind the Orbit (anatomy), orbit (eye socket). These openings have historically been used to track the evolution and affinities of re ...
. Fragments of the left
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 ...
and
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 ...
were found to have sockets for nine teeth, with four being in the premaxilla. The first premaxillary teeth start out small in size and progressively get larger with the fourth tooth being clearly enlarged. This was compared to the skull of ''Ornithosuchus'', which is defined with characteristics of two enlarged teeth in this similar area; the first two maxillary teeth. The teeth of ''Hesperosuchus'' are serrated in both the posterior and anterior edges which supports the fact that ''Hesperosuchus'' was a meat eating animal. With the two fragments of the jaws found, only 14 teeth in total were reported with five in the posterior fragment and nine in the anterior one. The basioccipital region is defined as typically
archosaur Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...
ian, with a rounded condyle, a rather elongated surface above it for the
medulla oblongata The medulla oblongata or simply medulla is a long stem-like structure which makes up the lower part of the brainstem. It is anterior and partially inferior to the cerebellum. It is a cone-shaped neuronal mass responsible for autonomic (involun ...
, and an extended ventral plate. These basioccipital characteristics are seen in extinct archosaurians such as primitive theropod dinosaurs as well as seen in crocodylians. It can be seen that the skull would have been relatively large which was compared to carnosaurian dinosaurs, which too had fairly large skulls. It can be said that both these groups were active and carnivorous as large skulls allow for wide gaping jaws to catch and attack prey. Such large skulls need to be light as a large antorbital opening is present and clearly shown in ''Hesperosuchus''.


Soft tissue reconstruction

Ziphodont dentition and
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 ...
-like pattern of linearly arranged jaw foramina of ''Hesperosuchus'' suggest a possible existence of lips that protected its teeth from the outside.


Discovery

''Hesperosuchus'' was discovered in upper Triassic rocks of Northern Arizona by Llewellyn I. Price, William B. Hayden, and Barnum Brown in the fall of 1929 and the summer of 1930. The specimen was then taken to a museum for Otto Falkenbach to carefully and precisely put together. Many different illustrations of the bones were done by Sydney Prentice from the Carnegie Museum of Pittsburgh. In addition, models and figures were also made by John LeGrand Lois Darling from the Museum Illustrators Corps. The exact location where the specimen was found is an area southeast of
Cameron, Arizona Cameron () is a census-designated place (CDP) in Coconino County, Arizona, United States, on the Navajo Nation. The population was 885 at the 2010 census. Most of the town's economy is tourist food and craft stalls, restaurants, and other serv ...
, close to the old Tanner Crossing of the
Little Colorado River The Little Colorado River () is a tributary of the Colorado River in the U.S. state of Arizona, providing the principal drainage from the Painted Desert region. Together with its major tributary, the Puerco River, it drains an area of about in ...
.Lucas, S.G., 1993, The Chinle Group: revised stratigraphy and chronology of Upper Triassic nonmarine strata in the western United States: Museum of Northern Arizona, Bulletin 59, p. 27-50. This area, in particular, is a very prevalent for finding many Triassic
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s.Blakey, R. C. and R. Gubitosa. 1983, Late Triassic paleogeography and depositional history of the Chinle Formation, southern Utah and northern Arizona: ''in'' Reynolds, M.W., and Dolly, E.D., eds., Mesozoic paleogeography of west-central U.S.: Society of Economic Paleontologists and Mineralogists, Rocky Mountain Section, Denver, p. 57–76. This area is about on top of the Moen kopi formation, in the portion of the
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 ...
Irmis, R.B., 2005, The vertebrate fauna of the Upper Triassic Chinle Formation in northern Arizona: Mesa Southwest Museum, Bulletin 9, p. 63- 88. where the Little Colorado River flows through a canyon. Along with ''Hesperosuchus'', many other specimens were found in the same general area. There were many
ganoid scale A fish scale is a small rigid plate that grows out of the skin of a fish. The skin of most jawed fishes is covered with these protective scales, which can also provide effective camouflage through the use of reflection and colouration, as w ...
s believed to have belonged to Triassic freshwater
holostean Holostei is a group of ray-finned bony fish. It is divided into two major clades, the Halecomorphi, represented by the single living genus, '' Amia'' with two species, the bowfins (''Amia calva'' and ''Amia ocellicauda''), as well as the Ginglymo ...
fish, several
phytosaur Phytosaurs (Φυτόσαυροι in Greek, meaning 'plant lizard') are an extinct group of large, mostly semiaquatic Late Triassic archosauriform or basal archosaurian reptiles. Phytosaurs belong to the order Phytosauria and are sometimes ref ...
teeth, and many small
stereospondyl The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecologi ...
vertebrae. In addition, a large number of small teeth were found, some which definitely belonged to ''Hesperosuchus'', and some belonging to
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
related animals. It is hypothesized that these other teeth may have belonged to animals that ''Hesperosuchus'' may have preyed upon. ''Hesperosuchus'' was a contemporary of ''
Coelophysis ''Coelophysis'' ( Traditional English pronunciation of Latin, traditionally; or , as heard more commonly in recent decades) is a genus of coelophysid Theropoda, theropod dinosaur that lived Approximation, approximately 215 to 201.4 million y ...
'', a primitive predatory
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 ...
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 ...
. ''Coelophysis'' was long thought to have been a cannibal, based on the presence of putative juvenile ''Coelophysis'' bones in the gut regions of a few adults. However, in some of these cases, it was later found that the "juvenile ''Coelophysis''" bones were actually those of a ''Hesperosuchus'' (or something very similar) instead. In 2001, a new specimen of ''H. agilis'' was described from the famous
Ghost Ranch Ghost Ranch is a retreat and education center in Rio Arriba County in north central New Mexico, United States. It is about 65 miles northwest of Santa Fe and 14 miles from Abiquiu, the nearest community. In the later 20th century, it was the ...
fossil site in New Mexico.


Paleoecology

''Hesperosuchus'' was a
terrestrial Terrestrial refers to things related to land or the planet Earth, as opposed to extraterrestrial. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on o ...
animal, where its speed and ability to run fast is the most advantageous as a fitness trait. Northern Arizona's landscape during the Triassic period was surrounded by numerous bodies of water like lakes and streams. This supports that ''Hesperosuchus'' likely lived close to water although being a full on land-dwelling animal. The ganoid scales found in the general area where ''Hesperosuchus'' was found belong to freshwater fish of the Triassic period, belonging to the genus ''
Semionotus ''Semionotus'' (from , 'mark' and , 'back') is an extinct genus of ray-finned fish found throughout Northern Pangaea (North America and Europe) during the late Triassic, becoming extinct in the Early Jurassic The Jurassic ( ) is a Geological ...
'' or ''
Lepidotes ''Lepidotes'' (from , 'covered with scales') (previously known as ''Lepidotus'') is an extinct genus of Mesozoic ray-finned fish. It has long been considered a wastebasket taxon, characterised by "general features, such as thick rhomboid scales ...
'', which lived in shallow lakes and streams. The phytosaur teeth and small stereospondyl vertebrae found near ''Hesperosuchus'' support the presence of lakes or streams crossing a flood plain. Also the many small teeth found, of which some belong to amphibians of the Triassic period, supports the occupying of near water habitats.


See also

*
Pseudosuchia Pseudosuchia, from Ancient Greek ψεύδος (''pseúdos)'', meaning "false", and σούχος (''soúkhos''), meaning "crocodile" is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely relat ...
*
Ornithosuchia 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 ...
*
Crocodilia Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
*
Archosaur Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...
ia *
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
*
Sphenosuchia Sphenosuchia is a suborder of basal crocodylomorphs that first appeared in the Triassic and occurred into the Middle Jurassic. Most were small, gracile animals with an erect limb posture. They are now thought to be ancestral to crocodyliforms ...
*
Euparkeria ''Euparkeria'' (; meaning "Parker's good animal", named in honor of W. K. Parker) is an Extinction, extinct genus of archosauriform reptile from the Triassic of South Africa. ''Euparkeria'' is close to the ancestry of Archosauria, the reptile gro ...


References

{{Taxonbar, from=Q2697976 Terrestrial crocodylomorphs Triassic crocodylomorpha Monotypic prehistoric reptile genera Late Triassic archosaurs of North America Chinle fauna Prehistoric pseudosuchian genera