Desmatosuchus Spurensis
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''Desmatosuchus'' (, from
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 ...
δεσμός ''desmos'' 'link' + σοῦχος ''soûkhos'' 'crocodile') 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
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 ...
belonging to the Order
Aetosauria 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 m ...
. It lived during 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 ...
.


Description

''Desmatosuchus'' was a large quadrupedal reptile upwards of to in lengthvon Baczko, M. B., Desojo, J. B., Gower, D. J., Ridgely, R., Bona, P., & Witmer, L. M. (2021)
New digital braincase endocasts of two species of Desmatosuchus and neurocranial diversity within Aetosauria (Archosauria: Pseudosuchia)
The Anatomical Record, 1–20. https://doi.org/10.1002/ar.24798
and in weight. Its
vertebral column The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a vertebrate in which the notochord (a flexible rod of uniform composition) found in all chordata, ...
had amphicoelus
centra Centra is a convenience shop chain that operates throughout Ireland. The chain operates as a symbol group owned by Musgrave Group, the food wholesaler, meaning the stores are all owned by individual franchisees. The chain has three different ...
and 3 sacral vertebrae. This archosaur's most distinguishing anatomical characteristics were its
scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eithe ...
e which possessed large acromion processes commonly referred to as "shoulder spikes". The forelimbs were much shorter than the hindlimbs, with
humeri The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a round ...
less than two-thirds the length of the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
s. The
pelvic girdle The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The p ...
consisted of a long pubis with a strong
symphysis A symphysis (, pl. symphyses) is a fibrocartilaginous fusion between two bones. It is a type of cartilaginous joint, specifically a secondary cartilaginous joint. # A symphysis is an amphiarthrosis, a slightly movable joint. # A growing together ...
in the middle, a plate-like
ischium The ischium () form ...
, a highly recurved ilium, and a deep, imperforate
acetabulum The acetabulum (), also called the cotyloid cavity, is a concave surface of the pelvis. The head of the femur meets with the pelvis at the acetabulum, forming the hip joint. Structure There are three bones of the ''os coxae'' (hip bone) that c ...
. The femurs were relatively long and straight, the ankles
crurotarsal A crurotarsal joint is one that’s situated between the bones of ''crus'', i.e. shin (tibia and fibula) and the proximal tarsal bones, i.e. astragalus and calcaneum. The ankle joint of therian mammals (marsupials and placentals) is a crurotarsal ...
, with calcaneal tubers that gave it large heels. Its skull was relatively small, on average about 37 centimeters long, 18 centimeters wide, and 15 centimeters high. The braincase was very firmly fused with the skull roof and palate. It had slender, forked
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 ...
e that turned up and expanded in the front, creating a shovel-like structure. ''Desmatosuchus'' is unique among aetosaurs in that its species are the only known aetosaurs that lacked teeth on their premaxillae. Their premaxillae fit loosely together with their
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 ...
e, indicating flexibility at that joint. Their maxilla contained 10 to 12 teeth. ''Desmatosuchus'' also had very thin
vomer The vomer (; lat, vomer, lit=ploughshare) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right maxill ...
s, which bounded the medial side of the internal nares. These internal nares were relatively large, roughly half the length of the entire palate. The lower jaw typically carried 5 or 6 teeth, and had a toothless beak on the end. The
dentary 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 ...
was about half the length of the lower jaw, with the front portion being toothless and covered by a horny sheath. Behind the dentary was a moderately large mandibular
fenestra A fenestra (fenestration; plural fenestrae or fenestrations) is any small opening or pore, commonly used as a term in the biological sciences. It is the Latin word for "window", and is used in various fields to describe a pore in an anatomical st ...
. Individuals of ''Desmatosuchus'' were heavily armored. The
carapace A carapace is a Dorsum (biology), dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tor ...
was made up of two rows of median
scute A scute or scutum (Latin: ''scutum''; plural: ''scuta'' "shield") is a bony external plate or scale overlaid with horn, as on the shell of a turtle, the skin of crocodilians, and the feet of birds. The term is also used to describe the anterior po ...
s surrounded by two more rows of lateral scutes. The lateral scutes had well-developed spine-like processes which pointed out laterally and dorso-posteriorly. There were typically five rows of spines, increasing in size anteriorly. The front spine was much larger, around 28 centimeters long, and was recurved. The fourth spine varies in length in each specimen, but remains shorter than the fifth in all of them. ''Desmatosuchus'' are the only aetosaurs known to have possessed spines like these.


Discovery and classification

The first ''Desmatosuchus'' discovery occurred in the late 19th century when E.D. Cope classified armor from the
Dockum Group The Dockum is a Late Triassic (approximately late Carnian through Rhaetian, or 223–200 Ma) geologic group found primarily on the Llano Estacado of western Texas and eastern New Mexico with minor exposures in southwestern Kansas, eastern Color ...
in
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
, USA, as the new species ''Episcoposaurus haplocerus''. Case later classified a partial skeleton found in the
Tecovas Formation The Tecovas Formation is a geological formation in Texas.Weishampel, David B; et al. (2004). "Dinosaur distribution (Late Triassic, North America)." In: Weishampel, David B.; Dodson, Peter; and Osmólska, Halszka (eds.): The Dinosauria, 2nd, Berk ...
as ''Desmatosuchus spurensis''. Since the localities of Cope and Case were only a few kilometers apart, the two taxa were synonymized into ''Desmatosuchus haplocerus'', the initial type species of the genus. A revision of ''Desmatosuchus'' by Parker (2008) found the
lectotype In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes the ...
of ''Episcoposaurus haplocerus'' to be referable to ''Desmatosuchus'' but indeterminate at the species level. Therefore, ''E. haplocerus'' was considered to be a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium'' it may be impossible to determine whether a s ...
'' and ''D. spurensis'' was named 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 ...
of the genus. Two species were accepted as valid: ''D. spurensis'' and ''D. smalli'', named after Brian J. Small for his contribution to the study of this genus. ''Desmatosuchus chamaensis'' is recognized as a distinct genus, but there is some dispute about whether the name '' Heliocanthus'' or '' Rioarribasuchus'' applies. The following
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 ...
is simplified, after an analysis presented by Julia B. Desojo, Martin D. Ezcurra and Edio E. Kischlat (2012).


Paleobiology

Bones and armor pieces of ''Desmatosuchus'' are abundant in the Dockum formation, Chinle formation, and Post quarry, indicating that they were widespread and abundant during the Late Triassic. It is possible that ''Desmatosuchus'' traveled in herds or family units. This is evidenced by several findings of multiple ''Desmatosuchus'' skeletons in relatively small areas. ''Desmatosuchus'' had blunt, bulbous, slightly recurved teeth. Furthermore, they are believed to have had
homodont In anatomy, a heterodont (from Greek, meaning 'different teeth') is an animal which possesses more than a single tooth morphology. In vertebrates, heterodont pertains to animals where teeth are differentiated into different forms. For example ...
dentition. This, combined with its shovel like snout, indicate that ''Desmatosuchus'' fed by digging up soft vegetation. This method of feeding is further evidenced by its toothless premaxilla and dentary tip, which were covered in horny sheaths. These sheaths protected the bones and could be used for cutting or holding objects. It is believed that ''Desmatosuchus'' dug for food in the soft mud near bodies of water due to the abundance of lakes and rivers in the Dockum area and the fact that ''Desmatosuchus'' scutes are often found among parts of other reptiles that are known to have fed along waterways. It is unknown whether or not ''Desmatosuchus'' replaced their teeth and, if so, how. The low number of ''Desmatosuchus'' teeth that have been discovered indicates that they were only held in place by soft tissue connections. The jaw articulation point is below the tooth line, holding its upper and lower tooth rows parallel while biting in a way that is reminiscent of
ornithischian Ornithischia () is an extinct order 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 st ...
dinosaurs. The armor and spikes of ''Desmatosuchus'' were its only ways to defend itself from predators. The lateral spike rows showed variation in size among individuals, especially the second most anterior spike. This spike was always shorter than the one in front of it, but to what extent varied drastically. This variation may indicate
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
. It has also been hypothesized as a form of sexual display. Aside from this armor, ''Desmatosuchus'' was defenseless from attacks from carnivores. Several ''Desmatosuchus'' bones have been found amongst skeletons of ''
Postosuchus ''Postosuchus'', meaning "Crocodile from Post", is an extinct genus of rauisuchid reptiles comprising two species, ''P. kirkpatricki'' and ''P. alisonae'', that lived in what is now North America during the Late Triassic. ''Postosuchus'' is a ...
'', indicating predation by ''Postosuchus''. The herd nature of ''Desmatosuchus'' apparently did little to discourage predators, as ''Postosuchus'' along with several other Late Triassic carnivores also traveled in groups. Most thecodonts of the Late Triassic lacked certain pelvic features that aided locomotion, such as a deep acetabulum or a crest over the acetabulum. This, in spite of their upright posture, rendered them only slightly more mobile than sprawling reptiles. ''Desmatosuchus'' possessed both of these features, along with its long femur and elongate pubis, making it more mobile than most thecodonts of its time. This mobility, along with its size, abundance, and specialized beak made it the chief herbivore in the Dockum area. It has also been suggested that ''Desmatosuchus'' could have been omnivorous or even an insectivore. This is because of several similarities between ''Desmatosuchus'' and
armadillos Armadillos (meaning "little armored ones" in Spanish) are New World placental mammals in the order Cingulata. The Chlamyphoridae and Dasypodidae are the only surviving families in the order, which is part of the superorder Xenarthra, along w ...
. For instance, both groups are armored. They possess long snouts that lack teeth on the end. Also, there is evidence of bees, wasps, and termites in the Late Triassic, meaning that ''Desmatosuchus'' had access to insects that armadillos prey on. Their teeth are somewhat similar in shape, although armadillos have more peg-like teeth. Both ''Desmatosuchus'' and armadillos typically carry around 6 teeth on their dentaries. Both armadillos and ''Desmatosuchus'' have hypertrophied processes present on their limb bones, which indicates large limb muscles. This connection is more tenuous, however, since ''Desmatosuchus'' have a crest over their hind limbs but lack one on their forelimbs, meaning that they likely didn't have the musculature for digging with their forelimbs the way armadillos do. In spite of these parallels, the general consensus is still that ''Desmatosuchus'' was most likely herbivorous.


References

{{Taxonbar, from=Q132727 Aetosaurs of North America Chinle fauna Late Triassic pseudosuchians Prehistoric pseudosuchian genera