Typothorax Coccinarum
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''Typothorax'' 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 typothoracine
aetosaur 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 ...
that lived in 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 ...
. Its remains have been found in
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
. Two species are known: ''T. coccinarum'', 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 ...
, and ''T. antiquum''.


Description

''Typothorax'' was an
aetosaur 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 ...
, a
pseudosuchian Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Prior to ...
distantly related to modern crocodilians. Unlike modern crocodilians, aetosaurs were herbivorous. ''Typothorax'' and other aetosaurs possess small, leaf-shaped teeth that were unsuited for a diet consisting of meat.Martz, J.W. 2002. The morphology and ontogeny of Typothorax coccinarum (Archosauria, Stagonolepididae) from the Upper Triassic of the American southwest. M.S. thesis, Geosciences, Texas Tech University, Lubbock, 279 pp. Unlike some aetosaurs such as ''
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 ...
'', ''Typothorax'' does not have large shoulder spikes. It does, however, have a pair of enlarged spikes on the neck projecting from the third row of scutes. It has lateral scutes that bear horns that are posteriorly hooked along its back, while its sides and underbelly are covered with ornamented scutes. Although
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s of aetosaurs are not as common as other Triassic
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 ...
, with their armor plates being the most common, ''Typothorax'' has been represented by fewer skeletal elements than other aetosaurs. ''Typothorax'' was about long and weighed . The vertebral column of ''Typothorax'' is shortened, with individual vertebrae being reduced in length. However, the
osteoderm Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amp ...
s that overly the vertebrae are not shortened. Instead, they are reduced in number so that each dorsal paramedian osteoderm (osteoderm that covers the back) overlies several dorsal vertebrae. In nearly all other crurotarsans, there is one row of osteoderms per vertebra. ''T. coccinarum'' has around 20 rows of presacral osteoderms and about 26 presacral vertebrae. If the cervical spikes of ''Typothorax'' are homologous to those of ''Desmatosuchus'', it is likely that rows of osteoderms were removed from the front. This is because in ''Desmatosuchus'' the spikes are present in the fifth row, while in ''Typothorax'' they are present in the third. In ''T. coccinarum'', there are 10
thoracic The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the crea ...
columns and four caudal columns of ventral osteoderms on the underside. Unlike all other aetosaurs, ''Typothorax'' possesses spiked osteoderms on the underside of the tail, near the
cloaca In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and birds, a ...
. This region is not covered by osteoderms to the extent that it is in other aetosaurs such as ''
Aetosaurus ''Aetosaurus'' is an extinct genus of pseudosuchian reptile belonging to the order Aetosauria. It is generally considered to be the most primitive aetosaur. Three species are currently recognized: ''A. ferratus'', the type species from Germany an ...
'' and '' Coahomasuchus''.


History

''Typothorax'' was one of the earliest vertebrates named from the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
of western North AmericaCope, E.D. 1875. The geology of New Mexico. Philadelphia Academy of Natural Sciences Proceedings, pp. 263-267. but has been poorly understood since that naming.
Paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
s have found the characteristic armor plates for decades,Case, E.C. 1922. New reptiles and stegocephalians from the Upper Triassic of western Texas. Carnegie Institution of Washington Publication, no. 321 (October, 1922). Carnegie Institution of Washington, Washington D.C., 84 pp.Long, R.A., and Murry P.A. 1995. Late Triassic (Carnian and Norian) tetrapods from the southwestern United States. New Mexico Museum of Natural History and Science Bulletin 4, 254 pp. but only recently has a comprehensive study of ''Typothorax'' appeared. It has been found in
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 ...
, in 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 Ne ...
, as well as in
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ker ...
and
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 ...
in the
Bull Canyon Formation The Cooper Canyon Formation is a geological formation of Norian age in Texas and New Mexico.Weishampel, David B; et al. (2004).Dinosaur distribution (Late Triassic, North America)" In: Weishampel, David B.; Dodson, Peter; and Osmólska, Halszka (ed ...
of 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 ...
.


Paleobiology

Like all aetosaurs and many other early crurotarsans, ''Typothorax'' had erect hindlimbs held beneath the body. This is evident by a straight
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 ...
, an anteriorly directed pes (foot), and the projection of the lateral surface of the ilium over the femur. Because the length of the femur is almost equal to that of the tibia and fibula (lower leg) and
astragalus ''Astragalus'' is a large genus of over 3,000 species of herbs and small shrubs, belonging to the legume family Fabaceae and the subfamily Faboideae. It is the largest genus of plants in terms of described species. The genus is native to tempe ...
and
calcaneum In humans and many other primates, the calcaneus (; from the Latin ''calcaneus'' or ''calcaneum'', meaning heel) or heel bone is a bone of the tarsus of the foot which constitutes the heel. In some other animals, it is the point of the hock. St ...
(ankle), ''Typothorax'' was probably slow-moving. The forelimbs are reduced in size and were directed outward in a sprawling position. This posture is also seen in
ankylosaur Ankylosauria is a group of herbivorous dinosaurs of the order Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. ...
s,
ceratopsia Ceratopsia or Ceratopia ( or ; Greek: "horned faces") is a group of herbivorous, beaked dinosaurs that thrived in what are now North America, Europe, and Asia, during the Cretaceous Period, although ancestral forms lived earlier, in the Jurassic. ...
ns, and the early
cynodont The cynodonts () (clade Cynodontia) are a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 mya), and extensively diversified after the Permian–Triassic extinction event. Cynodonts had a wide variety ...
''
Procynosuchus ''Procynosuchus'' (Greek: "Before dog crocodile") is an extinct genus of cynodonts from the Late Permian. It is considered to be one of the earliest and most basal cynodonts. It was 60 cm (2 ft) long. Remains of ''Procynosuchus'' have ...
''. Several aspects of the forelimbs have been interpreted as adaptations to digging. Like many digging
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
s, the
radius In classical geometry, a radius ( : radii) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', meaning ray but also the ...
is significantly shorter than the
humerus 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 roun ...
. Like other aetosaurs, there is a prominent
deltopectoral crest Deltopectoral may refer to; * Clavipectoral triangle, also known as the deltopectoral triangle * Deltopectoral groove * Deltopectoral lymph nodes One or two deltopectoral lymph nodes (or infraclavicular nodes) are found beside the cephalic vein, b ...
on the humerus. The manus is short and wide, a characteristic of digging animals. There is also an entepicondyle on the humerus, which is the origin of forearm pronator and manual flexor muscles. ''Typothorax'' also possesses a relatively long
olecranon process The olecranon (, ), is a large, thick, curved bony eminence of the ulna, a long bone in the forearm that projects behind the elbow. It forms the most pointed portion of the elbow and is opposite to the cubital fossa or elbow pit. The olecranon ...
for the insertion of the '' M. triceps'' muscle, but not as long as those of digging animals. Heckert ''et al.'' (2010) concluded that ''Typothorax'' was not specifically adapted for a
fossorial A fossorial () animal is one adapted to digging which lives primarily, but not solely, underground. Some examples are badgers, naked mole-rats, clams, meerkats, and mole salamanders, as well as many beetles, wasps, and bees. Prehistoric eviden ...
lifestyle but had an ability to dig that other aetosaurs didn't have. The upturned snout of ''Typothorax'' and other aetosaurs suggests that they may have searched for food by rooting around in soil.


Classification

''Typothorax'' belongs to a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
of aetosaurs called the Typothoracinae, within the
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
Aetosaurinae Aetosaurinae is one of the two main clades of aetosaurs, the other being Desmatosuchia. It is a stem-based taxon defined as all aetosaurs more closely related to ''Aetosaurus'' than ''Desmatosuchus''. Aetosaurinae currently comprises ''Aetosaurus ...
. It is closely related to ''
Redondasuchus ''Redondasuchus'' is an extinct genus of aetosaur. It may be a junior synonym of '' Typothorax coccinarum'', another aetosaur. ''Redondasuchus'' is a member of the clade Typothoracisinae within the subfamily Aetosaurinae, and lived during the mi ...
''. ''Redondasuchus reseri'' was first named in 1991, but was later synonymized with ''T. coccinarum'' in 1995 on the basis of the similarity between the osteoderms used to describe the species and the anterior osteoderms of ''T. coccinarum''. Martz (2002) suggested that ''R. reseri'' belongs to ''Typothorax'', but is still its own species, called ''T. reseri''. More recently, the distinctiveness of ''Redondasuchus'' from ''Typothorax'' has been supported with the description of a new species, ''R. rineharti'', in 2006. With new skeletal material from ''T. coccinarum'', Heckert ''et al.'' (2010) claim that ''Redondasuchus'' is distinct from ''Typothorax'' because it has strongly flexed paramedian osteoderms, while ''Typothorax'' has more gently arching paramedian osteoderms. Parker (2013) redescribed 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 type species of ''Typothorax'', ''T. coccinarum''. It was found to be too fragmentary to be diagnosable and therefore represents 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 ...
''. '' Episcoposaurus horridus'', the type species of its genus usually considered to be a
junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...
of ''T. coccinarum'', is the next available name for material referred to ''T. coccinarum''. Therefore, other species of ''Typothorax'' as well as material referred to ''T. coccinarum'', can be reassigned to ''Episcoposaurus''. Nevertheless, to preserve taxonomic stability Parker (2013) suggested to attempt to petition for a
neotype 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 ...
for ''T. coccinarum'' from well preserved specimens which have been previously referred to it.


References


External links


''Typothorax''
in the
Paleobiology Database The Paleobiology Database is an online resource for information on the distribution and classification of fossil animals, plants, and microorganisms. History The Paleobiology Database (PBDB) originated in the NCEAS-funded Phanerozoic Marine Pale ...
*Jeff Martz'
Master's Thesis
from
Texas Tech University Texas Tech University (Texas Tech, Tech, or TTU) is a public research university in Lubbock, Texas. Established on , and called Texas Technological College until 1969, it is the main institution of the five-institution Texas Tech University Sys ...
. {{Taxonbar, from=Q1012908 Aetosaurs of North America Chinle fauna Prehistoric pseudosuchian genera