Typothorax
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''Typothorax'' 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 typothoracine
aetosaur Aetosaurs () are heavily armored reptiles belonging to the extinct order (biology), order Aetosauria (; from Ancient Greek, Greek, (aetos, "eagle") and (, "lizard")). They were medium- to large-sized Omnivore, omnivorous or Herbivore, herbivor ...
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 a ...
. Its remains have been found in
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
. Two species are known: ''T. coccinarum'', the
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
, and ''T. antiquum''.


Description

''Typothorax'' was an
aetosaur Aetosaurs () are heavily armored reptiles belonging to the extinct order (biology), order Aetosauria (; from Ancient Greek, Greek, (aetos, "eagle") and (, "lizard")). They were medium- to large-sized Omnivore, omnivorous or Herbivore, herbivor ...
, a pseudosuchian 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 language, 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 ''Desma ...
'', ''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 preserve ...
s of aetosaurs are not as common as other Triassic archosaurs, with their armor plates being the most common, ''Typothorax'' has been represented by fewer skeletal elements than other aetosaurs. The largest known specimen of ''T. coccinarum'', described in 2023, would have been similar in size to ''Desmatosuchus'', measuring over long, though it has been identified as a skeletally immature individual (16 to 19 years old) based on the open neurocentral suture and the lines of arrested growth. However, the authors warn that, since smaller individuals have co-ossified neurocentral sutures, they may not be a reliable factor in determining maturity in aetosaurs. So this individual still may be an adult. The vertebral column of ''Typothorax'' is shortened, with individual vertebrae being reduced in length. However, the osteoderms 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 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. This region is not covered by osteoderms to the extent that it is in other aetosaurs such as '' Aetosaurus'' and '' Coahomasuchus''.


History

''Typothorax'' was one of the earliest vertebrates named from the
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 ...
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. Paleontologists 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 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 ...
, in the Chinle Formation, as well as in
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 ...
and
Texas Texas ( , ; or ) is the most populous U.S. state, state in the South Central United States, South Central region of the United States. It borders Louisiana to the east, Arkansas to the northeast, Oklahoma to the north, New Mexico to the we ...
in the Bull Canyon Formation of the Dockum Group.


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 (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
, 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 and calcaneum (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 ankylosaurs, ceratopsians, and the early
cynodont Cynodontia () is a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 Megaannum, mya), and extensively diversified after the Permian–Triassic extinction event. Mammals are cynodonts, as are their extin ...
'' Procynosuchus''. Several aspects of the forelimbs have been interpreted as adaptations to digging. Like many digging
tetrapod A tetrapod (; from Ancient Greek :wiktionary:τετρα-#Ancient Greek, τετρα- ''(tetra-)'' 'four' and :wiktionary:πούς#Ancient Greek, πούς ''(poús)'' 'foot') is any four-Limb (anatomy), limbed vertebrate animal of the clade Tetr ...
s, the
radius In classical geometry, a radius (: radii or radiuses) of a circle or sphere is any of the line segments from its Centre (geometry), center to its perimeter, and in more modern usage, it is also their length. The radius of a regular polygon is th ...
is significantly shorter than the humerus. Like other aetosaurs, there is a prominent deltopectoral crest 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 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 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 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 ...
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 botanical subfamily names with "-oideae", and zo ...
Aetosaurinae. It is closely related to '' Redondasuchus''. ''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 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 ...
''. '' Episcoposaurus horridus'', the type species of its genus usually considered to be a junior synonym 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 for ''T. coccinarum'' from well preserved specimens which have been previously referred to it.


References


External links


''Typothorax''
in the Paleobiology Database *Jeff Martz'
Master's Thesis
from Texas Tech University. {{Taxonbar, from=Q1012908 Aetosaurs of North America Chinle fauna Prehistoric pseudosuchian genera Taxa named by Edward Drinker Cope