Archeopelta
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''Archeopelta'' 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
carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose nutrition and energy requirements are met by consumption of animal tissues (mainly mu ...
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 ...
from the late Middle or early Late
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 ...
period (late
Ladinian The Ladinian is a stage and age in the Middle Triassic series or epoch. It spans the time between Ma and ~237 Ma (million years ago). The Ladinian was preceded by the Anisian and succeeded by the Carnian (part of the Upper or Late Triassic ...
to early
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 ...
stage). It was a 2 m (6 ft) long
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
which lived in what is now southern Brazil. Its exact phylogenetic placement within Archosauriformes is uncertain; it was originally classified as a doswelliid, but subsequently it was argued to be an
erpetosuchid Erpetosuchidae is an extinct family of pseudosuchian archosaurs. Erpetosuchidae was named by D. M. S. Watson in 1917 to include ''Erpetosuchus''. It includes the type species '' Erpetosuchus granti'' from the Late Triassic of Scotland, ''Erpetos ...
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 ...
.


Discovery

It is only known from 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 ...
CPEZ-239a, which consists of partial
skeleton A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal fra ...
(including
vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
e, partial right front and hind limbs, a partial hip, and an undetermined bone which may be part of a tibia) 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 ...
. It was found in the Santa Maria 1 Sequence, previously known as the
Santa Maria Formation The Santa Maria Formation is a sedimentary rock formation found in Rio Grande do Sul, Brazil. It is primarily Carnian in age (Late Triassic), and is notable for its fossils of cynodonts, " rauisuchian" pseudosuchians, and early dinosaurs and othe ...
, in Chiniquá region, São Pedro do Sul of
Rio Grande do Sul Rio Grande do Sul (, ; ; "Great River of the South") is a Federative units of Brazil, state in the South Region, Brazil, southern region of Brazil. It is the Federative units of Brazil#List, fifth-most populous state and the List of Brazilian s ...
State. It was first named by Julia B. Desojo, Martín D. Ezcurra and César L. Schultz in
2011 The year marked the start of a Arab Spring, series of protests and revolutions throughout the Arab world advocating for democracy, reform, and economic recovery, later leading to the depositions of world leaders in Tunisia, Egypt, and Yemen ...
and 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 ...
is ''Archeopelta arborensis''. The generic name comes from ''archaios'', ancient in
Greek Greek may refer to: 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 of all kno ...
and ''pelta'', shield, in reference to its thick
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 amph ...
s. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
is derived from ''arbor'', tree in
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
, in reference to Sanga da Árvore where the fossils were found.


Description

CPEZ-239a's braincase is low, without an occipital neck. The exoccipitals do not meet each other medially. The paraoccipital processes extend outwards and slightly downwards, and the supraoccipital has a ridge. Uniquely, a very deep fossa is present in the corner of the opisthotics. The exit of the
hypoglossal nerve The hypoglossal nerve, also known as the twelfth cranial nerve, cranial nerve XII, or simply CN XII, is a cranial nerve that innervates all the extrinsic and intrinsic muscles of the tongue except for the palatoglossus, which is innervated b ...
is a single opening. The basal tubera are very low and separated by a deep notch. The parabasisphenoid is short and its exits for the
internal carotid arteries The internal carotid artery is an artery in the neck which supplies the anterior and middle cerebral circulation. In human anatomy, the internal and external carotid arise from the common carotid artery, where it bifurcates at cervical verteb ...
are small and pushed to the rear edge of the bone. The basipterygoid processes are close to each other and project anterolaterally. The exit for the
facial nerve The facial nerve, also known as the seventh cranial nerve, cranial nerve VII, or simply CN VII, is a cranial nerve that emerges from the pons of the brainstem, controls the muscles of facial expression, and functions in the conveyance of ta ...
is present on the vestibule, and the lamina separating the metotic foramen and the
fenestra ovalis The oval window (or ''fenestra vestibuli'' or ''fenestra ovalis'') is a connective tissue membrane-covered opening from the middle ear to the cochlea of the inner ear. Vibrations that contact the tympanic membrane travel through the three ossicl ...
is very thin. The back vertebrae are only slightly elongated and are not constricted from the sides. The diapophyses are thick, subrectangular, and elongated. The
prezygapophyses The articular process or zygapophysis ( + apophysis) of a vertebra is a projection of the vertebra that serves the purpose of fitting with an adjacent vertebra. The actual region of contact is called the ''articular facet''.Moore, Keith L. et al. ...
are short while the neural spines are long and oval-shaped in cross section. The first 'primordial' sacral vertebra (likely the second sacral) is low and wide, with characteristically-shaped sacral ribs. The sacral ribs expand anteroposteriorly at their tips, with the anterior expansions being large and subtriangular. The first primordial's unique prezygapophyses are very large and circular, with their faces pointing upward and inwards. On the other hand, the postzygapophyses are short, downwards-pointing, and connected by a V-shaped hyposphene. Although incomplete, the second 'primordial' sacral (likely the third sacral) is also low, with sacral ribs similarly shaped to those of the first 'primordial' sacral. The
humeral head The humerus (; : humeri) 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 ...
is offset from the humeral shaft, and the right
humerus The humerus (; : humeri) 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 (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
as a whole is wide and distally tapering (although missing the distal portion). A thin bone has been interpreted to be a right ilium, with an unusual illiac blade which is S-shaped in posterior view. However, this interpretation is uncertain and the bone's shape may be a result of postmortem deformation. The right
ischium The ischium (; : is ...
has a long and deep pubic peduncle but a very short illiac peduncle. The ischial shaft is thin and the lower edge bends towards the midline. The right
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 ...
is S-shaped from the front, with a transversely very wide distal end and poorly developed condyles and tubercules. The right
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
is anteroposteriorly wide but distally tapering and missing its distal tip. An unusual rod-like bone may be the distal part of a tibia. Several
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 amph ...
s are preserved with CPEZ-239a. At least two rows of osteoderms attached to each neural spine of the vertebrae, and there is evidence that additional rows of lateral osteoderms were also present. The osteoderms are very thick and quadrangular in shape, with straight posterior and medial borders and rounded anterior and lateral borders. They were rough and covered in deep, circular pits, with each possessing an anterior articular lamina (a smooth area where the preceding osteoderm would have overlapped the front edge of the following osteoderm). Although they had serrated edges, they did not possess a raised keel or peak (a dorsal prominence) on their surface. Small, circular plates attached to the femur may be appendicular osteoderms, although poor preservation makes this uncertain.


Classification

Upon the initial description of ''Archeopelta'', it was placed as a close relative of ''
Doswellia ''Doswellia'' is an extinct genus of archosauriform from the Late Triassic of North America. It is the most notable member of the family Doswelliidae, related to the Proterochampsidae, proterochampsids. ''Doswellia'' was a low and heavily built c ...
'' in the newly created family Doswellidae. This referral was due to the structure of its osteoderms, which were very similar to those of ''Doswellia''. In addition, ''Archeopelta'' shared several other features with ''Doswellia'' which were unknown in ''
Tarjadia ''Tarjadia'' is an extinct genus of erpetosuchid pseudosuchian, distantly related to modern crocodilians. It is known from a single species, ''T. ruthae'', first described in 1998 from the Middle Triassic Chañares Formation in Argentina. Parti ...
'', which was rather incomplete at the time of ''Archeopelta's'' description. Among these features are the wide first primordial sacral, a long and laterally deflected illiac blade, and anterolaterally-projected basipterygoid processes. This is the
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
from the study, after Desojo, Ezcurra & Schultz, 2011: In 2013, Lucas, Spielmann, and Hunt claimed that ''Archeopelta'' was a
junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...
of ''Tarjadia'' due to a lack of distinguishing features between the two. However, Ezcurra (2016) provided several differences between the two genera. For example, ''Archeopelta'' has unconstricted dorsal vertebrae and a ridge on the supraoccipital, while ''Tarjadia'' has the opposite traits. Nevertheless, Ezcurra observed that both genera had very thick osteoderms, and he considered that both of them were doswellids more closely related to each other than to ''Doswellia''. New specimens of ''Tarjadia'' described in 2017 provided a
phylogenetic analysis In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
that argued that ''Tarjadia'' and ''Archeopelta'' were not doswellids, but rather
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 ...
n
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 ...
s of the family
Erpetosuchidae Erpetosuchidae is an extinct family of pseudosuchian archosaurs. Erpetosuchidae was named by D. M. S. Watson in 1917 to include ''Erpetosuchus''. It includes the type species '' Erpetosuchus granti'' from the Late Triassic of Scotland, ''Erpetos ...
. The strict consensus tree of this analysis is given below. Erpetosuchid osteoderms are similar to Doswellid osteoderms, which explains the earlier classification of ''Tarjadia'' and ''Archeopelta''. Even in non-parmonious phylogenetic trees of the analysis which forced them to be recovered as doswellids, ''Tarjadia'' and ''Archeopelta'' still formed a clade with ''Erpetosuchus''. Despite the change in ''Archeopelta's'' classification, it still forms a clade with ''Tarjadia'' as an erpetosuchid. This is due to both of them possessing a very short or absent occipital neck and osteoderms without a dorsal prominence.


References

{{Portal bar, Brazil, Paleontology, Reptiles Erpetosuchidae Triassic reptiles of South America Triassic Brazil Fossils of Brazil Santa Maria Formation Fossil taxa described in 2011 Prehistoric pseudosuchian genera