Parringtonia
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''Parringtonia'' is an extinct
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
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 ...
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 ...
within the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Erpetosuchidae, known from 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 ...
''Parringtonia gracilis''. It is known from a single specimen,
NHMUK The Natural History Museum in London is a museum that exhibits a vast range of specimens from various segments of natural history. It is one of three major museums on Exhibition Road in South Kensington, the others being the Science Museum and ...
R8646, found from the
Anisian In the geologic timescale, the Anisian is the lower stage (stratigraphy), stage or earliest geologic age, age of the Middle Triassic series (stratigraphy), series or geologic epoch, epoch and lasted from million years ago until million years ag ...
-age
Manda Formation The Manda Formation (also known as the Manda Beds) is a Middle Triassic (Anisian?) or possibly Late Triassic (Carnian?) Formation (stratigraphy), geologic formation in Tanzania. It preserves fossils of many terrestrial vertebrates from the Triassi ...
of
Tanzania Tanzania, officially the United Republic of Tanzania, is a country in East Africa within the African Great Lakes region. It is bordered by Uganda to the northwest; Kenya to the northeast; the Indian Ocean to the east; Mozambique and Malawi to t ...
. This specimen, like most archosaur material from the Manda Formation, is fragmentary, including only a
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 ...
and a few postcranial bones. They show similarities with those of another archosaur called ''
Erpetosuchus ''Erpetosuchus'' is an extinct genus of pseudosuchian from the Late Triassic. The type species of ''Erpetosuchus'' is ''E. granti''. It was first described by E. T. Newton in 1894 for remains found in northeastern Scotland, including four specim ...
'', known from the Middle Triassic of Scotland and the eastern United States. The phylogenetic placement of ''Parringtonia'' and ''Erpetosuchus'' are uncertain; some studies placed them close to the group
Crocodylomorpha 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 ...
, which includes all modern crocodylians and many extinct forms that diversified after the Triassic, but this relationship has more recently been questioned.


Description

NHMUK R8646 consists of a right maxilla or upper jaw bone, a left
scapula The scapula (: 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 either side ...
or shoulder blade, part of what might be the
ischium The ischium (; : is ...
bone of the pelvis, five complete and four partial
dorsal vertebra In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebrae of intermediate size between the cervical and lumbar ve ...
e, three
caudal 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 spinal ...
e, and five
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. Some vertebrae show suture lines where different parts have begun to fuse, indicating that the individual was immature when it died. NHMUK R8646 most closely resembles the bones of ''Erpetosuchus granti'' from Scotland and ''Erpetosuchus'' sp. from the eastern United States. The teeth are restricted to the front half of the maxillae in ''Parringtonia'' and ''Erpetosuchus'', and the back of the maxilla is thicker than it is tall. ''Parringtonia'' has five tooth sockets, ''Erpetosuchus granti'' only four, and ''Erpetosuchus'' sp. six or more. Unlike ''Erpetosuchus'', ''Parringtonia'' has a foramen or hole on the outer surface of the maxilla. The scapula of ''Parringtonia'' differs in that it has a small bump or
tubercle In anatomy, a tubercle (literally 'small tuber', Latin for 'lump') is any round nodule, small eminence, or warty outgrowth found on external or internal organs of a plant or an animal. In plants A tubercle is generally a wart-like projectio ...
over its shoulder socket. Both ''Parringtonia'' and ''Erpetosuchus'' have a groove that runs along the top of the
neural arch 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 spinal ...
of each vertebra.


Classification and phylogeny

When first described by
Friedrich von Huene Baron Friedrich Richard von Hoyningen-Huene (22 March 1875 – 4 April 1969) was a German nobleman paleontologist who described a large number of dinosaurs, more than anyone else in 20th-century Europe. He studied a range of Permo-Carbonife ...
in 1939, ''Parringtonia'' was assigned to a group called
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 ...
, which included crocodile-like Triassic archosaurs (although the name has recently been reinstated for the clade including all crocodile-line archosaurs). In 1970 and 1976, ''Parringtonia'' was referred to the family Erpetosuchidae because of similarities between its
scapula The scapula (: 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 either side ...
and that of the better known ''Erpetosuchus''. In both genera, the scapula is very thin and forward-curving. ''Parringtonia'' and ''Erpetosuchus'' are similar in size, and 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 amph ...
s of both genera are similar in shape and sculpture. However, the osteoderms of ''Parringtonia'' are comparable to a number of other small
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 and cannot diagnose it as an erpetosuchid alone. Since ''Parringtonia'' lacks all of the
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to ...
or unique characteristics of ''Erpetosuchus'' including an
otic notch Otic notches are invaginations in the posterior margin of the skull roof, one behind each orbit. Otic notches are one of the features lost in the evolution of amniotes from their tetrapod ancestors. The notches have been interpreted as part of an ...
at the back of the skull and a large
antorbital fenestra An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with Archosauriformes, archosauriforms, first appearing during the Triassic Period. Among Extant ...
set in a deep fossa on the snout, its classification as an erpetosuchid was tentative at first. ''Parringtonia'' was redescribed in 2012 by Nesbitt & Butler and included in a phylogenetic analysis along with ''Erpetosuchus''. The analysis confirmed that ''Parringtonia'' and ''Erpetosuchus'' were
sister taxa In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
in their own clade, which was designated Erpetosuchidae. Three
synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ...
or shared characteristics were identified for Erpetosuchidae: teeth restricted to the anterior half of the maxilla, a posterior half of the maxilla that is thicker than it is tall, and no tooth serrations. Other possible synapomorphies were considered tentative and not included within the analysis. Both taxa share a sharp ridge on the lateral margin of the
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 ...
that marks the lower extent of an opening called the antorbital fossa. Below this ridge, the external surface of the maxilla slopes inward (medially) toward the edge of the jaw. In ''Erpetosuchus'', this medially inclined external surface of the maxilla continues back onto the
jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
bone so that a large part of the external surface of the jugal faces downward. This morphology unites the North American and European specimens of ''Erpetosuchus'' with ''Parringtonia gracilis''. Other potential synapomorphies include a hypertrophied or enlarged tuber at the bottom of the scapula that is thought to be the attachment point of the
triceps brachii muscle The triceps, or triceps brachii (Latin for "three-headed muscle of the arm"), is a large muscle on the back of the upper limb of many vertebrates. It consists of three parts: the medial, lateral, and long head. All three heads cross the elbow jo ...
. Unlike other archosauriforms that have a small tuber in the same location, the size of the tuber in erpetosuchids is exceptionally large in relation to the overall size of the scapula. The forward-curving scapula that was first noted in 1976 as a shared erpetosuchid feature was noted to be present in several other archosaurs such as ''
Postosuchus kirkpatricki ''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 m ...
'', reducing its utility as a
synapomorphy In phylogenetics, an apomorphy (or derived trait) is a novel Phenotypic trait, character or character state that has evolution, evolved from its ancestral form (or Plesiomorphy and symplesiomorphy, plesiomorphy). A synapomorphy is an apomorphy sh ...
. While the 2012 analysis strongly supported the inclusion of ''Parringtonia'' within Erpetosuchidae, the position of Erpetosuchidae within Archosauria was uncertain. Erpetosuchidae formed a
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tree ...
or unresolved evolutionary relationship with several other archosaur groups, including
Avemetatarsalia Avemetatarsalia (meaning "bird metatarsals") is a clade of diapsid Reptile, reptiles containing all archosaurs more closely related to birds than to crocodilians. The two most successful groups of avemetatarsalians were the dinosaurs and pterosau ...
,
Ornithosuchidae Ornithosuchidae is an extinct family (biology), family of Pseudosuchia, pseudosuchian archosaurs (distant relatives of modern Crocodilia, crocodilians) from the Triassic period. Ornithosuchids were quadrupedal and facultatively bipedal (e.g. like ...
,
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 ...
and ''
Revueltosaurus ''Revueltosaurus'' ("Revuelto lizard") is an extinct genus of suchian pseudosuchian from Late Triassic (late Carnian to middle Norian stage) deposits of New Mexico, Arizona and North Carolina, United States. Many specimens, mostly teeth, have bee ...
'', ''
Ticinosuchus ''Ticinosuchus'' is an extinct genus of suchian archosaur from the Middle Triassic (Anisian– Ladinian) of Switzerland and Italy. Description One of only a handful of fossil reptiles that have been found in Switzerland, ''Ticinosuchus'' (mea ...
'' and
Paracrocodylomorpha Paracrocodylomorpha is a clade of pseudosuchian archosaurs. The clade includes the diverse and unusual group Poposauroidea as well as the generally carnivorous and quadrupedal members of Loricata, including modern crocodylians. Paracrocodylomor ...
, ''
Gracilisuchus ''Gracilisuchus'' (meaning "slender crocodile") is an extinct genus of tiny pseudosuchian (a group which includes the ancestors of crocodilians) from the Late Triassic of Argentina. It contains a single species, ''G. stipanicicorum'', which is pl ...
'', and ''
Turfanosuchus ''Turfanosuchus'' is a genus of archosauriform reptile, likely a Gracilisuchidae, gracilisuchid archosaur, which lived during the Middle Triassic (Anisian) of northwestern China. The type species, ''T. dabanensis'', was described by Yang Zhongjia ...
''. The removal of ''Gracilisuchus'' and ''Turfanosuchus'' from the analysis resulted in Erpetosuchidae nesting within the clade
Suchia Suchia (from Ancient Greek σούχος (''soúkhos''), meaning "crocodile") is a clade of archosaurs containing the majority of pseudosuchians (crocodilians and their extinct relatives). It was defined as the least inclusive clade containing ''A ...
as the sister-taxon of
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 ...
plus ''
Revueltosaurus ''Revueltosaurus'' ("Revuelto lizard") is an extinct genus of suchian pseudosuchian from Late Triassic (late Carnian to middle Norian stage) deposits of New Mexico, Arizona and North Carolina, United States. Many specimens, mostly teeth, have bee ...
'' clade, but not as a sister taxon of Crocodylomorpha as had previously been proposed.


References


External links


''Parringtonia''
in the
Paleobiology Database The Paleobiology Database (PBDB) is an online resource for information on the distribution and classification of fossil animals, plants, and microorganisms. History The Paleobiology Database originated in the NCEAS-funded Phanerozoic Marine Pale ...
{{Taxonbar, from=Q16987513 Erpetosuchidae Prehistoric pseudosuchian genera Triassic archosaurs Anisian life Middle Triassic reptiles of Africa Triassic Tanzania Fossils of Tanzania Fossil taxa described in 1939