Asperoris

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Asperoris
Temporal range: Middle Triassic, 245–242  Ma
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Asperoris mnyama.png
Front part of the skull
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauromorpha
Clade: Crocopoda
Clade: Archosauriformes
Genus: Asperoris
Nesbitt et al., 2013
Type species
Asperoris mnyama
Nesbitt et al., 2013

Asperoris is an extinct genus of archosauriform reptile known from the Middle Triassic Manda Beds of southwestern Tanzania. It is the first archosauriform known from the Manda Beds that is not an archosaur. However, its relationships with other non-archosaurian archosauriforms are uncertain. It was first named by Sterling J. Nesbitt, Richard J. Butler and David J. Gower in 2013 and the type species is Asperoris mnyama. Asperoris means "rough face" in Latin, referring to the distinctive rough texture of its skull bones. [1]

Contents

Discovery

Location of the discovery Lifua Member of the Manda beds.png
Location of the discovery

Asperoris is known solely from the well-preserved but incomplete holotype skull NHMUK PV R36615, which includes the right premaxilla and maxilla bones of the upper jaw, the right nasal, prefrontal, frontal, postfrontal, and parietal bones of the top of the skull, and part of the right postorbital bone behind the eye socket, as well as three unidentified skull fragments. NHMUK PV R36615 was discovered by a joint 1963 expedition of the Natural History Museum and the University of London to eastern Zambia and western Tanzania (then northern Rhodesia and Tanganyika, respectively). Asperoris comes from Manda Beds of the Ruhuhu Basin in Songea Urban District of southwestern Tanzania, which dates back to the late Anisian stage of the Middle Triassic. Based on field notes, NHMUK PV R36615 was found on August 23 in a drainage of the Hita River between the Njalila and Hiasi rivers also known as locality U9/1 of the Lifua Member of Manda Beds, near the remains of dicynodonts. It was fully described in 2013 and assigned to a new genus and species, Asperoris mnyama, by Sterling J. Nesbitt, Richard J. Butler and David J. Gower in the journal PLoS One. The generic name is derived from asper, meaning "rough", and oris, meaning "face" in Latin in reference to its rough and uniquely sculptured skull. The specific name mnyama means "beast" in Swahili. [1]

Description

Restored skull Asperoris.png
Restored skull

Like most archosauriforms from the Manda Beds, Asperoris is known from very fragmentary remains. NHMUK PV R36615 is distinguished from some other archosauriform material in the beds by the lack of a depression called an antorbital fossa on the surfaces of its maxilla and premaxilla. Although it can not be directly compared with Stagonosuchus , Hypselorhachis , Nyasasaurus , Teleocrater and an unnamed suchian, its inferred phylogenetic position is not consistent with it belonging to any of these taxa. The total length of the skull is estimated to have been 50 centimetres (20 in) in length. A unique characteristic or autapomorphy of Asperoris is the rough texture of its skull bones, particularly the frontal. The skull roof of Asperoris is relatively thick compared to those of other archosauriforms and its antorbital fenestra, a hole in the side of the skull in front of the eye socket, is relatively narrow. [1]

Relationships

Skull roof Asperoris skull roof.png
Skull roof

Asperoris belongs to a clade or evolutionary grouping of reptiles called Archosauriformes, which includes Archosauria (the clade including living crocodilians and birds) and their extinct, mostly Triassic, relatives. It has several features that place it outside Archosauria with non-archosaurian archosauriforms, including the lack of an antorbital fossa and the possible presence of a postparietal bone at the back of the skull, which is not found in archosaurs. However, it also has features that place it among the closest relatives of archosaurs, including the presence of an antorbital fenestra and the lack of a hole called the parietal foramen on the skull roof. Asperoris shares with a group of archosauriforms called erythrosuchids the presence of slot in the lower part of the nasal bone that fits into a projection of the premaxilla beneath it. It also has a thecodont dentition, meaning that its teeth fit into deep sockets in the jaw. A thecodont dentition is seen in all non-archosaurian archosauriforms except proterosuchids, and is characteristic of the clade. [1]

Premaxilla Asperoris premaxilla.png
Premaxilla

Because of the poor preservation of NHMUK PV R36615 and the phylogenetic analysis used in the study (which focuses on relationships of Triassic archosaurs), the phylogenetic relationships of Asperoris are uncertain. Nesbitt, Butler and Gower's 2013 phylogenetic analysis resulted in a strict consensus tree with Asperoris in a polytomy or unresolved phylogenetic relationship with Erythrosuchus africanus , an erythrosuchid, Vancleavea campi , an aquatic archosauriform, the proterosuchid clade, and the clade including Euparkeria capensis , phytosaurs, and Archosauria. Other phylogenetic trees produced in the analysis placed Asperoris as the sister taxon (closest relative) of either Erythrosuchus africanus or Euparkeria capensis. A sister-taxon relationship with Erythrosuchus is more likely because it is based on a derived feature, the slot in the nasal, while the relationship with Euparkeria is less likely because it is based only on characteristics inherited from archosauriform ancestors (plesiomorphies). [1]

Asperoris was also featured in a phylogenetic analysis by Martin Ezcurra in 2016. Most parts of Ezcurra's analysis omitted this genus due to its incompleteness, but in versions which did feature it, it was found in a polytomy with Yarasuchus , Dongusuchus , Dorosuchus , and Euparkeria at the base of a clade which also includes proterochampsians and archosaurs. Ezcurra named this broad clade Eucrocopoda. The five-genus polytomy is resolved into a clearer system of clades when Asperoris is omitted, as the taxon's lack of postcranial features reduced the analysis's clarity. [2] Yarasuchus and Dongusuchus are now considered to be aphanosaurs, part of a group of archosaurs at the base of the branch that leads to pterosaurs and dinosaurs (including birds) but not crocodiles. [3]

Related Research Articles

<span class="mw-page-title-main">Archosauriformes</span> Clade of reptiles

Archosauriformes is a clade of diapsid reptiles encompassing archosaurs and some of their close relatives. It was defined by Jacques Gauthier (1994) as the clade stemming from the last common ancestor of Proterosuchidae and Archosauria. Phil Senter (2005) defined it as the most exclusive clade containing Proterosuchus and Archosauria. Archosauriforms are a branch of archosauromorphs which originated in the Late Permian and persist to the present day as the two surviving archosaur groups: crocodilians and birds.

<i>Euparkeria</i> Extinct genus of reptiles

Euparkeria is an extinct genus of archosauriform reptile from the Triassic of South Africa. Euparkeria is close to the ancestry of Archosauria, the reptile group that includes crocodilians, pterosaurs, and dinosaurs.

<i>Teleocrater</i> Extinct genus of reptiles

Teleocrater is a genus of avemetatarsalian archosaur from the Middle Triassic Manda Formation of Tanzania. The name was coined by English paleontologist Alan Charig in his 1956 doctoral dissertation, but was only formally published in 2017 by Sterling Nesbitt and colleagues. The genus contains the type and only species T. rhadinus. Uncertainty over the affinities of Teleocrater have persisted since Charig's initial publication; they were not resolved until Nesbitt et al. performed a phylogenetic analysis. They found that Teleocrater is most closely related to the similarly enigmatic Yarasuchus, Dongusuchus, and Spondylosoma in a group that was named the Aphanosauria. Aphanosauria was found to be the sister group of the Ornithodira, the group containing dinosaurs and pterosaurs.

<i>Erythrosuchus</i> Extinct genus of reptiles

Erythrosuchus is an extinct genus of archosauriform reptiles from the Triassic of South Africa. Remains have been found from the Cynognathus Assemblage Zone of the Beaufort Group in the Karoo of South Africa.

<i>Proterosuchus</i> Extinct genus of reptiles from the Early Triassic of South Africa

Proterosuchus is an extinct genus of archosauriform reptiles that lived during the Early Triassic. It contains three valid species: the type species P. fergusi and the referred species P. alexanderi and P. goweri. All three species lived in what is now South Africa. The genus was named in 1903 by the South African paleontologist Robert Broom. The genus Chasmatosaurus is a junior synonym of Proterosuchus.

<i>Erpetosuchus</i> Extinct genus of reptiles

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 specimens from the latest Carnian Lossiemouth Sandstone Formation. Additional remains of Erpetosuchus have been found in the New Haven Formation of Connecticut in the eastern United States, although they were not attributed to the species E. granti. The relationship of Erpetosuchus to other archosaurs is uncertain. In 2000 and 2002, it was considered a close relative of the group Crocodylomorpha, which includes living crocodylians and many extinct relatives. However, this relationship was questioned in a 2012 analysis that found the phylogenetic placement of Erpetosuchus to be very uncertain.

<span class="mw-page-title-main">Proterochampsidae</span> Extinct family of reptiles

Proterochampsidae is a family of proterochampsian archosauriforms. Proterochampsids may have filled an ecological niche similar to modern crocodiles, and had a general crocodile-like appearance. They lived in what is now South America in the Middle and Late Triassic.

<i>Turfanosuchus</i> Extinct genus of reptiles

Turfanosuchus is a genus of archosauriform reptile, likely a gracilisuchid archosaur, which lived during the Middle Triassic (Anisian) of northwestern China. The type species, T. dabanensis, was described by C.C. Young in 1973, based on a partially complete but disarticulated fossil skeleton found in the Kelamayi Formation of the Turfan Basin.

Yarasuchus is an extinct genus of avemetatarsalian archosaur that lived during the Anisian stage of the Middle Triassic of India. The genus was named and described in 2005 from a collection of disarticulated but fairly complete fossil material found from the Middle Triassic Yerrapalli Formation. The material is thought to be from two individuals, possibly three, with one being much more complete and articulated than the other. The type and only species is Y. deccanensis. Yarasuchus was a quadruped roughly 2–2.5 metres (6.6–8.2 ft) long, with an elongated neck and tall spines on its vertebrae. Unlike other quadrupedal Triassic reptiles, the limbs and shoulders of Yarasuchus were slender, and more like those of ornithodirans.

Dongusuchus is an extinct genus of archosaur. Fossils have been found from the Donguz Formation outcropping on the banks of the Donguz River in the Orenburg Oblast of Russia. They are associated with a fossil assemblage called the Eryosuchus Fauna, named after the capitosaurid Eryosuchus, the most common organism found from the assemblage. The locality dates back to the Anisian and early Ladinian stages of the Middle Triassic.

<i>Parringtonia</i> Extinct genus of reptiles

Parringtonia is an extinct genus of Triassic archosaur within the family Erpetosuchidae, known from the type species Parringtonia gracilis. It is known from a single specimen, NHMUK R8646, found from the Anisian-age Manda Formation of Tanzania. This specimen, like most archosaur material from the Manda Formation, is fragmentary, including only a maxilla and a few postcranial bones. They show similarities with those of another archosaur called Erpetosuchus, 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, which includes all modern crocodylians and many extinct forms that diversified after the Triassic, but this relationship has more recently been questioned.

<i>Yonghesuchus</i> Extinct genus of reptiles

Yonghesuchus is an extinct genus of Late Triassic archosaur reptile. Remains have been found from the early Late Triassic Tongchuan Formation in Shanxi, China. It is named after Yonghe County, the county where fossils were found. Currently only one species, Y. sangbiensis, is known. The specific name refers to Sangbi Creek, as fossils were found in one of its banks.

Koilamasuchus is an extinct genus of indeterminate archosauriform from the Triassic of Argentina. It is based on an external mold of a partial postcranial skeleton from the Quebrada de los Fósiles Formation. Due to its incomplete nature, the relationships of this reptile are difficult to establish. Originally described as a non-archosaur archosauriform, later studies tentatively considered it a doswelliid or a suchian archosaur.

<span class="mw-page-title-main">Suchia</span> Clade of reptiles

Suchia is a clade of archosaurs containing the majority of pseudosuchians. It was defined as the least inclusive clade containing Aetosaurus ferratus, Rauisuchus tiradentes, Prestosuchus chiniquensis, and Crocodylus niloticus by Nesbitt (2011). Generally the only pseudosuchian group which is omitted from Suchia is the family Ornithosuchidae, although at least one analysis classifies ornithosuchids as close relatives of erpetosuchids and aetosaurs. Phytosaurs are also excluded from Suchia, although it is not certain whether they qualify as pseudosuchians in the first place.

<span class="mw-page-title-main">Doswelliidae</span> Extinct family of reptiles

Doswelliidae is an extinct family of carnivorous archosauriform reptiles that lived in North America and Europe during the Middle to Late Triassic period. Long represented solely by the heavily-armored reptile Doswellia, the family's composition has expanded since 2011, although two supposed South American doswelliids were later redescribed as erpetosuchids. Doswelliids were not true archosaurs, but they were close relatives and some studies have considered them among the most derived non-archosaurian archosauriforms. They may have also been related to the Proterochampsidae, a South American family of crocodile-like archosauriforms.

<span class="mw-page-title-main">Proterochampsia</span> Extinct clade of reptiles

Proterochampsia is a clade of early archosauriform reptiles from the Triassic period. It includes the Proterochampsidae and probably also the Doswelliidae. Nesbitt (2011) defines Proterochampsia as a stem-based taxon that includes Proterochampsa and all forms more closely related to it than Euparkeria, Erythrosuchus, Passer domesticus, or Crocodylus niloticus. Therefore, the inclusion of Doswelliidae in it is dependent upon whether Doswellia and Proterochampsa form a monophyletic group to the exclusion of Archosauria and other related groups.

<span class="mw-page-title-main">Erpetosuchidae</span> Extinct family of reptiles

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, Erpetosuchus sp. from the Late Triassic of eastern United States and Parringtonia gracilis from the middle Middle Triassic of Tanzania; the group might also include Dyoplax arenaceus from the Late Triassic of Germany, Archeopelta arborensis and Pagosvenator candelariensis from Brazil and Tarjadia ruthae from Argentina.

Nundasuchus is an extinct genus of crurotarsan, possibly a suchian archosaur related to Paracrocodylomorpha. Remains of this genus are known from the Middle Triassic Manda beds of southwestern Tanzania. It contains a single species, Nundasuchus songeaensis, known from a single partially complete skeleton, including vertebrae, limb elements, osteoderms, and skull fragments.

<i>Polymorphodon</i> Extinct genus of reptiles

Polymorphodon is an extinct genus of archosauriform reptile from the Middle Triassic of Germany. The only known species is Polymorphodon adorfi, discovered in Lower Keuper deposits at a quarry in Eschenau, Germany. Polymorphodon is notable for its heterodont dentition, with long and conical premaxillary teeth followed by thin maxillary teeth with large serrations. Maxillary teeth near the back of the mouth are short and leaf-shaped, similar to some living and extinct reptiles with a herbivorous or omnivorous diet. This may suggest that Polymorphodon had some reliance on plants in its diet, a rarity among basal archosauriforms, most of which are carnivores.

<i>Mambawakale</i> Extinct genus of reptiles

Mambawakale is a genus of large sized basal paracrocodylomorph, possibly a poposauroid, from the Manda Beds of Tanzania. It was informally named Pallisteria before being officially published under its current name almost 60 years after its discovery. It contains a single species, Mambawakale ruhuhu.

References

  1. 1 2 3 4 5 Nesbitt, S. J.; Butler, R. J.; Gower, D. J. (2013). Joger, Ulrich (ed.). "A New Archosauriform (Reptilia: Diapsida) from the Manda Beds (Middle Triassic) of Southwestern Tanzania". PLOS ONE. 8 (9): e72753. Bibcode:2013PLoSO...872753N. doi: 10.1371/journal.pone.0072753 . PMC   3785487 . PMID   24086264.
  2. Ezcurra, Martín D. (2016-04-28). "The phylogenetic relationships of basal archosauromorphs, with an emphasis on the systematics of proterosuchian archosauriforms". PeerJ. 4: e1778. doi: 10.7717/peerj.1778 . ISSN   2167-8359. PMC   4860341 . PMID   27162705.
  3. Nesbitt, Sterling J.; Butler, Richard J.; Ezcurra, Martín D.; Barrett, Paul M.; Stocker, Michelle R.; Angielczyk, Kenneth D.; Smith, Roger M. H.; Sidor, Christian A.; Niedźwiedzki, Grzegorz (2017). "The earliest bird-line archosaurs and the assembly of the dinosaur body plan" (PDF). Nature. 544 (7651): 484–487. Bibcode:2017Natur.544..484N. doi:10.1038/nature22037. ISSN   1476-4687. PMID   28405026. S2CID   262955669.