Doswelliidae

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Doswellids
Temporal range: Middle - Late Triassic, 242–220  Ma
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Possible Rhaetian records if Vancleavea is a member.
Doswellia kaltenbachi life restoration.png
Life restoration Doswellia kaltenbachi , the most well known dosweliid
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauromorpha
Clade: Archosauriformes
Clade: Proterochampsia
Family: Doswelliidae
Weems, 1980
Genera

Doswelliidae is an extinct family of carnivorous archosauriform reptiles that lived in North America and Europe during the Middle to Late Triassic period. [1] Long represented solely by the heavily-armored reptile Doswellia , the family's composition has expanded since 2011, although two supposed South American doswelliids ( Archeopelta and Tarjadia ) 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. [2] They may have also been related to the Proterochampsidae, a South American family of crocodile-like archosauriforms. [3]

Contents

Description

Doswelliids are believed to be semiaquatic carnivores similar to crocodilians in appearance, as evidenced by their short legs and eyes and nostrils which are set high on the head, though the putative member Scleromochlus has been interpreted as a frog-like hopper by one study. [4] They had long bodies and tails, and their front legs were shorter than their hind legs. Unlike in some other groups of archosauriforms, doswelliids retain teeth on the pterygoid, on the roof of the mouth. Although Vancleavea had a short and deep skull, most doswelliids had slender and elongated snouts, similar to other members of Proterochampsia. Advanced doswelliids possessed dorsal ribs which splay outwards (rather than downwards), making their bodies wide and low. [5]

Doswelliids were armored with multiple rows of bony scutes (osteoderms) on their backs. With the exception of Vancleavea, which had many different forms of smooth osteoderms, doswelliid osteoderms were characteristically covered by deep, circular pits. There is also a smooth area (an anterior articular lamina) on the front edge of each osteoderm where the preceding osteoderm overlaps. This combination of osteoderm features is also present in erpetosuchids and some aetosaurs, although the osteoderms of the latter group differ in the arrangement of the pits and the fact that the anterior articular lamina is formed by a raised bar. [3] Doswellia had at least ten rows of osteoderms, creating a flattened carapace-like armor plate on its back. Jaxtasuchus had lighter armor, with only four rows. [5]

Classification

The family was originally named by R. E. Weems in 1980 and was placed in its own suborder, Doswelliina. The Doswelliidae has long been considered a monospecific family of basal archosauriforms represented by Doswellia kaltenbachi from the Late Triassic of North America. [6]

However, a 2011 cladistic analysis by Desojo, Ezcurra, & Schultz recovered the newly named Brazilian genus Archeopelta as well as the enigmatic Argentinian archosauriform Tarjadia as close relatives of Doswellia, within a monophyletic Doswelliidae. These authors defined the family as the most inclusive clade containing all archosauromorphs more closely related to Doswellia kaltenbachi than to Proterochampsa barrionuevoi, Erythrosuchus africanus, Caiman latirostris, (the broad-snouted caiman) or Passer domesticus (the house sparrow). [2]

Within the next few years, several other genera of archosauriforms were classified as Dosweliids. A second New Mexican species of Doswellia was described in 2012; [7] however, this species was subsequently transferred to the separate doswelliid genus Rugarhynchos . [8] Two additional dosweliids were named in 2013: Jaxtasuchus salomoni based on several skeletons found in the Ladinian-age Lower Keuper of Germany, [5] and Ankylosuchus chinlegroupensis based on fragments of four vertebrae, parts of the skull and of a limb bone from the early Carnian Colorado City Formation. [9]

Desojo, Ezcurra, & Schultz (2011)'s analysis placed Doswellidae as the closest large monophyletic clade to Archosauria, with only the Chinese archosauriform Yonghesuchus nested closer to archosaurs. [2] However, a phylogenetic analysis by Ezcurra (2016) recovered Doswelliidae alongside the family Proterochampsidae within the clade Proterochampsia, which was found to be the sister taxon of Archosauria. The unusual aquatic archosauriform Vancleavea was also referred to Doswelliidae in this analysis. [3] Subsequently, Ezcurra et al. (2017) excluded Archeopelta and Tarjadia from Doswelliidae, considering them to be archosaurs of the family Erpetosuchidae instead. [10] Litorosuchus , an aquatic archosauriform which is considered a close relative of Vancleavea, may also be a doswelliid if Vancleavea is a member of the family.

The following cladogram is after Ezcurra (2016): [3]

  Eucrocopoda  

Dorosuchus neoetus

Euparkeria capensis (Euparkeriidae)

Dongusuchus efremovi

Yarasuchus deccanensis

  Archosauria  

Avemetatarsalia

  Pseudosuchia  

Phytosauria

The rest of Pseudosuchia (incl. Koilamasuchus )

  Proterochampsia  
 Doswelliidae 

Vancleavea campi

Archeopelta arborensis

Tarjadia ruthae

Doswellia kaltenbachi

Jaxtasuchus salomoni

  Proterochampsidae  

Proterochampsa barrionuevoi

Proterochampsa nodosa

Cerritosaurus binsfeldi

Tropidosuchus romeri

  Rhadinosuchinae  

Pseudochampsa ischigualastensis

Gualosuchus reigi

Chanaresuchus bonapartei

Rhadinosuchus gracilis

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<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.

<span class="mw-page-title-main">Aetosaur</span> Extinct order of heavily armoured reptiles

Aetosaurs are heavily armored reptiles belonging to the extinct order Aetosauria. They were medium- to large-sized omnivorous or herbivorous pseudosuchians, part of the branch of archosaurs more closely related to crocodilians than to birds and other dinosaurs. All known aetosaurs are restricted to the Late Triassic, and in some strata from this time they are among the most abundant fossil vertebrates. They have small heads, upturned snouts, erect limbs, and a body ornamented with four rows of plate-like osteoderms. Aetosaur fossil remains are known from Europe, North and South America, parts of Africa, and India. Since their armoured plates are often preserved and are abundant in certain localities, aetosaurs serve as important Late Triassic tetrapod index fossils. Many aetosaurs had wide geographic ranges, but their stratigraphic ranges were relatively short. Therefore, the presence of particular aetosaurs can accurately date a site in which they are found.

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

Proterosuchidae is an early family of basal archosauriforms whose fossils are known from the Late Permian and the Early Triassic. The highest diversity of genera is known from European Russia, but fossils are also known from South Africa, India, China, Australia, Brazil and possibly Argentina. The name comes from Greek πρότερο- ("first") and σοῦχος ("crocodile").

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

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 proterochampsids. Doswellia was a low and heavily built carnivore which lived during the Carnian stage of the Late Triassic. It possesses many unusual features including a wide, flattened head with narrow jaws and a box-like rib cage surrounded by many rows of bony plates. The type species Doswellia kaltenbachi was named in 1980 from fossils found within the Vinita member of the Doswell Formation in Virginia. The formation, which is found in the Taylorsville Basin, is part of the larger Newark Supergroup. Doswellia is named after Doswell, the town from which much of the taxon's remains have been found. A second species, D. sixmilensis, was described in 2012 from the Bluewater Creek Formation of the Chinle Group in New Mexico; however, this species was subsequently transferred to a separate doswelliid genus, Rugarhynchos. Bonafide Doswellia kaltenbachi fossils are also known from the Chinle Formation of Arizona.

<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>Proterochampsa</i> Extinct genus of reptiles

Proterochampsa is an extinct genus of proterochampsid archosauriform from the Late Triassic of South America. The genus is the namesake of the family Proterochampsidae, and the broader clade Proterochampsia. Like other proterochampsids, Proterochampsa are quadruped tetrapods superficially similar in appearance to modern crocodiles, although the two groups are not closely related. Proterochampsids can be distinguished from other related archosauriformes by characters such as a dorsoventrally flattened, triangular skull with a long, narrow snout at the anterior end and that expands transversally at the posterior end, asymmetric feet, and a lack of postfrontal bones in the skull, with the nares located near the midline. Proterochampsa is additionally defined by characters of dermal sculpturing consisting of nodular protuberances on the skull, antorbital fenestrae facing dorsally, and a restricted antorbital fossa on the maxilla. The genus comprises two known species: Proterochampsa barrionuevoi and Proterochampsa nodosa. P. barrionuevoi specimens have been discovered in the Ischigualasto Formation in northwestern Argentina, while P. nodosa specimens have been found in the Santa Maria supersequence in southeastern Brazil. The two species are distinct in several characters, including that P. nodosa has larger, more well-developed nodular protuberances, a more gradually narrowing snout, and a higher occiput than P. barrionuevoi. Of the two, P. nodosa is thought to have less derived features than P. barrionuevoi.

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

Luperosuchus is an extinct genus of loricatan pseudosuchian reptile which contains only a single species, Luperosuchus fractus. It is known from the Chañares Formation of Argentina, within strata belonging to the latest Ladinian stage of the late Middle Triassic, or the earliest Carnian of the Late Triassic. Luperosuchus was one of the largest carnivores of the Chañares Formation, although its remains are fragmentary and primarily represented by a skull with similarities to Prestosuchus and Saurosuchus.

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>Chanaresuchus</i> Extinct genus of reptiles

Chanaresuchus is an extinct genus of proterochampsid archosauriform. It was of modest size for a proterochampsian, being on average just over a meter in length. The type species is Chanaresuchus bonapartei was named in 1971. Its fossils were found in from the early Carnian-age Chañares Formation in La Rioja Province, Argentina. Chanaresuchus appears to be one of the most common archosauriforms from the Chañares Formation due to the abundance of specimens referred to the genus. Much of the material has been found by the La Plata-Harvard expedition of 1964-65. Chanaresuchus is the most well-described proterochampsid in the subfamily Rhadinosuchinae.

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

Vancleavea is a genus of extinct, armoured, non-archosaurian archosauriforms from the Late Triassic of western North America. The type and only known species is V. campi, named by Robert Long & Phillip A Murry in 1995. At that time, the genus was only known from fragmentary bones including osteoderms and vertebrae. However, since then many more fossils have been found, including a pair of nearly complete skeletons discovered in 2002. These finds have shown that members of the genus were bizarre semiaquatic reptiles. Vancleavea individuals had short snouts with large, fang-like teeth, and long bodies with small limbs. They were completely covered with bony plates known as osteoderms, which came in several different varieties distributed around the body. Phylogenetic analyses by professional paleontologists have shown that Vancleavea was an archosauriform, part of the lineage of reptiles that would lead to archosaurs such as dinosaurs and crocodilians. Vancleavea lacks certain traits which are present in most other archosauriforms, most notably the antorbital, mandibular and supratemporal fenestrae, which are weight-saving holes in the skulls of other taxa. However, other features clearly support its archosauriform identity, including a lack of intercentra, the presence of osteoderms, an ossified laterosphenoid, and several adaptations of the femur and ankle bones. In 2016, a new genus of archosauriform, Litorosuchus, was described. This genus resembled both Vancleavea and more typical archosauriforms in different respects, allowing Litorosuchus to act as a transitional fossil linking Vancleavea to less aberrant archosauriforms.

<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.

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

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. Partial remains have been found from deposits that are Anisian-Ladinian in age. Long known mostly from osteoderms, vertebrae, and fragments of the skull, specimens described in 2017 provided much more anatomical details and showed that it was a fairly large predator. Tarjadia predates known species of aetosaurs and phytosaurs, two Late Triassic groups of crurotarsans with heavy plating, making it one of the first heavily armored archosaurs. Prior to 2017, most studies placed it outside Archosauria as a member of Doswelliidae, a family of heavily armored and crocodile-like archosauriforms. The 2017 specimens instead show that it belonged to the Erpetosuchidae.

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

Archeopelta is an extinct genus of carnivorous archosaur from the late Middle or early Late Triassic period. It was a 2 m (6 ft) long predator 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 archosaur.

<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.

Jaxtasuchus is an extinct genus of armored doswelliid archosauriform reptile known from the Middle Triassic of the Erfurt Formation in Germany. The type species, Jaxtasuchus salomoni, was named in 2013 on the basis of several incomplete skeletons and other isolated remains. Like other doswelliids, members of the genus were heavily armored, with four longitudinal rows of bony plates called osteoderms covering the body. Jaxtasuchus is the first doswelliid known from Europe and is most closely related to Doswellia from the Late Triassic of the eastern United States. However, it was not as specialized as Doswellia, retaining several generalized archosauriform characteristics and having less armor. Jaxtasuchus fossils have been found in aquatic mudstones alongside fossils of temnospondyl amphibians, crustaceans, and mollusks, suggesting that Jaxtasuchus was semiaquatic like modern crocodilians.

The Colorado City Formation is a Late Triassic geologic formation in the Dockum Group of Texas, United States. It has previously been known as the Iatan Member, Colorado City Member or 'Pre-Tecovas Horizon'.

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

Rugarhynchos is an extinct genus of doswelliid archosauriform from the Late Triassic of New Mexico. The only known species is Rugarhynchos sixmilensis. It was originally described as a species of Doswellia in 2012, before receiving its own genus in 2020. Rugarhynchos was a close relative of Doswellia and shared several features with it, such as the absence of an infratemporal fenestra and heavily textured skull bones. However, it could also be distinguished by many unique characteristics, such as a thick diagonal ridge on the side of the snout, blunt spikes on its osteoderms, and a complex suture between the quadratojugal, squamosal, and jugal. Non-metric multidimensional scaling and tooth morphology suggest that Rugarhynchos had a general skull anatomy convergent with some crocodyliforms, spinosaurids, and phytosaurs. However, its snout was somewhat less elongated than those other reptiles.

References

  1. Sues, Hans-Dieter; Desojo, Julia B.; Ezcurra, Martín D. (2013-01-01). "Doswelliidae: a clade of unusual armoured archosauriforms from the Middle and Late Triassic". Geological Society, London, Special Publications. 379 (1): 49–58. Bibcode:2013GSLSP.379...49S. doi:10.1144/SP379.13. ISSN   0305-8719. S2CID   129887410.
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  4. Bennett, S.C. (2020). "Reassessment of the Triassic archosauriform Scleromochlus taylori: neither runner nor biped, but hopper". PeerJ. 8: e8418. doi:10.7717/peerj.8418. ISSN 2167-8359.
  5. 1 2 3 Schoch, R. R.; Sues, H. D. (2013). "A new archosauriform reptile from the Middle Triassic (Ladinian) of Germany". Journal of Systematic Palaeontology. 12: 113–131. doi:10.1080/14772019.2013.781066. S2CID   85401373.
  6. R. E. Weems (1980). "An unusual newly discovered archosaur from the Upper Triassic of Virginia, U.S.A." Transactions of the American Philosophical Society. New Series. 70 (7): 1–53. doi:10.2307/1006472. JSTOR   1006472.
  7. Heckert, Andrew B.; Lucas, Spencer G.; Spielmann, Justin A. (2012). "A new species of the enigmatic archosauromorph Doswellia from the Upper Triassic Bluewater Creek Formation, New Mexico, USA". Palaeontology. 55 (6): 1333–1348. doi:10.1111/j.1475-4983.2012.01200.x.
  8. Brenen M. Wynd; Sterling J. Nesbitt; Michelle R. Stocker; Andrew B. Heckert (2020). "A detailed description of Rugarhynchos sixmilensis, gen. et comb. nov. (Archosauriformes, Proterochampsia), and cranial convergence in snout elongation across stem and crown archosaurs". Journal of Vertebrate Paleontology. 39 (6): e1748042. doi:10.1080/02724634.2019.1748042. S2CID   219917329.
  9. Lucas, S.G.; Spielmann, J.A.; Hunt, A.P. (2013). "A new doswelliid archosauromorph from the Upper Triassic of West Texas" (PDF). In Tanner, L.H.; Spielmann, J.A.; Lucas, S.G. (eds.). The Triassic System. New Mexico Museum of Natural History and Science Bulletin. Vol. 61. pp. 382–388. Archived from the original (PDF) on 2013-12-24. Retrieved 2016-05-02.
  10. Martín D. Ezcurra; Lucas E. Fiorelli; Agustín G. Martinelli; Sebastián Rocher; M. Belén von Baczko; Miguel Ezpeleta; Jeremías R. A. Taborda; E. Martín Hechenleitner; M. Jimena Trotteyn; Julia B. Desojo (2017). "Deep faunistic turnovers preceded the rise of dinosaurs in southwestern Pangaea". Nature Ecology & Evolution. 1 (10): 1477–1483. doi:10.1038/s41559-017-0305-5. PMID   29185518. S2CID   10007967.