Doswellia Temporal range: Late Triassic, | |
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Life restoration of Doswellia kaltenbachi in mid-stride | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauromorpha |
Clade: | Archosauriformes |
Clade: | † Proterochampsia |
Family: | † Doswelliidae |
Genus: | † Doswellia Weems, 1980 |
Type species | |
Doswellia kaltenbachi Weems, 1980 |
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. [1] 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 (formerly known as the Falling Creek 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; [1] however, this species was subsequently transferred to a separate doswelliid genus, Rugarhynchos . [2] Bonafide Doswellia kaltenbachi fossils are also known from the Chinle Formation of Arizona. [3]
The most complete specimens of Doswellia were discovered in 1974 during the construction of a sewage treatment plant in Doswell, Virginia. A party led by James Kaltenbach (the namesake of Doswellia kaltenbachi) unearthed a large block containing a partial skeleton, including numerous vertebrae, ribs, osteoderms (bony plates), and other bones. This block, USNM 244214, has been designated the holotype of D. kaltenbachi. Two additional slabs were later unearthed at the same site and almost certainly pertained to the same individual. One of these slabs contained additional vertebrae and ribs while the other contained a partial skull and mandible. These additional slabs were collectively termed the paratype, USNM 214823. An isolated right jugal found at the site (USNM 437574) was also referred to the species. [4]
In the 1950s and 1960s, several additional bones (including vertebrae, osteoderms, a dentary, and a femur) were unearthed near Ashland, a little south of Doswell. These specimens were initially believed to have belonged to phytosaurs, but were recognized as pertaining to Doswellia after the Doswell specimens were found. The Ashland specimens are cataloged as USNM 186989 and USNM 244215. [5] Assorted osteoderms and vertebrae from the Otis Chalk and Colorado City Formation in Texas, as well as the Monitor Butte Formation in Utah have also been assigned to Doswellia. [6] [1] Fossils referable to Doswellia cf. kaltenbachi are known from Petrified Forest National Park in Arizona. They were found in the Blue Mesa Member of the Chinle Formation, making the Arizona remains among the youngest in the genus. [3]
In 2012, a new species of archosauriform was described and referred to Doswellia, as the second species Doswellia sixmilensis. The holotype of this species was NMMNH P-61909, an incomplete skeleton including skull fragments, osteoderms, vertebrae, and possible limb fragments. It was found in strata of the Bluewater Creek Formation exposed at Sixmile Canyon in McKinley County, New Mexico. [1] A 2020 redescription of "Doswellia" sixmilensis determined that its supposed snout fragments represented an entire skull of a related doswelliid. Once major anatomical differences were discovered, the species was given its own genus, Rugarhynchos , in 2020. [2]
Doswellia possesses many highly derived features in its skeleton. The skull is low and elongated with a narrow snout and wide temporal region behind the eye sockets. The temporal region is unusual in that it is euryapsid, which means that the lower of the two temporal holes on either side of the skull has closed. The jugal bone has expanded into the region the lower temporal opening would normally occupy. Paired squamosal bones extend beyond the skull's back margin to form small horn-like projections. The skull of Doswellia lacks several bones found in other archosauriforms, including the postfrontals, tabulars, and postparietals. [4]
The body of Doswellia is also distinctive. The neck is elongated and partially covered by a fused collection of bony scutes called a nuchal plate. The ribs in the front part of the torso project horizontally from the spine and then bend at nearly 90-degree angles to give the body of Doswellia a box-like shape. The blade-like ilium bone of the hip also projects horizontally. Rows of osteoderms stretched from the nuchal plate to the tail. At least ten rows covered the widest part of Doswellia's back. [4]
The quadratojugal and surangular bones (on the cranium and lower jaw, respectively) were both incorporated into the jaw joint, reinforcing the joint and preventing side-to-side or front-to back movement. As a result, the jaws of Doswellia would have been incapable of any notable form of movement other than vertical scissor-like snapping. In addition, the expanded back of the skull and very deep rear part of the lower jaw likely housed muscles that could let the jaw both open and close with a high amount of force. This contrasts with modern crocodilians, which have a powerful bite but a much weaker ability to open their jaws. Nevertheless, the high amount of sculpturing in the skull of Doswellia is similar to the skulls of modern crocodilians. It is likely an adaptation to minimize stresses in the skull during a powerful bite. [5]
The pointed teeth, long snout, and upward-pointing eyes of Doswellia are support for the idea that it was an aquatic carnivore. In addition, its relatively compact osteoderms are also evidence for an aquatic lifestyle. [7] However, it may not necessarily have been strictly aquatic, as these features are also found in animals such as Parasuchus , a phytosaur which is known to have preyed on terrestrial reptiles such as Malerisaurus . Other possible food sources include crustaceans, bivalves, and (most speculatively) large flying or hopping insects. It is also conceivable that it was capable of limited burrowing either for shelter (as in alligators) or defense, partially burying itself to keep its armor exposed yet protect its soft underside. This technique is used by modern armadillos, echidnas and Cordylus lizards. The front limbs are unknown in Doswellia, so there is no direct evidence for burrowing adaptations. [5]
The neck of Doswellia was long and flexible, although also heavily armored, so it was likely incapable of bending above the horizontal, instead probably being used more for downwards and lateral (side-to-side) movement. The body was also probably incapable of moving up and down to much of an extent due to the extensive and overlapping armor plating which characterizes the genus. The chest would have been much more likely to have flexed laterally (like most living reptiles and amphibians) while the animal was walking. The first few tail vertebrae were similar to the body vertebrae, so the front of the tail would probably have been held level with the body. The rest of the tail would have been more capable of bending downward, but lacked many adaptations for lateral movement. This means that, if Doswellia was an aquatic predator, it probably would not have used its tail for swimming as in modern crocodilians. Although limb material is not well known in Doswellia, material that is preserved suggests that both the front and rear legs were strongly built. Although the bizarre downward pointing hip could have given Doswellia an upright posture as in dinosaurs (including the armored ankylosaurs), various other primitive features suggest that it was more likely to have been sprawling or semi-sprawling most of the time. [5]
In 2017, an osteoderm from the Doswellia holotype was given a histological analysis to study growth patterns. The analysis concluded that the osteoderm formed by "intramembraneous ossification" due to the lack of structural fibers within it. This means that the bone of the osteoderm formed from a soft layer of periosteal tissue, rather than fibrous tendons or cartilage. Growth marks within the bone indicate that the holotype specimen of Doswellia died at 13 years of age. Perhaps the most unique aspect of Doswellia's osteoderm development lies in the fact that the ridges formed from the bone instead of the pits. In most prehistoric armored animals with pitted osteoderms, the pitting pattern formed due to specific spots of the bone being reabsorbed, creating pits. This even holds true to other doswelliids such as Jaxtasuchus . However, the studied osteoderm of Doswellia shows no evidence for reabsorption of specific areas, instead showing increased amounts of bone growth in the web of ridges which surround the pits. Although certain "rauisuchians" (non-crocodylomorph paracrocodylomorph archosaurs) also have osteoderms which form from bone growth in specific areas, their osteoderms are relatively smooth rather than pitted. Vancleavea , a supposed relative of Doswellia which also had its osteoderms analyzed, differed from the genus in multiple ways. [7]
The type species, Doswellia kaltenbachi, was described by Weems in 1980. [5] Weems placed Doswellia within Thecodontia, a group of archosaurs that traditionally included many Triassic archosaurs. He placed the genus within its own family, Doswelliidae, and suborder, Dosweliina. Parrish (1993) placed Doswellia among the most primitive of the crurotarsans, a group that includes crocodilians and their extinct relatives. More recently, Dilkes and Sues (2009) proposed a close relationship between Doswellia and the early archosauriform family Proterochampsidae. Desojo et al. (2011) added the South American archosauriforms Tarjadia and Archeopelta to Doswelliidae, and found support for Dilkes and Sues' classification in their own phylogenetic analysis. [8] Although Tarjadia and Archeopelta are now considered to be erpetosuchids only distantly related to Doswellia, the family Doswelliidae is still considered valid due to other taxa (such as Jaxtasuchus from Germany and Ankylosuchus from Texas) being considered close relatives of Doswellia.
Gracilisuchus is an extinct genus of tiny pseudosuchian from the Late Triassic of Argentina. It contains a single species, G. stipanicicorum, which is placed in the clade Suchia, close to the ancestry of crocodylomorphs. Both the genus and the species were first described by Alfred Romer in 1972.
Riojasuchus is an extinct genus of ornithosuchid archosaur from the Late Triassic (Norian) of Argentina. Ornithosuchidae was a widespread family of facultatively bipedal pseudosuchians with adaptations for scavenging. Riojasuchus is notable as one of the youngest and most complete members of the family. The type and only known species, Riojasuchus tenuisceps, was named and described by José Bonaparte in 1967. It was one of the first of many well-preserved Triassic archosaurs to be discovered in Argentina. The holotype specimen, PVL 3827, was found in the Los Colorados Formation of the Ischigualasto-Villa Unión Basin in northwestern Argentina.
Saurosuchus is an extinct genus of large loricatan pseudosuchian archosaurs that lived in South America during the Late Triassic period. It was a heavy, ground-dwelling, quadrupedal carnivore, likely being the apex predator in the Ischigualasto Formation.
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.
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.
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.
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.
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.
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.
Rhadinosuchus is an extinct genus of proterochampsian archosauriform reptile from the Late Triassic. It is known only from the type species Rhadinosuchus gracilis, reposited in Munich, Germany. The fossil includes an incomplete skull and fragments of post-cranial material. Hosffstetter (1955), Kuhn (1966), Reig (1970) and Bonaparte (1971) hypothesized it to be synonymous with Cerritosaurus, but other characteristics suggest it is closer to Chanaresuchus and Gualosuchus, while it is certainly different from Proterochampsa and Barberenachampsa. The small size indicates it is a young animal, making it hard to classify.
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.
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.
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.
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.
Pamelaria is an extinct genus of allokotosaurian archosauromorph reptile known from a single species, Pamelaria dolichotrachela, from the Middle Triassic of India. Pamelaria has sprawling legs, a long neck, and a pointed skull with nostrils positioned at the very tip of the snout. Among early archosauromorphs, Pamelaria is most similar to Prolacerta from the Early Triassic of South Africa and Antarctica. Both have been placed in the family Prolacertidae. Pamelaria, Prolacerta, and various other Permo-Triassic reptiles such as Protorosaurus and Tanystropheus have often been placed in a group of archosauromorphs called Protorosauria, which was regarded as one of the most basal group of archosauromorphs. However, more recent phylogenetic analyses indicate that Pamelaria and Prolacerta are more closely related to Archosauriformes than are Protorosaurus, Tanystropheus, and other protorosaurs, making Protorosauria a polyphyletic grouping.
Ankylosuchus is an extinct doswelliid archosauromorph reptile from the Late Triassic of Texas. Ankylosuchus is a monotypic genus and the type species is Ankylosuchus chinlegroupensis. It was named in 2013 on the basis of a fossil specimen including fragments of four vertebrae, parts of the skull, and part of a limb bone. These remains come from the Colorado City Formation, which dates to the early Carnian age of the Late Triassic. A. chinlegroupensis is named after the Chinle Group, a stratigraphic group that many Late Triassic formations of the southwestern United States have often been placed under, although a recent revision in stratigraphic nomenclature favors it being called the Dockum Group, which would make the species name a misnomer. Ankylosuchus is similar to other doswelliids in having heavy armor consisting of thick bony plates called osteoderms that interlock tightly and are irregularly pitted. It differs from other doswelliids in that the pits on the osteoderms are deeper and some osteoderms are fused to those that lie laterally to them.
Litorosuchus is a genus of armored, semiaquatic archosauriform reptile from the Middle Triassic of China, closely related to the morphologically similar Vancleavea. It contains one species, L. somnii.
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.
Syntomiprosopus is an extinct genus of archosauriform, possibly a crocodylomorph from the Late Triassic period of Arizona. The type and only known species is S. sucherorum. Syntomiprosopus was unusually short-snouted, comparable to the Late Cretaceous notosuchian Simosuchus, and is regarded as an example of convergent evolution between Triassic stem-archosaurs and Cretaceous archosaurs.