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Vancleavea

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.

History
and Arizona, but the most complete specimens hail from the Coelophysis Quarry at Ghost Ranch. Vancleavea was first discovered in 1962 from the Petrified Forest Member of the Petrified Forest National Park and initially described by Long and Murry in 1995. At that time the only described specimen was the holotype, PEFO 2427. Since then, a number of remains have been found. A 2009 reevaluation of the genus by Nesbitt et al. formally described two additional specimens, GR 138 and 139. GR 138 is particularly notable due to being a nearly complete and articulated skeleton preserving a variety of osteoderms in the positions they would have been in during life. It was discovered at the Coelophysis Quarry in north-central New Mexico (Ghost Ranch), US, and was prepared at the Ruth Hall Museum of Paleontology in Abiquiú, New Mexico before its formal description. Vancleavea is a fairly common occurrence in most levels of the Chinle Formation, however, due to the poorly preserved remains, it is difficult to compare specimens across stratigraphic levels. ==Description==
Description
The most complete specimen of Vancleavea (GR 138) was around 1.2 m (3.9 ft) in length. However, isolated bones have shown that members of the genus could grow larger than GR 138. In particular, MCCDM 1745 from New Mexico may have reached a total length of . Imbricating osteoderms cover the entire body, the limbs are relatively short, and the skull is highly ossified. The nares open dorsally (i.e. nostrils face upward) and the jaw contains enlarged caniniform fangs. Each osteoderm possesses a pronounced central keel and an anterior projection. The ilium of Vancleavea resembles those of unrelated drepanosaurs. The rear edge of the skull is somewhat difficult to interpret even in well-preserved specimens such as GR 138. The upper rear corner of the temporal fenestra is formed by the squamosal bone, which connects to the back of the postorbital bone and the parietal bone, as well as the braincase. The front edge of the squamosal possesses a deep 'pocket', while the lower portion of the bone expands into a large downward-pointing 'hood'. A thin structure known as a ventral process stretches downwards, forming the rear edge of the temporal fenestra. This ventral process connects to the quadratojugal bone, which itself contacts the rear branch of the jugal and forms the rear lower corner of the temporal fenestra. The quadrate bone, which forms the cranium's contribution to the jaw joint, is located inwards from the squamosal's ventral process. The quadrate not only contacts the lower jaw, but also connects to the inner face of the quadratojugal and is overlapped by the 'hood' of the squamosal. The appearance of the quadrate is yet another unique aspect of Vancleavea. In most basal archosauriforms, the quadrate is tall and straight, but in Vancleavea it is short, stout, and arched forwards. Above the occipital condyle is the foramen magnum, a very large hole where the spinal cord exits the braincase. The upper rear portion of the braincase (as well as potentially the upper edge of the foramen magnum) is formed by the supraoccipital bone. As with other archosauriforms, the rear edge of this bone is acutely angled, terminating in a large keel edged by smooth areas for the attachment of muscles that raise the head. The side of the supraoccipital and a keel on the underside of the parietal bones meet along the side of the braincase, where they contact a downward-pointing spur known as the paroccipital process. The prootic bone, which forms the upper front part of the braincase, possesses a forward-pointing pit for the trigeminal nerve, with the area below the pit being smooth as in advanced archosauriforms. The front of the braincase also has a long and thick forward-pointing bone known as a laterosphenoid. This bone arches outwards but converges along the midline at its front tip. The dorsals (back vertebrae) are also long, and also possess "pinched" centra. Unlike the procoelous cervicals, the dorsals are amphicoelous with both the front and rear surface being concave. A pair of large and cylindrical outgrowths extend perpendicular to each centrum, at the front edge of their respective vertebrae. These outgrowths, currently identified as diapophyses, would have connected to ribs. Some dorsals of Vancleavea are also characteristic in the possession of a shallow groove which extends down the underside of each centrum, bordered by a pair of keels. A large, plate-like structure known as a neural spine juts out of the top of each vertebra. These neural spines are thin seen from the front but broad when seen from the side. They are roughly keystone-shaped, with notches at both the upper front and rear corner, as well as a convex upper portion covered with small grooves. The lower portion of the ilium is mostly formed by the acetabulum (hip socket), and has a triangular lower edge which wedges between the other two bones of the hip. The pubis is incomplete, but possesses a knob-shaped rear branch (posterior process) which extends under the acetabulum. This branch contacts the ischium along a straight edge, although part of their connection is left open as a notch which partially or completely enclosed the obturator foramen. The ischium is fan-shaped and points backwards and downwards. When seen from the front, the two ischia (on either side of the body) converge at the rear edges of each. Although the majority of the acetabulum is formed by the lower portion of the ilium, a noticeable portion is also formed by the upper portion of the ischium and a tiny sliver of the socket is formed by the upper portion of the pubis. Leg The top part of the S-shaped femur (thigh bone) is paddle-like and oval-shaped in cross-section. It includes a noticeable, but not strongly offset 'head'. The upper portion of the wide rear/inner face of the femur sometimes has a small ridge. This ridge, which is not present in every femur referred to Vancleavea, is believed to be an area for the attachment of the powerful caudofemoralis muscle which helps to pull the leg back. This ridge is believed to be synonymous with the fourth trochanter of archosaurs as well as the internal trochanter of basal archosauriforms. The midshaft of the femur is circular in cross-section. The lower portion of the femur includes both a medial condyle (which connects to the tibia) and a lateral condyle (which connects to the fibula), although neither of these knobs are particularly strongly developed. In general, the femur of Vancleavea is intermediate in structure between basal archosauriforms and archosaurs. The tibia and fibula (lower leg bones) are both slightly curved away from the center of the leg. They are also thinnest at their respective midshafts. The front edge of the tibia possesses a sharp ridge known as a cnemial crest, which extends about two thirds the way down the shaft. The fibula is thinner and somewhat twisted compared to the tibia. Ankle and foot view (i.e. with the tibial and fibular joints facing the viewer), with the calcaneum in blue and the astragalus in pink. Like most other archosauromorphs, the ankle is formed by two notable bones: the calcaneum, which connects to the fibula on the outer side of the heel, and the astragalus, which connects to both the tibia and the fibula on the inner part of the heel. The astragalus has facets for the tibia and fibula as well as a gap between the two facets, known as a non-articular notch. The facet for the tibia stretches down the inner edge of the astragalus, but not along the rear edge. This is similar to the situation in basal archosauriforms but contrasts with that of more advanced taxa which have a more sprawling tibial facet. The outer edge of the astragalus (which connects to the calcaneum) is also convex like basal archosauriforms. The calcaneum is small and triangular. The outer edge has a spur which points outwards and slightly downwards. Although this spur (formally known as a calcaneal tuber) is common in many archosauriforms, that of Vancleavea is unique in having a 'squashed' shape, with the upper/front and lower/rear surfaces meeting each other at a rounded outer apex. Similar to advanced archosauriforms, Vancleavea only possessed two distal tarsal bones in the ankle (apart from the calcaneum and astragalus), rather than four as in more basal members of the group. The main portion of the foot is formed by four thin metatarsal bones and a much shorter fifth metatarsal, each of which probably connect to a short toe. However, the bones of the foot are jumbled even in complete specimens such as GR 138, making it difficult to assign a specific location to any metatarsal or toe bone. Osteoderms The body is heavily armored, coated in various bony plates known as osteoderms. There are five to six distinct osteoderm morphotypes (i.e. variants) which are expressed at different parts of the body. Hunt, Lucas, & Spielmann (2005) were the first paleontologists to use morphotypes to describe Vancleavea osteoderms, but their designations differed from those in later studies such as Nesbitt et al. (2009), which are used here. The throat region, between the shoulder and jaws, possesses "morphotype A" osteoderms, which are teardrop-shaped and sometimes keeled, with a pointed front tip and rounded rear edge. "Morphotype B" osteoderms cover the back and sides of the body. They all tightly overlap and possess front spurs and low keels, but the osteoderms that form several rows on the back are more symmetrical and diamond-shaped. The osteoderms on the sides, on the other hand, are asymmetrical, with a smaller lower portion (below the keel) that slants forwards, a front spur that dips slightly downwards, and a rear tip that is more pointed than those of the back osteoderms. The osteoderms that cover the belly, known as "morphotype C" osteoderms, are of a type unique to Vancleavea. They are large, boxy, and rounded, with large keels and heavily serrated front edges. This morphotype has been compared to the armor of ankylosaurs. The upper side of the tail also possesses very characteristic and unusual osteoderms, designated as "morphotype D". These are elongated plates that stick up from the tail, with one plate per vertebra. They are wide at their base but flattened (from the side) elsewhere, and also acquire a swept-back orientation. In life, they would probably have been connected by soft tissue to form a tail fin to assist swimming. The morphotype D osteoderms are largest towards the front of the tail and gradually decrease in height and overall size towards the tip. The sides of the tail are covered with morphotype B osteoderms. The limbs are covered with "morphotype E" osteoderms. These plates are smaller, rounder and thinner than the osteoderms of the body. A sixth type of osteoderm, which may or may not belong to Vancleavea, was also reported in 1995. This osteoderm, of which only the base is known, is comparable to the tail spike of a Stegosaurus. Large, spike-like osteoderms were not present in GR 138, which was otherwise a complete skeleton with all of the other osteoderms in the same positions they were in life. It is conceivable that the "Stegosaurus spike" osteoderm was actually misplaced from the skeleton of an aetosaur, although the spikes of aetosaurs known from Vancleavea-bearing fossil sites differed from the "Stegosaurus spike". ==Classification==
Classification
}} }} Before the genus was described 1995, an additional specimen consisting of fragmentary skull material found by Charles Camp in 1923 was suggested to belong to a proterochampsid. However, this material was later referred to a new taxon, Acallosuchus rectori, after having been shown to be distinct from the postcranial material found in 1962. In the initial description of Vancleavea, the genus was referred to Neodiapsida incertae sedis along with A. rectori. Controversy still remains as to whether or not the specimens referred to this genus are representative of a "single species-level taxon or a clade of closely related taxa that lived through much of the Late Triassic of North America, given the poor fossil record of the taxon." However, the discovery of Litorosuchus, the closest relative of Vancleavea, has cast doubt on this hypothesis. ==Paleobiology==
Paleobiology
Vancleavea has features that suggest a semi-aquatic lifestyle. These include the long body, short limbs, and deep tail. Vancleavea is unique among archosauriforms and tetrapods in general in that the fin-like tail is deepened by elongated osteoderms rather than tall neural spines. ==References==
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