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Seymouria

Seymouria is an extinct genus of seymouriamorph from the Early Permian of North America and Europe. Although they were amphibians, Seymouria were well-adapted to life on land, with many reptilian features—so many, in fact, that Seymouria was first thought to be a primitive reptile. It is primarily known from two species, Seymouria baylorensis and Seymouria sanjuanensis. The type species, S. baylorensis, is more robust and specialized, though its fossils have only been found in Texas. On the other hand, S. sanjuanensis is more abundant and widespread. This smaller species is known from multiple well-preserved fossils, including a block of six skeletons found in the Cutler Formation of New Mexico, and a pair of fully grown skeletons from the Tambach Formation of Germany, which were fossilized lying next to each other.

History
Early history as a putative reptile Fossils of Seymouria were first found near the town of Seymour, in Baylor County, Texas (hence the name of the type species, Seymouria baylorensis, referring to both the town and county). The earliest fossils of the species to be collected were a cluster of individuals acquired by C.H. Sternberg in 1882. However, these fossils would not be properly prepared and identified as Seymouria until 1930. American paleontologist S.W. Williston later described a nearly complete skeleton in 1911, and noted that "Desmospondylus anomalus", a taxon he had recently named from fragmentary limbs and vertebrae, likely represented juvenile or even embryonic individuals of Seymouria. Likewise, English paleontologist D.M.S. Watson noted in 1918 that Conodectes, a dubious genera named by Edward Drinker Cope back in 1896, was likely synonymous with Seymouria. Robert Broom (1922) argued that the genus should be referred to as Conodectes since that name was published first, but Alfred Romer (1928) objected, noting that the name Seymouria was too popular within the scientific community to be replaced. During this time, Seymouria was generally seen as a very early reptile, part of an evolutionary grade known as "cotylosaurs", which also included many other stout-bodied Permian reptiles or reptile-like tetrapods. Proposed amphibian affinities Many paleontologists were uncertain about Seymouria's allegiance with the reptiles, noting many similarities with the embolomeres, which were unquestionably "labyrinthodont" amphibians. This combination of features from reptiles (i.e. other "cotylosaurs") and amphibians (i.e. embolomeres) was evidence that Seymouria was central to the evolutionary transition between the two groups. Regardless, not enough was known about its biology to conclude which group it was truly part of. Broom (1922) supported a placement among the Amphibia, but most studies around this time tentatively considered it an extremely "primitive" reptile; these included a comprehensive redescription of material referred to the species, published by Theodore E. White in 1939. However, indirect evidence that Seymouria was not biologically reptilian started to emerge by the 1940s. Around this time, several newly described genera were linked to Seymouria as part of the group Seymouriamorpha. Some seymouriamorphs, such as Kotlassia, had evidence of aquatic habits, and even Seymouria itself had occasionally been argued to possess lateral lines, sensory structures only usable underwater. and Romer (1947) each reversed their stance on Seymouria's classification, placing it among the amphibians rather than the reptiles. Perhaps the most damning evidence came in 1952, when Czech paleontologist Zdeněk Špinar reported gills preserved in juvenile fossils of the seymouriamorph Discosauriscus. This unequivocally proved that seymouriamorphs had an aquatic larval stage, and thus were amphibians, biologically speaking. Nevertheless, the numerous similarities between Seymouria and reptiles supported the idea that seymouriamorphs were close to the ancestry of amniotes. Additional species and fossils block containing six Seymouria sanjuanensis skeletons, described by Berman, Reisz, & Eberth (1987)|left In 1966, Peter Paul Vaughn described an assortment of Seymouria skulls from the Organ Rock Shale of Utah. These remains represented a new species, Seymouria sanjuanensis. Fossils of this species are now understood to be more abundant and widespread than those of Seymouria baylorensis. Several more species were later named by Paul E. Olson, although their validity has been more questionable than that of S. sanjuanensis. For example, Seymouria agilis (Olson, 1980), known from a nearly complete skeleton from the Chickasha Formation of Oklahoma, was reassigned by Michel Laurin and Robert R. Reisz to the parareptile Macroleter in 2001. Seymouria grandis, described a year earlier from a braincase found in Texas, has not been re-referred to any other tetrapod, but it remains poorly known. Langston (1963) reported a femur indistinguishable from that of S. baylorensis in Permian sediments at Prince Edward Island on the Eastern coast of Canada. Seymouria-like skeletal remains are also known from the Richards Spur Quarry in Oklahoma, as first described by Sullivan & Reisz (1999).File:Seymouria Tambach Lovers.jpg|thumb|318x318px|A cast of the "Tambach lovers" Seymouria sanjuanensis specimen first described by Berman et al. (2000) The Tambach Formation has also produced the developmentally youngest known fossils of Seymouria, assisting comparisons to Discosauriscus, which is known primarily from juveniles. == Description ==
Description
Seymouria individuals were robustly-built animals, with a large head, short neck, stocky limbs, and broad feet. The sensory apparatus of the skull also deserves mention for an array of unique features. The orbits (eye sockets) were about midway down the length of the skull, although they were a bit closer to the snout in juveniles. They were more rhomboidal than the circular orbits of other seymouriamorphs, with an acute front edge. The palate (roof of the mouth) had some similarities with both amniote and non-amniote tetrapods. On the one hand, it retained a few isolated large fangs with maze-like internal enamel folding, as is characteristic for "labyrinthodont" amphibians. On the other hand, the vomer bones at the front of the mouth were fairly narrow, and the adjacent choanae (holes leading from the nasal cavity to the mouth) were large and close together, as in amniotes. The palate is generally solid bone, with only vestigial interpteryoid vacuities (a pair of holes adjacent to the midline) separated by a long and thin cultriform process (the front blade of the base of the braincase). Apart from the fangs, the palate is also covered with small denticles radiating out from the rear part of the pterygoid bones. Seymouria has a few amniote-like characteristics of the palate, such as the presence of a prong-like outer rear branch of the pterygoid (formally known as a transverse flange) as well as an epipterygoid bone which is separate from the pterygoid. However, these characteristics have been observed in various non-amniote tetrapods, so they do not signify its status as an amniote. Postcranial skeleton The vertebral column is fairly short, with a total of 24 vertebrae between the hip and skull. In addition, later studies found that the atlas intercentrum was divided into a left and right portion, more like that of amphibian-grade tetrapods. Unlike almost all other Paleozoic tetrapods (amniote or otherwise), Seymouria completely lacks any bony remnants of scales or scutes, not even the thin, circular belly scales of other seymouriamorphs. The pectoral (shoulder) girdle has several reptile-like features. For example, the scapula and coracoid (bony plates which lie above and below the shoulder socket, respectively) are separate bones, rather than one large shoulder blade. Likewise, the interclavicle was flat and mushroom-shaped, with a long and thin "stem". The humerus (forearm bone) was shaped like a boxy and slightly twisted L, with large areas for muscle attachment. This form, which has been described as "tetrahedral", is plesiomorphic for tetrapods and contrasts with the slender hourglass-shaped humerus of amniotes. On the other hand, the lower part of the humerus also has a reptile-like adaptation: a hole known as an entepicondylar foramen. The radius was narrowest at mid-length. The ulna is similar, but longer due to the possession of a pronounced olecranon process, as is common in terrestrial tetrapods but rare in amphibious or aquatic ones. The carpus (wrist) has ten bones, and the hand has five stout fingers. The carpal bones are fully developed and closely contact each other, another indication of terrestriality. The phalanges (finger bones) decrease in size towards the tip of the fingers, where they each end in a tiny, rounded segment, without a claw. The phalangeal formula (number of phalanges per finger, from thumb to little finger) is 2-3-4-4-3. Two sacral (hip) vertebrae were present, though only the first one possessed a large, robust rib which contacted the ilium (upper blade of the hip). Some studies have argued that there was only one sacral vertebra, with the supposed second sacral actually being the first caudal due to having a shorter, more curved rib than the first sacral. Each ilium is low and teardrop-shaped when seen from the side, while the underside of the hip as a whole is formed by a single robust puboischiadic plate, which is rectangular when seen from below. Both the hip and shoulder sockets were directed at 45 degrees below the horizontal. The femur is equally stout as the humerus, and the tibia and fibula are robust, hourglass-shaped bones similar to the radius and ulna. The tarsus (ankle) incorporates 11 bones, intermediate between earlier tetrapods (which have 12) and amniotes (which have 8 or fewer). The five-toed feet are quite similar to the hands, with phalangeal formula 2-3-4-5-3. There were only about 20 caudal (tail) vertebrae at most. Past the base of the tail, the caudals start to acquire bony spines along their underside, known as chevrons. These begin to appear in the vicinity of the third to sixth caudal, depending on the specimen. Ribs are only present within the first five or six caudals; they are long at the base of the tail but diminish soon afterwards and typically disappear around the same area the chevrons appear. Differences between species Seymouria baylorensis and Seymouria sanjuanensis can be distinguished from each other based on several differences in the shape and connections between the different bones of the skull. For example, the downturned flange of bone above the otic notch (sometimes termed the "tabular horn" or "otic process") is much more well-developed in S. baylorensis than in S. sanjuanensis. In the former species, it acquires a triangular shape (when seen from the side) as it extends downwards more extensively towards the rear of the skull. In S. sanjuanensis, the postfrontal bone contacts the parietal bone by means of an obtuse, wedge-like suture, while the connection between the two bones is completely straight in S. baylorensis. Some authors have argued that the postparietals of S. baylorensis were smaller than those of S. sanjuanensis, but some specimens of S. sanjuanensis (for example, the "Tambach lovers") also had small postparietals. In addition, the "Tambach lovers" have a quadratojugal bone which is more similar to that of S. baylorensis rather than S. sanjuanensis. The combination of features from both species in these specimens may indicate that the two species are part of a continuous lineage, rather than two divergent evolutionary paths. Likewise, some differences relating to the proportions of the rear of the skull may be considered to be an artifact of the fact that most S. sanjuanensis specimens were not fully grown prior to the discovery of the "Tambach lovers", which were adult members of the species. Nevertheless, several traits are still clearly differentiated between the two species. The lacrimal bone, in front of the eyes, only occupies the front edge of the orbit in S. baylorensis. Conversely, specimens of S. sanjuanensis have a branch of the lacrimal which extends a small distance under the orbit. In S. sanjuanensis, much of the rear edge of the orbit is formed by the chevron-shaped postorbital bone, which is more rectangular in S. baylorensis. The shape of the lacrimal and postorbital of S. sanjuanensis closely corresponds to the condition in other seymouriamorphs, while the condition in S. baylorensis is more unique and derived. The tooth-bearing maxilla bone, which forms the side of the snout, is also distinctively unique in S. baylorensis. In S. sanjuanensis, the maxilla was low, with many sharp, closely spaced teeth extending along its length. This condition is similar to other seymouriamorphs. However, S. baylorensis has a taller snout, and its teeth are generally much larger, less numerous, and less homogenous in size. The palate is generally similar between the two species, although the ectopterygoids are more triangular in S. baylorensis and rectangular in S. sanjuanensis. ==Paleobiology==
Paleobiology
Lifestyle Romer (1928) was among the first authors to discuss the biological implications of Seymouria's skeleton. He argued that the robust limbs and wide-set body supported the idea that it was a strong, terrestrial animal with a sprawling gait. However, he also noted that Permian trackways generally support the idea that terrestrial tetrapods from this time period were not belly-draggers, but instead were strong enough to keep their bodies off of the ground. As with other paleontologists around the time, Romer assumed that Seymouria had a reptilian (or amniote) mode of reproduction, with eggs laid on dry land and protected from the elements by an amnion membrane. ==References==
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