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Thrinaxodon is an extinct genus of which lived in what are now South Africa and Antarctica during the Early Triassic. The genus contains a single species, T. liorhinus.
Similar to other therapsids, Thrinaxodon adopted a semi-sprawling posture, an intermediary form between the sprawling position of basal and the more upright posture present in current mammals.Blob R. 2001. Evolution of hindlimb posture in non-mammalian therapsids: biomechanical tests of paleontological hypotheses. 27(1): 14-38. Thrinaxodon is prevalent in the fossil record, and one of the specimens represent the oldest known record of burrowing behavior among cynodonts.
On the skull roof of Thrinaxodon, the fronto-nasal suture represents an arrow shape instead of the general transverse process seen in more primitive skull morphologies. The prefrontals, which are slightly anterior and ventral to the frontals exhibit a very small size and come in contact with the post-orbitals, frontals, nasals and lacrimals. More posteriorly on the skull, the parietals lack a sagittal crest. The cranial roof is the narrowest just posterior to the parietal foramen, which is very nearly circular in shape. The temporal crests remain quite discrete throughout the length of the skull. The temporal fenestra have been found with ossified fasciae, giving evidence of some type of a temporal muscle attachment.
The upper jaw contains a secondary palate which separates the nasal passages from the rest of the mouth, which would have given Thrinaxodon the ability to breathe uninterrupted, even if food had been kept in its mouth. This adaptation would have allowed the Thrinaxodon to mash its food to a greater extent, decreasing the amount of time necessary for digestion. The maxillae and palatines meet medially in the upper jaw developing a midline suture. The maxillopalatine suture also includes a posterior palatine foramen. The large palatal roof component of the vomer in Thrinaxodon is just dorsal to the choana, or interior nasal passages. The pterygoid bones extend in the upper jaw and enclose small interpterygoid vacuities that are present on each side of the cultriform processes of the parasphenoids. The parasphenoid and basisphenoid are fused, except for the most anterior/dorsal end of the fused bones, in which there is a slight separation in the trabecular attachment of the basisphenoid.
The otic region is defined by the regions surrounding the temporal fenestrae. Most notable is evidence of a deep recess that is just anterior to the fenestra ovalis, containing evidence of smooth muscle interactions with the skull. Such smooth muscle interactions have been interpreted to be indicative of the tympanum and give the implications that this recess, in conjunction with the fenestra ovalis, outline the origin of the ear in Thrinaxodon. This is a new synapomorphy as this physiology had arisen in Thrinaxodon and had been conserved through late Cynodontia. The stapes contained a heavy cartilage plug, which was fit into the sides of the fenestra ovalis; however, only one half of the articular end of the stapes was able to cover the fenestra ovalis. The remainder of this pit opens to an "un-ossified" region which comes somewhat close to the cochlear recess, giving one the assumption that inner ear articulation occurred directly within this region.
The skull of Thrinaxodon is an important transitional fossil which supports the simplification of synapsid skulls over time. The most notable jump in bone number reduction had occurred between Thrinaxodon and Probainognathus, a change so dramatic that it is most likely that the fossil record for this particular transition is incomplete. Thrinaxodon contains fewer bones in the skull than that of its ancestors.
Upper incisors in T. liorhinus assume a backwards directed cusp, being curved and pointed at their most distal point, and becoming broader and rounder as they reach their proximal insertion point into the premaxilla. The fourth upper incisor is roughly homologous with a small canine tooth in form, but is positioned too far anteriorly to be a functional canine - thus ruling it out as an instance of convergent evolution. Lower incisors possess a very broad base, which is progressively reduced, heading distally towards the tip of the tooth. The lingual face of the lower incisors is most often concave while the labial face is often convex, and these lower incisors are oriented anteriorly, except in some cases for the third lower incisor, which can assume a more dorsoventral orientation. The incisors are, for the most part, single functional teeth encompassing a broad, cone-like morphology. The canines of T. liorhinus possess small dorsoventrally-directed facets on their surfaces, which appear to be involved with occlusion (dentition alignment in upper- and lower jaw closure). Each canine possesses a replacement canine located within the jaw, posterior to the existing canine, neither of the replacement or functional canine teeth possess any serrated margins only the small facets. It is important to note that the lower canine is directed almost vertically (dorsoventrally) while the upper canine is directed slightly anteriorly.
The upper and lower postcanines in T. liorhinus share some common features but also vary quite a fair amount in comparison to one another. The first postcanine (just posterior to the canine) is most often smaller than the other postcanines and is most often bicuspid. Including the first postcanine, if any of the other postcanines are bicuspid, then it is safe to assume that the posterior accessary cusp is present and that that tooth will not have any cingular or labial cusps. If, however, the tooth is tricuspid, then there is a chance of cingular cusps developing, if this occurs then the anterior cusp will be the first to appear and will be the most pronounced cusp. In the upper postcanines, there should be no occurrence of any teeth possessing more than three cusps, and there is no occurrence of any labial cusps on the upper postcanines. The majority of upper postcanines in the juvenile Thrinaxodon are bicuspid, while only one of these upper teeth are tricuspid. The upper postcanines of an intermediate (between juvenile and adult) Thrinaxodon are all tricuspid with no labial or cingular cusps. The adult upper postcanines retain the intermediate physiologies and possess only tricuspid teeth; however, it is possible for cingular cusps to develop in these adult teeth. The ultimate (posterior-most) upper canine is often the smallest of all canines in the entire jaw system. Little data is known of the juvenile and intermediate forms of the lower postcanines in Thrinaxodon, but the adult lower postcanines all possess multiple (any value more than three) cusps as well as the only appearance of labial cusps. Some older specimens have been found that possess no multiple-cups lower canines, possibly a response to old age or teeth replacement.
Thrinaxodon shows one of the first occurrences of replacement teeth in cynodonts. This was discerned by the presence of replacement pits, which are situated lingual to the functional tooth in the incisors and postcanines. While a replacement canine does exist, more often than not it is not erupted and the original functional canine remains.
Within the femur of Thrinaxodon, there is no major region of the bone that is made of parallel-fibred tissues; however, there is a small ring of parallel-fibred bone within the mid-cortex. The remainder of the femur is made of fibro-lamellar tissue; however, the globular osteocyte lacunae become much more organized and the primary osteons assume less vasculature than many other bones as you begin to approach the subperiosteal surface. The femur contains very few bony trabeculae. The humerus differs from the femur in many regards, one of which being that there is a more extensive network of bony trabeculae in the humerus near the meduallary cavity of the bone. The globular osteocyte lacunae become more flattened as you get closer and closer to the midshaft of the humerus. While the vasculature is present, the humerus contains no secondary osteons. The radii and ulnae of Thrinaxodon represent roughly the same histological patterns. In contrast to the humerii and femora, the parallel-fibred region is far more distinct in the distal bones of the forelimb. The medullary cavities are surrounded by multiple layers of very poorly vascularized endosteal lamellar tissue, along with very large cavities near the medullary cavity of the metaphyses.
Thrinaxodon was initially believed to be isolated to that region. Other fossils in South Africa were recovered from the Normandien and Katberg Formations. It had not been until 1977 that additional fossils of Thrinaxodon had been discovered in the Fremouw Formation of Antarctica. Upon its discovery there, numerous experiments were done to confirm whether or not they had found a new species of Thrinaxodontidae, or if they had found another area which T. liorhinus called home. The first experiment was to evaluate the average number of pre-sacral vertebrae in the Antarctic vs African Thrinaxodon. The data actually showed a slight difference between the two, in that the African T. liorhinus contained 26 presacrals, while the Antarctic Thrinaxodon had 27 pre-sacrals. In comparison to other cynodonts, 27 pre-sacrals appeared to be the norm throughout this sub-section of the fossil record. The next step was to evaluate the size of the skull in the two different discovery groups, and in this study they found no difference between the two, the first indication that they may in fact be of the same species. The ribs were the final physiology to be cross-examined, and while they portrayed slight differences in the expanded ribs, against one another, the most important synapomorphy remained consistent between the two, and that was that the intercostal plates overlapped with one another. These evaluations led to the conclusion that they had not found a new species of Thrinaxodontidae, but yet they had found that Thrinaxodon occupied two different geographical regions, which today are separated by an immense expanse of ocean. This discovery was one of many to support the idea of a connected land mass, and that during the early Triassic, Africa and Antarctica must have been linked in some way, shape or form.
The bone histology of Thrinaxodon indicates that it most likely had very rapid bone growth during juvenile development, and much slower development throughout adulthood, giving rise to the idea that Thrinaxodon reached peak size very early in its life.Botha J. Chinsamy A. 2005. Growth patterns of Thrinaxodon liorhinus, a non-mammalian cynodont from the lower Triassic of South Africa. Paleontology. 48(2): 385-394.
There is strong evidence that juvenile Thrinaxodon were cared for by their parents, as evidenced by fossilised aggregations of adults with multiple juvenile individuals of a similar size, likely indicating these juveniles were the same age and belonged to the same clutch.
The earliest discovery of a burrowing Thrinaxodon places the specimen found around 251 million years ago, a time frame surrounding the Permian–Triassic extinction event. Much of these fossils had been found in the flood plains of South Africa, in the Karoo Basin. This behavior had been seen at a relatively low occurrence in the pre-Cenozoic, dominated by therapsids, early-Triassic cynodonts and some early Mammalia. Thrinaxodon was in fact the first burrowing cynodont that has been found, showing similar behavioral patterns to that of Trirachodon. The first burrowing vertebrate on record was the dicynodont synapsid Diictodon, and it is possible that these burrowing patterns had passed on to the future cynodonts due to the adaptive advantage of burrowing during the extinction. The burrow of Thrinaxodon consists of two laterally sloping halves, a pattern that has only been observed in burrowing non-mammalian Cynodontia. The changes in vertebral/rib anatomy that arose in Thrinaxodon permit the animals to a greater range of flexibility, and the ability to place their snout underneath their hindlimbs, an adaptive response to small living quarters, in order to preserve warmth and/or for aestivation purposes.
A Thrinaxodon burrow contained an injured temnospondyl, Broomistega. The burrow was scanned using a synchrotron, a tool used to observe the contents of the burrows in this experiment, and not damage the intact specimens. The synchrotron revealed an injured Rhinesuchidae, Broomistega putterilli, showing signs of broken or damaged limbs and two skull perforations, most likely inflicted by the canines of another carnivore. The distance between the perforations was measured in relation to the distance between the canines of the Thrinaxodon in question, and no such relation was found. Therefore, we may assume that the temnospondyl found refuge in the burrow after a traumatic experience and the T. liorhinus allowed it to stay in its burrow until they both ultimately met their respective deaths. Interspecific shelter sharing is a rare anomaly within the fossil record; this T. liorhinus shows one of the first occurrences of this type of behavior in the fossil record, but it currently is unknown if the temnospondyl inhabited the burrow before or after the death of the nesting Thrinaxodon.
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