A therapsid is a member of the clade Therapsida which is a major group of eupelycosaurian synapsids that includes mammals and their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more underneath the body, as opposed to the sprawling posture of many reptiles and salamanders.
Therapsids evolved from "pelycosaurs", specifically within the Sphenacodontia, more than 279.5 million years ago. They replaced the "pelycosaurs" as the dominant large land animals in the Guadalupian through to the Early Triassic. In the aftermath of the Permian–Triassic extinction event, therapsids declined in relative importance to the rapidly diversifying reptiles during the Middle Triassic.
The therapsids include the cynodonts, the group that gave rise to mammals (Mammaliaformes) in the Late Triassic, around 225 million years ago. Of the non-mammalian therapsids, only cynodonts survived beyond the end of the Triassic, with the only other remaining group of therapsids to have survived into the Late Triassic, the dicynodonts, becoming extinct towards the end of the period. The last surviving group of non-mammaliaform cynodonts were the Tritylodontidae, which became extinct during the Early Cretaceous.
Characteristics
Jaw and teeth
Therapsids' temporal fenestrae were larger than those of the pelycosaurs. The jaws of some therapsids were more complex and powerful, and the teeth were differentiated into frontal incisors for nipping, great lateral canines for puncturing and tearing, and molars for shearing and chopping food.
Posture
Therapsid legs were positioned more vertically beneath their bodies than were the sprawling legs of reptiles and pelycosaurs. Also compared to these groups, the feet were more symmetrical, with the first and last toes short and the middle toes long, an indication that the foot's axis was placed parallel to that of the animal, not sprawling out sideways. This orientation would have given a more mammal-like gait than the lizard-like gait of the pelycosaurs.
Physiology
The physiology of therapsids is poorly understood. Most Permian therapsids had a pineal foramen, indicating that they had a parietal eye like many modern reptiles and amphibians. The parietal eye serves an important role in thermoregulation and the circadian rhythm of ectotherms, but is absent in modern mammals, which are endothermic. Near the end of the Permian, dicynodonts, therocephalians, and cynodonts show parallel trends towards loss of the pineal foramen, and the foramen is completely absent in probainognathian cynodonts. Evidence from oxygen isotopes, which are correlated with body temperature, suggests that most Permian therapsids were ectotherms and that endothermy evolved convergently in dicynodonts and cynodonts near the end of the Permian. In contrast, evidence from histology suggests that endothermy is shared across Therapsida, whereas estimates of blood flow rate and lifespan in the mammaliaform Morganucodon suggest that even early mammaliaforms had reptile-like metabolic rates. Evidence for respiratory turbinates, which have been hypothesized to be indicative of endothermy, was reported in the therocephalian Glanosuchus, but subsequent study showed that the apparent attachment sites for turbinates may simply be the result of distortion of the skull.
Integument
The evolution of integument in therapsids is poorly known, and there are few fossils that provide direct evidence for the presence or absence of fur. The most basal synapsids with unambiguous direct evidence of fur are docodonts, which are mammaliaforms very closely related to crown-group mammals. Fossilized facial skin from the dinocephalian Estemmenosuchus has been described as showing that the skin was glandular and lacked both scales and hair.
Coprolites containing what appear to be hairs have been found from the Permian. Though the source of these hairs is not known with certainty, they may suggest that hair was present in at least some Permian therapsids.
The closure of the pineal foramen in probainognathian cynodonts may indicate a mutation in the regulatory gene Msx2, which is involved in both the closure of the skull roof and the maintenance of hair follicles in mice. This suggests that hair may have first evolved in probainognathians, though it does not entirely rule out an earlier origin of fur.
