The ancestry of the Theromorpha themselves is also shrouded in mystery. Attempts have been made to connect them with the Permian Protorosaurus, Palaeohatteria, and Eryops. On the other hand, some retain various Stegocephalous reminiscences (e.g. the roofed-in condition of the skull by membrane-bones, amongst which, besides others, supratemporals and postorbitals can be recognised; occurrence of cleithra in Pareiasaurus; distinct epiotic bones in Elginia). Although they have died out as a group, they have perhaps given rise to several side-branches, one of which (leaving aside the question of Mammalian origin) seems to have flourished as the Dinosauria.
We divide the Theromorpha into four orders, which are, however, liable to run into each other, and it is reasonably to be hoped that many forms may be discovered which will connect not only these provisional orders with each other, but also with other sub-classes.
Order I. PAREIASAURI
Cranium completely roofed in by membrane-bones. The only foramina are the nostrils, orbits, and the interparietal foramen. The teeth are comparatively small, and stand in even series in both jaws.
Pareiasaurus, several species from the Karroo sandstone of South Africa. P. baini was an extremely clumsy brute, of most uncouth appearance, standing between 2 and 3 feet high, and measuring with the short tail nearly 8 feet in length. The skull is very massive, 18 inches long and slightly broader, with a rugose, deeply pitted surface. The teeth are thickly enamelled, serrated at the margin, with many pointed cusps; those of the vomer, palatines, and pterygoids are recurved and arranged in several longitudinal rows. There is a small incisive foramen in the premaxilla; the choanae lie within the pterygoids. The palate has a pair of large lateral vacuities. Between the squamosal and quadrate is a small foramen, as in Belodon and Sphenodon. The nares are terminal, bordered behind by the nasals, and divided by the premaxillaries. The occipital condyle is a single knob, but the lateral occipital bones also partake in its formation. The shoulder-girdle is strong. The scapula slants backwards, is broad, and possesses a longitudinal spine, an almost exclusively Mammalian character. The scapula, coracoid and precoracoid are fused together, and are united ventrally with those of the other side. There is a T-shaped interclavicle, a pair of clavicles, and a pair of slender, long cleithra, which extend along the upper anterior margin of the scapulae. The humerus possesses enormous crests. The broad ilium is attached to two, or perhaps three, sacral ribs. The acetabulum is closed. The pubes and ischia are united into one broad mass of bone, and the obturator-foramina seem to be just large enough to permit of the passage of the nerve. Both fore- and hind-limbs are plantigrade and five-toed. The tibia articulates with one large bone, which is supposed to represent the united astragalus and calcaneum, the latter being without an indication of a prominent heel, although there is a tendency to develop the crurotarsal into the chief joint. The number of vertebrae amounts to eighteen presacrals, eight to ten of which are cervicals. There are two or three sacral and about twenty-four mostly shortened caudal vertebrae. The latter possess intercentral wedges and chevron-bones; wedges occur also between the cervical and some thoracic vertebrae. Some of the posterior cervical ribs are very peculiar–straight, broadened out, turned backwards, partly overlapped by one another, and 18 inches long, recalling the first two ribs of the crocodiles. Sternum and abdominal ribs are unknown.
Elginia mirabilis.–The skull (Fig. 54, A, p. [280])–nothing else is known–indicates one of the most remarkable reptiles hitherto found on this side of the Atlantic. It was discovered in the Red Sandstone of Elgin (Lower Trias). The skull reminds us in its general shape and by its spikes and horns of the little American Iguanoid lizard, Phrynosoma. The length of the cranium is about 6 inches, the distance between the tips of the two largest horns measures 9 inches. The teeth are small and resemble those of an Iguana in their shape and finely serrated edges, indicating herbivorous habits, but there are also several rows of smaller teeth on the palate, the configuration of which is not unlike that of Sphenodon. The top and sides of the skull, except the interparietal foramen, the orbits, and nostrils, are completely encased by rugose, pitted, dermal bones, most of them with strange, horn-like spikes. In the encasement of the temporal region can be discerned a postfrontal, parietal and squamosal, a conically projecting epiotic, a postorbital and supratemporal, a jugal and a quadrato-jugal, which latter almost completely covers the quadrate bone. The interparietal foramen lies far forwards, almost on a level with the orbits. The nostrils are terminal, surrounded by the short nasals, the maxillaries and the premaxillaries, which latter divide them.
Order II. THERIODONTIA.
The cranium is not roofed in, but shows a pair of large supratemporal fossae, bordered below by the zygoma, which is formed mainly by the squamoso-jugal bridge, and is shut off from the orbit by the postfrontal joining the bridge. The teeth are differentiated into incisors, canines, and molars (Fig. 54, C, p. [280]). The lower canines close in front of the upper.
Cynognathus, Karroo formation of South Africa. C. crateronotus has a skull about 16 inches long, looking like that of a ferocious Carnivore; there are four incisors, huge canines, and nine molars, the latter with serrated edges and anterior and posterior cusps. The wide supratemporal fossa is bordered and closed behind by the broad lateral extension of the parietal, which joins a similar extension of the squamosal bone. The latter is very long, extending to the postfrontal and to a bone which, bordering the orbit posteriorly, is either an upward branch of the jugal, or a postorbital bone; the latter interpretation is made probable by the occurrence of a suture with the jugal in C. platyceps. The jugal bone is very long, beginning at the quadrate, running along the squamosal, and forming the lower border of the orbit.
The number of vertebrae is large, there being as many as twenty-nine presacrals, six of which belong to the cervical region. The atlas is fused with the axis; most of the thoracic ribs articulate partly upon the intercentra. The lumbar ribs are very peculiar; they are much expanded horizontally, and overlap each other, forming thereby intercostal foramina. The broad ilium is attached to three or four sacral ribs. The acetabulum is closed. The ventral side of the pelvis shows a broad symphysis and has a pair of obturator-foramina. The scapula is large, directed backwards, and shows a distinct, very Mammalian spine; it is fused with the coracoid and precoracoid.