Captorhinus
The outlines of the skulls of Captorhinus differ considerably from those of the skulls of the primitive captorhinomorph Protorothyris. Watson (1954:335, Fig. 9) has shown that in the morphological sequence, Protorothyris—Romeria—Captorhinus, there has been flattening and rounding of the skull-roof and loss of the primitive "square-cut" appearance in transverse section. The quadrates in Captorhinus are farther from the midline than in Protorothyris, and the adductor chambers in Captorhinus are considerably wider than they were primitively. Additionally, the postorbital region of Captorhinus is relatively longer than that of Protorothyris, a specialization that has increased the length of the chambers within.
In contrast with these dimensional changes there has been little shift in the pattern of the dermal bones that roof the adductor chambers. The most conspicuous modification in Captorhinus is the absence of the tabular. This element in Protorothyris was limited to the occiput and rested without sutural attachment upon the squamosal (Watson, 1954:338); later loss of the tabular could have had no effect upon the origins of muscles from inside the skull roof. Changes in pattern that may have modified the origin of the adductors in Captorhinus were correlated with the increase in length of the parietals and the reduction of the supratemporals. Other changes that were related to the departure from the primitive romeriid condition of the adductors included the development of a coronoid process, the flattening of the quadrate-articular joint, and the development of the peculiar dentition of Captorhinus.
The adductor chambers of Captorhinus are large. They are covered dorsally and laterally by the parietal, squamosal, postfrontal, postorbital, quadratojugal and jugal bones. The chamber extends medially to the braincase, but is not limited anteriorly by a bony wall. The occiput provides the posterior limit. The greater part of the adductor chambers lies mediad of the mandibles and thus of the Meckelian fossae; consequently the muscles that arise from the dermal roof pass downward and outward to their insertion on the mandibular rami.
Mandible
The mandibular rami of Captorhinus are strongly constructed. Each ramus is slightly convex in lateral outline. Approximately the anterior half of each ramus lies beneath the tooth-row. This half is roughly wedge-shaped in its lateral aspect, reaching its greatest height beneath the short posterior teeth.
The posterior half of each ramus is not directly involved in supporting the teeth, but is associated with the adductor musculature and the articulation of the ramus with the quadrate. The ventral margin of this part of the ramus curves dorsally in a gentle arc that terminates posteriorly at the base of the retroarticular process. The dorsal margin in contrast sweeps sharply upward behind the teeth and continues posteriorly in a long, low, truncated coronoid process.
A prominent coronoid process is not found among the more primitive members of the suborder, such as Limnoscelis, although the mandible commonly curves upward behind the tooth-row in that genus. This area in Limnoscelis is overlapped by the cheek when the jaw is fully adducted (Romer, 1956:494, Fig. 213), thereby foreshadowing the more extreme condition in Captorhinus.
The coronoid process in Captorhinus is not oriented vertically, but slopes inward toward the midline at approximately 45 degrees, effectively roofing the Meckelian fossa and limiting its opening to the median surface of each ramus. When the jaw was adducted, the coronoid process moved upward and inside the cheek. A space persisted between the process and the cheek because the process sloped obliquely away from the cheek and toward the midline of the skull. The external surface of the process presented an area of attachment for muscles arising from the apposing internal surface of the cheek.
Palate
The palate of Captorhinus is of the generalized rhynchocephalian type (Romer, 1956:71). In Captorhinus the pterygoids and palatines are markedly arched and the relatively large pterygoid flange lies almost entirely below the lower border of the cheek. The lateral edge of the flange passes obliquely across the anterior lip of the Meckelian fossa and abuts against the bottom lip of the fossa when the jaw is closed.
The palatines articulate laterally with the maxillary bones by means of a groove that fits over a maxillary ridge. This presumably allowed the halves of the palate to move up and down rather freely. The greatest amplitude of movement was at the midline. Anteroposterior sliding of the palate seems impossible in view of the firm palatoquadrate and quadrate-quadratojugal articulations.
The subtemporal fossa is essentially triangular, and its broad end is bounded anteriorly by the pterygoid flange. The fossa is lateral to much of the adductor chamber; consequently muscles arising from the parietals passed ventrolaterally, parallel to the oblique quadrate ramus of the pterygoid, to their attachment on the mandible.
Musculature
These osteological features indicate that the adductor muscles of the jaw in Captorhinus consisted of two primary masses (Figs. [1], [2], [3]). The first of these, the capitimandibularis, arose from the internal surface of the cheek and roof of the skull and inserted on the bones of the lower jaw that form the Meckelian canal and the coronoid process.
Fig. 1. Captorhinus. Internal aspect of skull, showing masseter, medial adductor, and temporal muscles. Unnumbered specimen, coll. of Robert F. Clarke. Richard's Spur, Oklahoma. × 2.
Fig. 2. Captorhinus. Internal aspect of skull, showing anterior and posterior pterygoid muscles. Same specimen shown in Fig. [1]. × 2.
The muscle was probably divided into a major medial mass, the temporal, and a lesser, sheetlike lateral mass, the masseter. The temporal was the largest of the adductors and arose from the lateral parts of the parietal, the dorsal parts of the postorbital, the most posterior extent of the postfrontal, and the upper parts of the squamosal. The muscle may have been further subdivided, but evidence for subordinate slips is lacking. The fibers of this mass were nearly vertically oriented in lateral aspect since the parts of the ramus that are available for their insertion lie within the anteroposterior extent of the adductor chamber. In anterior aspect the fibers were obliquely oriented, since the jaw and subtemporal fossa are lateral to much of the skull-roof from which the fibers arose.
The masseter probably arose from the quadratojugal, the jugal, and ventral parts of the squamosal, although scars on the quadratojugal and jugal are lacking. The squamosal bears an indistinct, gently curved ridge, passing upward and forward from the posteroventral corner of the bone and paralleling the articulation of the squamosal with the parietal. This ridge presumably marks the upper limits of the origin of the masseter from the squamosal.
Fig. 3. Captorhinus. Cross-section of right half of skull immediately behind the pterygoid flange, showing masseter, temporal, and anterior pterygoid muscles. Same specimen shown in Fig. [1]. × 2.
Fig. 4. Captorhinus. Internal aspect of left mandibular fragment, showing insertion of posterior pterygoid muscle. KU 8963, Richard's Spur, Oklahoma. × 2.8.
The masseter inserted on the external surface of the coronoid process, within two shallow concavities separated by an oblique ridge. The concavities and ridge may indicate that the muscle was divided into two sheets. If so, the anterior component was wedge-shaped in cross-section, and its thin posterior edge overlapped the larger mass that inserted on the posterior half of the coronoid process.
From a functional standpoint it is doubtful that a major component of the adductors arose from the quadrate wing of the pterygoid, for when the jaw is closed the Meckelian fossa is directly lateral to that bone. If the jaw were at almost any angle but maximum depression, the greatest component of force would be mediad, pulling the rami together and not upward. The mediad component would increase as the jaw approached full adduction. Neither is there anatomical evidence for an adductor arising from the quadrate wing of the pterygoid. The bone is smooth, hard, and without any marks that might be interpreted as muscle scars.
The internal adductor or pterygoid musculature in Captorhinus consisted of anterior and posterior components. The anterior pterygoid arose from the lateral edge and the dorsal surface of the pterygoid flange. The burred dorsal recurvature of the edge resembles that of the flange of crocodiles, which serves as part of the origin of the anterior pterygoid in those animals. In Captorhinus the attachment of the anterior pterygoid to the edge of the flange was probably tendinous, judging from the extent of the development of the edge of the flange. From the edge the origin extended medially across the dorsal surface of the flange; the ridging of this surface is indistinct, leading to the supposition that here the origin was more likely to have been fleshy than tendinous.
The anterior pterygoid extended obliquely backward and downward from its origin, passed medial to the temporal muscle and inserted on the ventral and medial surfaces of the splenial and angular bones beneath the Meckelian fossa. The spatial relationship between the palate and quadrate-articular joint indicate that the muscle was probably a minor adductor in Captorhinus.
When the jaw was adducted, the insertion of the anterior pterygoid was in a plane nearly level with the origin. Contraction of the anterior pterygoid when the jaw was in this position pulled the mandible forward and did not adduct it. Maximum depression of the mandible produced maximum disparity vertically between the levels of the origin and insertion. The force exerted by the anterior pterygoid upon the mandible when fully lowered most nearly approached the perpendicular to the long axes of the mandibular rami, and the resultant force acting on the mandible was adductive.
The adductive component of force therefore decreased as the jaw swung upward, with the result that the anterior pterygoid could only have been active in initiating adduction and not in sustaining it.
The evidence regarding the position and extent of the posterior pterygoid is more veiled. On the medial surface of the mandible, the prearticular and articular bones meet in a ridge that ventrally rims the glenoid cavity ([Fig. 4]). The ridge extends anteriorly and curves slightly in a dorsal direction and meets the Meckelian fossa. The curved part of the ridge is made of the prearticular bone alone. A small hollow above the ridge, anterior to the glenoid cavity, faces the medial plane of the skull and is bordered by the articular bone behind and above, and by the Meckelian fossa in front.
The surfaces of the hollow and the prearticular-articular ridge bear tiny grooves and ridges that seem to be muscle scars. The entire area of the hollow and its bordering features was probably the area of insertion of the posterior pterygoid.
However, the area of insertion lies mostly ventral to the articulating surface of the articular bone and extends but slightly in front of it. Seemingly little lever effect could be exercised by an adductor attaching in this position, namely, at the level of the fulcrum of the mandibular ramus.
The posterior pterygoid muscle probably arose from the anterior portion of the pterygoid wing of the quadrate, from a ridge on the ventromedial surface. From the relationship of the muscle to the articulation of the jaw with the skull, it may be deduced that the muscle was limited in function to the stabilization of the quadrate-articular joint by keeping the articular surfaces in close contact with each other and by preventing lateral slipping.
Finally there is evidence for an adductor between the temporal and masseter masses. The anterior dorsal lip of the Meckelian fossa supports a small knob to which this muscle attached, much as in Sphenodon (Romer, 1956:18, Fig. 12). Presumably the muscle was sheetlike and attached to the skull roof, medial to the attachment of the masseter.
A pseudotemporal may have been present, but evidence to indicate its extent and position is lacking. The muscle usually arises from the epipterygoid and nearby areas of the braincase and skull roof and inserts in the anterior parts of the fossa of the jaw. In Captorhinus the lateral wing of the pterygoid cuts across the fossa, effectively blocking it from the upper and medial parts of the skull, the areas of origin for the pseudotemporal.