The Adductor Muscles of the Jaw In Some Primitive Reptiles

by Richard C. Fox

Dimetrodon

The morphology of the skull of Dimetrodon closely resembles that of the primitive Haptodus (Haptodontinae, Sphenacodontidae), and "hence may be rather confidently described as that of the family as a whole" (Romer and Price, 1940:285). The major differences between the two genera are in the increased specialization of the dentition, the shortening of the lacrimal, and the development of long vertebral spines in Dimetrodon. The absence of gross differences in the areas of the skull associated with the groups of muscles with which this study is concerned, implies a similarity in the patterns of musculature between the two groups. Romer and Price suggest that Haptodus, although too late in time to be an actual ancestor, shows "all the common features of the Dimetrodon group on the one hand and the therapsids on the other." The adductors of the jaw of Dimetrodon were probably little changed from those of the Haptodontinae and represent a primitive condition within the suborder.

Dimetrodon and Captorhinus differ in the bones associated with the adductor mechanism; the area behind the orbit in Dimetrodon is relatively shorter, reducing the comparative longitudinal extent of the adductor chamber. Furthermore, the dermal roof above the adductor chamber slopes gently downward from behind the orbit to its contact with the occipital plate in Dimetrodon. Temporal fenestrae are, of course, present in Dimetrodon.

Musculature

The adductor musculature of the lower jaw in Dimetrodon was divided into lateral and medial groups (Figs. [5], [6]). The lateral division consisted of temporal and masseter masses. The temporal arose from the upper rim of the temporal opening, from the lateral wall of the skull behind the postorbital strut, and from the dorsal roof of the skull. The bones of origin included jugal, postorbital, postfrontal, parietal and squamosal. This division may also have arisen from the fascia covering the temporal opening (Romer and Price, 1940:53). The muscle passed into the Meckelian fossa of the mandible and inserted on the angular, surangular, prearticular, coronoid and dentary bones. Insertion on the lips of the fossa also probably occurred.

The lateral division arose from the lower rim of the temporal opening and from the bones beneath. Insertion was in the Meckelian fossa and on the dorsal surface of the adjoining coronoid process.

Fig. 5. Dimetrodon. Internal aspect of skull, showing masseter and temporal muscles. Skull modified from Romer and Price (1940). Approx. × 1/4.

The reconstruction of the progressively widening masseter as it traveled to the mandible follows from the progressively widening depression on the internal wall of the cheek against which the muscle must have been appressed. The depressed surface included the posterior wing of the jugal, the whole of the squamosal, and probably the anteriormost parts of the quadratojugal. Expansion of the muscle rostrally was prevented by the postorbital strut that protected the orbit (Romer and Price, 1940:53).

The sphenacodonts possess the primitive rhynchocephalian kind of palate. In Sphenodon the anterior pterygoid muscle arises from the dorsal surface of the pterygoid bone and from the adjacent bones. A similar origin suggests itself for the corresponding muscle, the second major adductor mass, in Dimetrodon.

From the origin the muscle passed posterodorsad and laterad of the pterygoid flange. Insertion was in the notch formed by the reflected lamina of the angular, as suggested by Watson (1948).

In Dimetrodon the relationship of the dorsal surface of the palate and the ventromedial surface of the mandible in front of the articulation with the quadrate is unlike that in Captorhinus. When the mandible of Dimetrodon is at rest (adducted), a line drawn between these two areas is oblique, between 30 and 40 degrees from the horizontal. Depression of the mandible increases this angle. The insertion of the anterior pterygoid is thus always considerably below the origin, permitting the muscle to be active throughout the movement of the mandible, from maximum depression to complete adduction. This was a major factor in adding substantially to the speed and power of the bite.

The presence and extent of a posterior pterygoid is more difficult to assess, because of the closeness of the glenoid cavity and the raised ridge of the prearticular, and the occupancy of at least part of this region by the anterior pterygoid. In some specimens of Dimetrodon the internal process of the articular is double (see Romer and Price, 1940:87, Fig. 16) indicating that there was a double insertion here. Whether the double insertion implies the insertion of two separate muscles is, of course, the problem. Division of the pterygoid into anterior and posterior portions is the reptilian pattern (Adams, 1919), and such is adhered to here, with the posterior pterygoid arising as a thin sheet from the quadrate wing of the pterygoid and the quadrate, and inserting by means of a tendon on the internal process of the articular, next to the insertion of the anterior pterygoid.

Fig. 6. Dimetrodon. Internal aspect of right cheek, showing anterior and posterior pterygoid muscles. Skull modified from Romer and Price (1940). Approx. × 1/4.

Watson (1948) has reconstructed the musculature of the jaw in Dimetrodon with results that are at variance with those of the present study. Watson recognized two divisions, an inner temporal and an outer masseteric, of the capitimandibularis, but has pictured them (830: Fig. 4; 831: Fig. 5C) as both arising from the inner surface of the skull roof above the temporal opening. But in Captorhinus the masseter arose from the lower part of the cheek close to the outer surface of the coronoid process. Watson has shown (1948:860, Fig. 17B) the same relationship of muscle to zygoma in Kannemeyeria sp. It is this arrangement that is also characteristic of mammals and presumably of Thrinaxodon. In view of the consistency of this pattern, I have reconstructed the masseter as arising from the lower wall of the cheek beneath the temporal opening.

Watson's reconstruction shows both the temporal and masseter muscles as being limited anteroposteriorly to an extent only slightly greater than the anteroposterior diameter of the temporal opening. The whole of the posterior half of the adductor chamber is unoccupied. More probably this area was filled by muscles. The impress on the inner surface of the cheek is evident, and the extent of both the coronoid process and Meckelian opening beneath the rear part of the chamber indicate that muscles passed through this area.

Watson remarked (1948:829-830) that the Meckelian opening in Dimetrodon "is very narrow and the jaw cavity is very small. None the less, it may have been occupied by the muscle or a ligament connected to it. Such an insertion leaves unexplained the great dorsal production of the dentary, surangular and coronoid. This may merely be a device to provide great dorsal-ventral stiffness to the long jaw, but it is possible and probable that some part of the temporal muscle was inserted on the inner surface of the coronoid. Indeed a very well-preserved jaw of D. limbatus? (R. 105: Pl. I, Fig. 2) bears a special depressed area on the outer surface of the extreme hinder end of the dentary which differs in surface modelling from the rest of the surface of the jaw, has a definite limit anteriorly, and may represent a muscle insertion. The nature of these insertions suggests that the muscle was already divided into two parts, an outer masseter and an inner temporalis." But, unaccountably, Watson's illustration (1948:830, Fig. 4) of his reconstruction limits the insertion of the temporal to the anterior limit of the Meckelian opening and a part of the coronoid process above it. No muscle is shown entering the Meckelian canal. It seems more likely that the temporal entered and inserted in the canal and on its dorsal lips. The masseter inserted lateral to it, over the peak of the coronoid process, and overlapping onto the dorsalmost portions of its external face, as Watson has illustrated (Plate I, middle fig.).

I am in agreement with Watson's reconstruction of the origins for both the anterior and posterior pterygoid muscles. On a functional basis, however, I would modify slightly Watson's placement of the insertions of these muscles. Watson believed that the jaw of Dimetrodon was capable of anteroposterior sliding. The articular surfaces of the jaws of Dimetrodon that I have examined indicate that this capability, if present at all, was surely of a very limited degree, and in no way comparable to that of Captorhinus. The dentition of Dimetrodon further substantiates the movement of the jaw in a simple up and down direction. The teeth of Dimetrodon are clearly stabbing devices; they are not modified at all for grinding and the correlative freedom of movement of the jaw that that function requires in an animal such as Edaphosaurus. Nor are they modified to parallel the teeth of Captorhinus. The latter's diet is less certain, but presumably it was insectivorous (Romer, 1928). With the requisite difference in levels of origin and insertion of the anterior pterygoid in Dimetrodon insuring the application of force throughout the adduction of the jaws, it would seem that the whole of the insertion should be shifted downward and outward in the notch. If this change were made in the reconstruction, the anterior pterygoid would have to be thought of as having arisen by a tendon from the ridge that Watson has pictured (1948:828, Fig. 3) as separating his origins for anterior and posterior pterygoids. The posterior pterygoid, in turn, arose by tendons from the adjoining lateral ridge and from the pterygoid process of Romer and Price. Tendinous origins are indicated by the limitations of space in this area, by the strength of the ridges pictured and reported by Watson, and by the massiveness of the pterygoid process of Romer and Price.