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NUMBER 89 271<br />

FIGURE 6.—Lateral surface of jaw adductors in lateral view: A, Rhea, Rhea<br />

americana; B, White-naped Crane, Grus vipio. Aex=M. adductor mandibulae<br />

extemus; Ap=M. adductor mandibulae posterior; Dm=depressor mandibulae<br />

muscle; Pss=pseudotem-poralis superficialis muscle; Pvl=ventrolateral portion<br />

of the pterygoid muscle. (B, after Kuular and Dzerzhinsky, 1994.)<br />

pression stresses mn from the bill tip through the premaxillary<br />

and maxillary bones to the palatine and then almost directly to<br />

the apex of the basipterygoid process. In the roof of the mouth,<br />

the ostrich possesses a broad gap that is closed only by skin.<br />

This patch of skin gains a gliding support from the parasphenoidal<br />

rostmm via a long, thin, anterior extension of the vomer.<br />

Source of the Medial Mandibular Process<br />

The processus mandibulae medialis is highly specific for<br />

Aves. For example, in Gobipteryx, a fossil Mongolian bird, Elzanowski<br />

(1974) regarded this process as a distinctly avian<br />

character. The functional properties of the muscular portion<br />

(ventromedial portion of the pterygoid muscle) inserting on its<br />

tip are influenced significantly by the particular position of the<br />

tip. It is placed extremely high in the sagittal plane, so corresponding<br />

muscular forces pass almost through the pivot of the<br />

quadrato-mandibular joint (Figure 9A) and therefore apply a<br />

negligible adductory component to the mandible as compared<br />

to the retractory one. In the frontal plane, the tip of the process<br />

is extremely close to the midline, and therefore those muscular<br />

forces tend to rotate the caudal part of mandibular branch and<br />

so expand the lower jaw as a whole (Figure 9B; Yudin, 1961).<br />

The functional conditions discussed above, however, do not<br />

seem to account for the first steps in the evolution of the medial<br />

mandibular process. There is a peculiarity in the paleognath<br />

jaw musculature that is more useful in this respect: the abovementioned<br />

M. retractor palatini. I suggest that this muscle may<br />

have arisen by a joining of two muscular units—the ventromedial<br />

portion of the pterygoid muscle and the depressor mandibulae<br />

muscle. Thus, their primitive interconnection via the caudal<br />

portion of the mandible formed a two-link chain that<br />

foreshadowed the recent M. retractor palatini (Figure 10). The<br />

cmcial event in their further evolution has been an optimization<br />

of their ability to exert a single force, which has been ensured<br />

by alignment of both muscular links, due to the displacement<br />

Ls Fr<br />

FIGURE 7.—Lateral view of tinamou skull (Tinamiformes), showing the loose articulation of the frontal with<br />

adjacent bones. Fr=frontal; Ls=laterosphenoid; Pa=parietal.

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