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268 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY<br />

compression<br />

FIGURE 1.—Lateral view depicting the combination of forces in the upper bill<br />

of a bird (Struthio sp.) that produce resistance against the push from the mandible<br />

when grasping food. Lateral view. F'=useful force applied to object,<br />

R=rectractory (active muscular) force, N=push transferred to the braincase via<br />

dorsal upper-bill stalk.<br />

traction (backward shift) of the palate. Therefore, it is called<br />

the retracting or retractory force. The ancestors of birds apparently<br />

had no obvious source of such a force. Birds, however,<br />

have evolved the following three sources of retracting force.<br />

1. M. retractor palatini (Figure 2) is the ventromedial part of<br />

the pterygoid muscle (of Moller, 1930, 1931), which is rather<br />

large in tinamous (Figure 2 A,B; Dzerzhinsky, 1983; Elzanowski,<br />

1987), ostriches (Figure 2 D,E), and especially in Apteryx<br />

(Figure 2 C). In paleognaths (sensu Pycraft, 1900) this muscle<br />

usually originates on the pterygoid and on the rear end of the<br />

vomer. Its caudal attachment is not situated at the midline on<br />

the base of the braincase, as in many neognaths (sensu Pycraft,<br />

1900), but more laterally, near the caudal attachment of the occipito-mandibular<br />

ligament, i.e., on the medial part of ala tym-<br />

FlGURE 2.—Retractor palatini muscle in ventral view: A,B, tinamou, Eudromia elegans; c, Kiwi, Apteryx sp.<br />

(scale=5 mm); D,E, Ostrich, Struthio camelus. aca=aponeurosis of insertion of M. pterygoideus caudalis;<br />

ad'=aponeurose of insertion of M. depressor mandibulae that is related to M. retractor palatini; art=aponeurosis<br />

of insertion of M. retractor palatini; Pt=pterygoid; Rtp=M. retractor palatini; Vbm=vomer (A,C,D, superficial<br />

layer; B,E, deeper layers) (A,B, after Dzerzhinsky, 1983.)

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