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SOCIETY OF VERTEBRATE PALEONTOLOGY, MEMOIR 3<br />

1 Ornithischia<br />

FIGURE 39. Facial trends in Ornithopoda. In the course <strong>of</strong> ornithopod evolution, the external antorbital fenestra becomes reduced and eventually<br />

closed at the level <strong>of</strong> Hadrosauridae. The antorbital cavity also becomes reduced and eventually completely internalized, as in crocodylomorphs.<br />

These trends are associated with restriction <strong>of</strong> the main paranasal air sinus in connection with the elaboration <strong>of</strong> the masticatory apparatus and<br />

enlargement <strong>of</strong> the nasal vestibule. Skull drawings modified from Galton (1974), Norman (1986), Weishampel and Horner (1990), Weishampel<br />

and Witmer (1990b), and Sereno (1991a).<br />

independently show the same basic trends, namely, like crocodylomorphs,<br />

reduction <strong>of</strong> the antorbital cavity and closure <strong>of</strong><br />

the external antorbital fenestra (Fig. 39). The primitive ornithischian<br />

condition, manifested by Lesothosaurus diagnosticus,<br />

is to have a small internal antorbital fenestra (essentially the<br />

ostium <strong>of</strong> the antorbital paranasal air sinus) and a relatively<br />

large external antorbital fenestra. Basal ornithopods, such as<br />

Heterodontosaurus tucki, display the first signs <strong>of</strong> the trend in<br />

that lateral laminae from the maxilla and lacrimal constrict the<br />

external antorbital fenestra. These laminae are even more extensive<br />

in hypsilophodontids, and the external antorbital fenestra<br />

is relatively small. In basal iguanodontians, the antorbital<br />

cavity and external fenestra are further reduced and displaced<br />

caudally. Finally, in hadrosaurids, the external fenestra is completely<br />

closed (sometimes partly <strong>cover</strong>ed by the jugal), and the<br />

antorbital cavity is internalized and relatively small.<br />

An important concurrent trend here is the expansion <strong>of</strong> the<br />

feeding apparatus, in particular, the dentition and its bony buttresses<br />

(see Weishampel, 1984, 1993 and references therein).<br />

Ornithopods show functional innovations indicative <strong>of</strong> exten-<br />

sive oral processing in association with herbivory, such as a<br />

transverse power stroke (achieved independently through differelit<br />

mechanisms in heterodontosaurids and euornithopods).<br />

Aspects <strong>of</strong> this masticatory trend are increases in the number<br />

<strong>of</strong> teeth (but a decrease in the relative size <strong>of</strong> each tooth), their<br />

packing in the jaws, and the relative size <strong>of</strong> the maxilla, culminating<br />

in the characteristic dental batteries <strong>of</strong> hadrosaurids.<br />

Another trend worth noting takes place in Iguanodontia, and<br />

this relates to expansion <strong>of</strong> the nasal vestibule with enlargement<br />

<strong>of</strong> the naris and resultant caudal displacement <strong>of</strong> the antorbital<br />

cavity. (There is also a marked trend for size increase, but such<br />

allometric effects are too complex to merit laboring the discussion<br />

here.)<br />

Therefore, the relationship between the trends in the antorbital<br />

cavity (and its enclosed sinus) and in other anatomical<br />

systems is relatively straightforward. As the relative volume <strong>of</strong><br />

the dentition and its buttresses increases, the relative volume <strong>of</strong><br />

the antorbital paranasal air sinus and its bony cavity decreases;<br />

and, in iguanodontians, as the nasal vestibule expands, the antorbital<br />

cavity becomes further reduced. As in crocodylo-

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