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316 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY<br />

5mm<br />

B<br />

5mm<br />

FIGURE 3.—Fregata magnificens Mathews (Neognathae). The antorbital area of the skull in A, lateral, and B, dorsolaterocaudal<br />

views. Each scale bar=5 mm. (j=jugal, l=lacrimal, m=maxilla, n=nasal, nf=nasal foramen,<br />

pl=palatine, u=uncinate (lacrimopalatine).)<br />

naths and in nine families representing six neognathous orders,<br />

including Fregatidae (Pelecaniformes), Procellariidae and Diomedeidae<br />

(Procellariiformes), Laridae and Alcidae (Charadriiformes),<br />

Cariamidae (Gruiformes), Accipitridae (Falconiformes),<br />

and Cuculidae and Musophagidae (Cuculiformes)<br />

(Burton, 1970; pers. obs.). Monophyly of this set of taxa is out<br />

of the question, and four of the included orders (Pelecaniformes,<br />

Procellariiformes, Charadriiformes, and Gmiformes)<br />

have been repeatedly considered to be among the earliest neognathous<br />

branches. In addition, strong evidence of the presence<br />

of the uncinate has been found in Hesperomis (Elzanowski,<br />

1991). This suggests that the presence of the uncinate is plesiomorphic<br />

at least for the neomithines and that an ancestral bone<br />

was present in the primitive, pre-neomithine birds.<br />

The origin of the uncinate has remained obscure. Frank<br />

(1954:232) suggested that it is an avian neomorph. Burton<br />

(1970) raised the possibility of its derivation from a ligament<br />

(which is more widespread than the bone) but admitted that this<br />

begs the question of the origin of the ligament. McDowell<br />

(1978) mapped the uncinate to the archosaurian ectopterygoid<br />

because this bone appeared to be the only possible avian homologue<br />

for a reptilian element that lies lateral to the pterygoid<br />

and caudal to the maxilla. Thus far, no intermediate condition<br />

has been found. The oviraptorid ectopterygoid provides at least<br />

a stmctural intermediate between the reptilian ectopterygoid<br />

and the avian uncinate. Whether it is an evolutionary intermediate<br />

remains to be decided in a broader phylogenetic analysis.<br />

The similarities to the avian uncinate cannot be homologous<br />

unless the oviraptorids are more closely related to the ornithurine<br />

birds than is Archaeopteryx because the latter possesses a<br />

typical theropodan, hooked ectopterygoid, which is preserved<br />

in a caudal position in the fifth (Eichstatt) skeleton.<br />

BASIPTERYGOID ARTICULATION.—The pterygoid has a basal<br />

process for the articulation with the cranial base in oviraptorids<br />

(Figure 1), Hesperomis, and the majority of those neognaths<br />

that have a basipterygoid articulation (sensu stricto) as distinguished<br />

from the rostropterygoid articulation of the galliforms<br />

and anatids (Weber, 1993). The basal process is well developed<br />

in many Charadriiformes and some Caprimulgiformes (Steatomithidae,<br />

Caprimulgidae, Nyctibiidae) and in all the Tumi-

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