46 SOCIETY OF VERTEBRATE PALEONTOLOGY, MEMOIR 3 ing into chambers within the nasal bone. As noted previously, these nasal recesses appear to be associated with the excavatio pneumatica in the maxillary ascending ramus. Currie and Zhao (1994a) noted that the two cavities <strong>of</strong> S. dongi are extensive but do not communicate. Monolophosaurus jiangi (Zhao and Currie, 1994) deserves special mention in that its nasal bones form a large hollow crest with two large apertures caudally that actually pass from one side to the other, establishing a contralateral communication between the antorbital cavities, as well as invading the body <strong>of</strong> the nasal; furthermore, two additional, more rostral, foramina pneumatize the nasal bone and then continue rostrally to pneumatize the premaxillae. Nasal pneumatic recesses have not been described for other theropods (except for oviraptorosaurs; see below), and they can be shown to be positively absent in Tyrannosauridae, Ornithomimosauria, Troodontidae, Erlikosaurus andrewsi (PST 10011 11; Clark et al., 1994), and Archaeopteryx lithographica (cast <strong>of</strong> Eichstatt specimen). As with the lacrimal and jugal recesses, nasal recesses are described in Deinonychus antirrhopus (YPM 5210, 5232; MOR 747) here for the first time (see also Witmer, 199%; Witmer and Maxwell, 1996). Ostrom (1969:19) described a "groove with three moderate-sized, oval foramina" within the nasal <strong>of</strong> D. antirrhopus but had "no explanation for this pattern." Comparison with other theropods (as well as further preparation) reveal that Ostrom's groove is the nasal antorbital fossa and the "oval foramina" are pneumatic foramina leading into small cavities. Palatine recesses are found in some theropods and typically are cavities or foramina in the dorsal surface <strong>of</strong> the palatine bone that are clearly associated with the antorbital cavity. These are absent in Ceratosauria and some large tetanurans such as Allosaurus fragilis. Where present, they invariably are caudal or caudolateral to the choana, just medial to the contact with the maxilla, and rostrolateral to the presumed pterygoideus fossa, if present. In Archaeopteryx sp. (cast <strong>of</strong> Solenh<strong>of</strong>er Aktien- Verein specimen; see also Elzanowski and Wellnh<strong>of</strong>er, 1996), the dromaeosaurids Deinonychus antirrhopus (YPM 5210, 5232; Fig. 32) and Velociraptor mongoliensis (cast <strong>of</strong> PIN 314318; Osm6lska, 1985), and the troodontid Saurornithoides mongoliensis (AMNH 6516), there is a well-developed fossa in this position, and, in at least D. antirrhopus, this fossa leads into a small cavity. In Sinraptor dongi, the fossa surrounds a moderately large foramen that leads into a cavity within the choanal process (Currie and Zhao, 1994a). Tyrannosaurids have <strong>of</strong>ten large foramina in the dorsal surface <strong>of</strong> the palatine that lead into a strutted cavity with the bone (Molnar, 1991; Carr, 1996). There may be just a single large foramen leading into the bone (e.g., Daspletosaurus torosus, CMN 8506; Tyrannosaurus rex, AMNH 5027) or sometimes two or more (e.g., Nanotyrannus lancensis, Gilmore, 1946; Albertosaurus sarcophagus, CMN 5601; Tarbosaurus bataar, Maleev, 1974). Few theropod taxa can be scored for palatine recesses, and thus the phylogenetic distribution <strong>of</strong> these features is problematic. If it were not for the clear presence <strong>of</strong> palatine recesses in Sinraptor dongi, it might have been possible to regard them as a synapomorphy <strong>of</strong> at least Maniraptora (assuming Holtz's [I9941 placement <strong>of</strong> Tyrannosauridae is correct) if not Coelurosauria. A discussion <strong>of</strong> the facial structure <strong>of</strong> theropods would not be complete without special mention <strong>of</strong> the peculiar Mongolian oviraptorosaurs (Barsbold et al., 1990). Oviraptor philoceratops and its relatives clearly manifest the osteological correlates permitting the inference <strong>of</strong> an antorbital air sac. Furthermore, their facial skeletons have been described as being extensively pneumatic by virtually all recent workers (Osmblska, 1976; Barsbold, 1983; Barsbold et al., 1990). In fact, the facial bones in some species are developed into a tall, cassowary-like crest formed by a fine lattice <strong>of</strong> bony trabeculae (Barsbold et al., 1990). Oviraptorosaurs were not discussed much in the above analysis <strong>of</strong> theropod accessory cavities because their skulls are so transformed that it is not always easy to identify homologous pneumatic structures. Virtually all <strong>of</strong> the facial elements have complex cavities within them. The ones in the maxilla, lacrimal, and nasal are associated with the antorbital cavity (Osm6lska, 1976), whereas the cavities within the premaxilla and others in the nasal and even the frontal bone are apparently connected with the nasal vestibule (Barsbold et al., 1990). Coelophysis bauri also merits further discussion in that it lacks all <strong>of</strong> the cavities and chambers in the facial bones, thus strongly resembling many non-dinosaurian archosaurs; yet it is regarded as a derived member <strong>of</strong> its clade, Ceratosauria (Rowe and Gauthier, 1990; Holtz, 1994). Since more basal ceratosaurians such as Ceratosaurus nasicornis and Dilophosaurus wetherilli are large-skulled forms manifesting some~<strong>of</strong> these cavities (and, presumably, pneumatic diverticula <strong>of</strong> an antorbital air sac), it is reasonable to suggest that C. bauri also possessed a large paranasal air sinus, but, by virtue <strong>of</strong> small skull size, did not develop pneumatic diverticula into its facial bones. Therefore, some aspects <strong>of</strong> facial pneumaticity may have a size-related (i.e., allometric) component. Sauropodomorpha-In the prosauropod Plateosaurus engelhardti (AMNH 6810), the nasal bone sends a lamina lateral to the maxillolacrimal contact that broadly overhangs the antorbital cavity (Fig. 12A). In fact, there is large, dorsal, C-shaped (laterally open) hiatus framed by the laminae <strong>of</strong> the maxilla and lacrimal that the nasal caps (Fig. 12D). That portion <strong>of</strong> the nasal ro<strong>of</strong>ing the aperture has a deeply excavated recess (Fig. 12C), and may be the closest approach among sauropodomorphs to a theropod-like bony accessory cavity. It is not known how widely this nasal recess is distributed among prosauropods, but since the nasal overhangs the antorbital cavity in Sellosaurus gracilis (Galton, 1985b) and perhaps Massospondylus carinatus (Gow et al., 1990), the recess also may be present in these forms. Ornithischia-The palatine <strong>of</strong> Lesothosaurus diagnosticus has two well-marked fossae on its dorsolateral surface. The caudal fossa was interpreted earlier as a muscular fossa. The rostral fossa (Fig. 7C, D), however, appears to be associated with the antorbital cavity and may well be a pneumatic fossa. It resembles in some respects the palatine fossa noted above for the small theropods Deinonychus antirrhopus (Fig. 32) and Velociraptor mongoliensis. Haubold (1990) described a deep rostral extension <strong>of</strong> the antorbital cavity within the maxilla <strong>of</strong> the basal thyreophoran Emausaurus ernsti (Fig. 18), which could be an accessory cavity. Similarly, in another basal thyreophoran, Scelidosaurus harrisonii, there is a moderately large foramen at the rostral apex <strong>of</strong> the maxillary antorbital fossa that could lead to an accessory cavity, but it is unknown whether it expands into a chamber within the bone. Clearer evidence <strong>of</strong> accessory cavities is available for protoceratopsians (Fig. 20C, D). In at least Bagaceratops rozhdestvenskyi and Protoceratops andrewsi, an accessory cavity is associated with the maxillary antorbital fossa (Maryanska and Osm6lska, 1975; Osm6lska, 1986). This "intramaxillary sinus" enters the bone via a slit or cleft (sometimes two slits as in AMNH 6466) within the floor <strong>of</strong> the maxillary recess. It subsequently expands, forming a cavity running the length <strong>of</strong> the maxilla dorsal to the tooth roots (Osm6lska, 1986; Fig. 20C, D). It is not known whether it occurs in other protoceratopsians, but Sternberg (1951:232) regarded the antorbital fossa (his "maxillary sinus") as being "much deeper" in Leptoceratops gracilis than in P. andrewsi, perhaps suggesting the presence <strong>of</strong> an accessory sinus in this form as well. Pterosauria-Many large pterodactyloids have spaces that can be interpreted as accessory cavities. For example, in Pteranodon longiceps (KUVP 976, USNM 13868), there is a clear
WZTMER-ANTORBITAL CAVITY OF ARCHOSAURS FIGURE 32. Deinonychus anrirrhopus. Right palatine <strong>of</strong> YPM 5210 in dorsal view, with interpretive drawing.
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