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The Skull of Velociraptor (Theropoda) from the Late Cretaceous of Mongolia

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The Skull of Velociraptor (Theropoda) from the Late Cretaceous of Mongolia The skull of Velociraptor (Theropoda) from the Late Cretaceous of Mongolia RINCHEN BARSBOLD and HALSZKA OSMOLSKA Barsbold, R. & Osmdlska, H. 1999. The skull of Velociraptor (Theropoda) from the Late Cretaceous of Mongolia. -Acta Palaeontologica Polonica 44,2, 189-219. The well preserved material of the Late Cretaceous dromaeosaurid, Velociraptor mon- goliensis, has allowed us to supplement earlier descriptions of the skull in this species. The skull of I? mongoliensis is similar to that of Deinonychus antirrhopus, but differs from the latter by: (1) laterally convex supratemporal arcade resulting in short, rounded supratemporal fenestra; (2) depressed nasal; (3) longer maxillary process of premaxilla; (4) lack of separate prefrontal, and (5)convex ventral border of the dentary. These differ- ences, especially that in the structure of the temporal region, support generic distinction of Deinonychus and Velociraptor.Skulls of other dromaeosaurids are compared. Key words: Dinosauria, Theropoda, Dromaeosauridae, Velociraptor,skull, mandible, Late Cretaceous, Gobi Desert, Mongolia. Rinchen Barsbold [barsgeodin@magicnet,mn],Institute of Geology, Mongolian Acad- emy of Sciences, Enkh Taivani Gudamji, Ulan Bator 210351, Mongolia. Halszka Osmdlska [[email protected]], Znstytut Paleobiologii PAN, ul. Twarda 51/55, PL-00-818 Warszawa, Poland. Introduction Dromaeosaurids were small to medium size theropods, except for Utahraptor and an undetermined dromaeosaurid from Japan (Azuma & Currie 1995), which were rela- tively large animals. They were cursorial, moderately fast carnivores with large, rostrolaterally facing eyes. There is taphonomic evidence (Ostrom 1990; Maxwell & Ostrom 1995) that at least Deinonychus antirrhopus may have hunted in packs. On the other hand, Velociraptor mongoliensis may have also been a carrion feeder (Osm6lska 1993; but see Kielan-Jaworowska & Barsbold 1972; Unwin et al. 1994; Fastovsky et al. 1997 for alternative interpretations). The Dromaeosauridae are Cretaceous mani- raptoran theropods, and eight monotypic genera: Adasaurus Barsbold, 1983, Deino- nychus Ostrom, 1969, Dromaeosaurus Matthew & Brown, 1922, Hulsanpes Osm61- ska, 1982, Omithodesmus Seeley, 1887, Sauromitholestes Sues, 1978, Utahraptor 190 Skull of Velociraptor: BARSBOLD & OSMOLSKA Kirkland, Burge, & Gaston, 1993, and Velociraptor Osborn, 1924 are presently as- signed to this family. Three of these genera (Adasaurus, Hulsanpes, and Ornitho- desmus), are based exclusively on incomplete postcrania. The most peculiar dromaeosaurid feature is the opisthopubic pelvis (Barsbold 1976), which distinguishes these dinosaurs from other theropods, except for the dis- tantly related therizinosauroids; the extremely long caudal zygapophyses and chev- rons are probably also common to all dromaeosaurids, but caudal vertebrae are known only in Deinonychus, Velociraptor, and Saurornitholestes (Dr P.J. Currie's personal communication 1999) The skull of V mongoliensis has been known for more than 70 years. Over this time, it has been described, illustrated or commented by several authors, among them Sues (1977a), Barsbold (1983), Paul (1988), and Ostrom (1969b, 1990). Up to now, Veloci- raptor is represented by the most complete and most numerous skulls and postcrania among dromaeosaurids (in addition to the here described material, there are numerous still not described specimens recently collected by the AMNH Asiatic Expeditions). It was Ostrom (1969a), who first recognised the close relationship of Velociraptor with the North American forms, Dromaeosaurus and Deinonychus, and assigned it to the Dromaeosauridae (= Dromaeosaurinae Matthew & Brown 1922). The skulls are largely complete in the two latter genera, whereas skulls of Saurornitholestes and Utahraptor are represented by a few bones each. The following description of the skull in V mongoliensis supplements the earlier descriptions by Osborn (1924), Sues (1977a), and Barsbold (1983). The skull data for D. antirrhopus, Dromaeosaurus albertensis, and Saurornitholestes langstoni used in the comparisons below are re- spectively from Ostrom (1969b), Colbert & Russell (1969), Currie (1995), Sues (1977a), and Witmer & Maxwell (1996), unless stated otherwise. Institutional abbreviations: AMNH, American Museum of Natural History, New York; GIN, Institute of Geology, Mongolian Academy of Sciences, Ulan Bator; PIN, Museum of Palaeontology, Russian Academy of Sciences, Moscow; ROM, Royal Ontario Museum, Toronto; TPM, Royal Tyrrell Museum of Palaeontology, Drumheller; ZPAL, Institute of Paleobiology, Polish Academy of Sciences, Warsaw. Systematic palaeontology Theropoda Marsh, 1881 Maniraptora Gauthier, 1986 Family Dromaeosauridae Matthew & Brown, 1922 Subfamily Velociraptorinae Barsbold, 1983 Velociraptor Osborn, 1924 Type species by monotypy: Velociraptor mongoliensis Osborn, 1924. Velociraptormongoliensis Osborn, 1924 Figs 1-8. Holotype: AMNH 6515, skull, mandible, manual digit I. ACTA PALAEONTOLOGICA POLONICA (44) (2) 191 Type horizon and locality: Djadokhta Formation (?early Campanian), Bayn Dzak, Ornnogov prov- ince, Gobi Desert, Mongolia. Material. -The present description of the skull of V: mongoliensis is based on several skulls pertaining to more or less complete skeletons from the Upper Cretaceous sand- stone deposits of the Mongolian Gobi. The specimens studied were found by the Polish-Mongolian Palaeontological Expeditions (specimens GIN 100125 and ZPAL MgD-U97, see Kielan-Jaworowska & Barsbold 1972; Gradziliski et al. 1977), by the Soviet-Mongolian Palaeontological Expeditions (specimen PIN 3 14318), by a Mongo- lian expedition (specimen GIN 100124, see Barsbold 1983) and by the Mongolian- -Japanese Palaeontological Expeditions (specimen GIN 10012000). Description of the postcrania preserved with these skulls will be published at a later date (Barsbold & Osm6lska in preparation). The following specimens derive from the Djadokhta Formation (?early Campanian, see Kielan-Jaworowska & Hurum 1997), TugrrE;zn-Shire,Omnogov, Mongolia: GIN 100124 - consists of an almost complete, articulated, but dorsoventrally flat- tened skull, both mandibular rami, and a few fragmentary postcranial bones. The premaxillae are damaged rostrally, as are the caudal ends of the nasals and the rostral tips of the frontals along their mutual contact; the shaft and the caudal end of the left lacrimal are missing; jugals, quadratojugals and quadrates are fragmentary; pterygoids and palatines are partly damaged, those on the left side having shifted caudally from their natural position; vomers are not exposed. The dentaries are positioned between the left and right maxillae and premaxillae; the postdentary portion of the right man- dibular ramus is fragmentary, the left surangular lacks its rostrodorsal part; the dentary teeth and articular region are not exposed. GIN 100125 - includes skeletons of two dinosaurs - V: mongoliensis and Proto- ceratops andrewsi. Remains of these dinosaurs are preserved in a position suggestive of combat and for that reason this specimen is widely known as one of the 'fighting di- nosaurs'. The V mongoliensis skeleton is complete and articulated. It includes the skull, adducted mandibles and postcranium. The snout and mandible are somewhat compressed laterally and the dentaries are forced under the skull obscuring the palate and rostral part of the basicranium. GIN 10012000 - is represented by the complete skeleton of a young individual. It includes skull with mandible and postcranium. PIN 314318 - almost complete skull with left mandibular ramus, lacking the right temporal region; tip of the rostrum is severely damaged. The Barun Goyot Formation (?late Campanian: Kielan-Jaworowska & Hurum 1997), Khulsan, Bayankhongor, Mongolia, yielded one specimen ZPAL MgD-U97 - included is the left, rostral half of the skull (exposed from the medial side) but lacks the premaxilla and tips of the vomers. Both mandibular rami are placed between the maxillae (as in all described V mongoliensis specimens with the mandible preserved); the left ramus lacks the caudal articular region and the tip of the dentary; the fragmen- tary right ramus includes the splenial and an incomplete dentary. Associated with the skull was a distal part of the left hind limb. The new, abundant Velociraptor material recently collected in Mongolia by the Mon- golian Academy of Sciences - American Museum of Natural History Expeditions (Norell & Makovicky 1998) has not been studied by the present authors. 192 Skull of Velociraptor: BARSBOLD & OSMOLSKA Revised diagnosis (based upon skull characters). - Skull shallow with long snout, preorbital length constituting 60% of total skull length (estimated at about 50% in Dr. albertensis and D. antirrhopus; proportions unknown in other drornaeosaurids); supratemporal fossa (and fenestra) subcircular, bound by laterally convex supra- temporal arcade (elongate, with straight arcade in D. antirrhopus; shape of fossa un- known in other drornaeosaurids); frontal long, almost four times longer than wide across the orbital portion, and almost four times as long as parietal [wider in S. langstoni, frontal length at most three times the width across the orbital portion; in Dr. albertensis frontal shorter, about twice as long as wide; parietal/frontal length ratio un- known in both these species; in D. antirrhopus, frontal only three times longer than wide and twice as long as parietal (Dr L.M. Witmer's personal communication 1999)l; rostral border of internal antorbital fenestra broadly rounded (subtriangular in
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