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Skull Structure and Evolution in Tyrannosaurid Dinosaurs

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Skull Structure and Evolution in Tyrannosaurid Dinosaurs Skull structure and evolution in tyrannosaurid dinosaurs PHILIP J. CURRIE, JØRN H. HURUM, and KAROL SABATH Currie, P.J., Hurum, J.H., and Sabath, K. 2003. Skull structure and evolution in tyrannosaurid dinosaurs. Acta Palaeonto− logica Polonica 48 (2): 227–234. Tyrannosauridae can be subdivided into two distinct subfamilies—the Albertosaurinae and the Tyrannosaurinae. Previ− ously recognized subdivisions Aublysodontinae and Shanshanosaurinae are rejected because they are based on insuffi− cient material and juvenile specimens. Our results are based upon a phylogenetic analysis using PAUP program (Swofford 1999) of 77 skull characters and seven genera (Albertosaurus, Alioramus, Daspletosaurus, Gorgosaurus, Nanotyrannus, Tarbosaurus, and Tyrannosaurus); with Allosaurus as outgroup. Of the 77 characters used, more than half were parsimony informative. Asingle most parsimonious tree was obtained with the Tree Length being 88. The analysis of cranial characters and comparison of postcranial features reveal that Tarbosaurus bataar is not the sister taxon of Tyrannosaurus rex (contra Holtz 2001). Their similarities are partially due to the fact that both are extremely large ani− mals. Thus, Tarbosaurus should be considered a genus distinct from Tyrannosaurus. Key words: Dinosauria, Theropoda, Coelurosauria, Tyrannosauridae, phylogeny, parsimonious analysis. Philip J. Currie [[email protected]], Royal Tyrrell Museum of Palaeontology, Box 7500, Drumheller, Alberta TOJ OYO, Canada; Jørn H. Hurum [[email protected]], Geologisk museum, Universitetet i Oslo, Postboks 1172 Blindern, N−0318 Oslo, Norway; Karol Sabath [[email protected]], Instytut Paleobiologii PAN, ul. Twarda 51/55, PL−00−818 Warszawa, Poland (currently: [[email protected]], Polish Geological Institute, ul. Rakowiecka 4, PL−00−975 Warszawa, Poland). Introduction Allosaurus Marsh, 1877 was chosen as a theropod generalized enough to serve as the outgroup. Siamotyrannus Buffetaut, To the general public, tyrannosaurids are amongst the best Suteethorn, and Tong, 1996 and Eotyrannus Hutt, Naish, known and most popular dinosaurs. Yet there have been rela− Martill, Barker, and Newbery, 2001 (Naish et al. 2001) have tively few monographic descriptions of specimens, and there both been described as primitive relatives of the Tyranno− is considerable disagreement about valid generic and specific sauridae. Although the fused, dorsally convex nasals alone names, and relationships. For example, Paul (1988) recog− demonstrate the probable relationship of the latter to tyranno− nized 14 tyrannosaurid species in five genera (Albertosaurus, saurids, we have not examined the original specimens and did Alioramus, Aublysodon, Indosuchus,andTyrannosaurus); not feel enough was known to meaningfully code the genera. Molnar et al. (1990) listed 12 species in at least eight genera For similar reasons, the Asian tyrannosaurid Alectrosaurus (Albertosaurus, Alectrosaurus, Alioramus, Chingkankou− Gilmore, 1933 was also excluded from this analysis. The saurus, Daspletosaurus, Nanotyrannus, Tarbosaurus,and holotype material of Alectrosaurus does not include any cra− Tyrannosaurus); Olshevsky (1995) referred to 19 species nial material (Gilmore 1933; Mader and Bradley 1989), and level taxa in at least 14 genera (Albertosaurus, Alectrosaurus, the skull of the better preserved Mongolian specimen (Perle Alioramus, Aublysodon, Daspletosaurus, Dinotyrannus, 1977) has been unavailable for years. Gorgosaurus, Jenghizkhan, Maleevosaurus, Nanotyrannus, Holtz (2001) and earlier authors recognised two sub− Shanshanosaurus, Stygivenator, Tarbosaurus,andTyranno− families (Aublysodontinae and Tyrannosaurinae) within the saurus); Holtz (2001) incorporated eleven species level taxa in Tyrannosauridae. Aublysodontinae Nopcsa, 1928 has al− at least eight genera (Albertosaurus, Alectrosaurus, Alio− ways been a vague subfamily established on the basis of a ramus, Aublysodon, Daspletosaurus, Gorgosaurus, Shan− tooth genus (Aublysodon Leidy, 1868). Although several shanosaurus,andTyrannosaurus). skeletons have been assigned to this genus, the recent discov− The purpose of this paper is to examine the intra−relation− eries of juveniles of well−established tyrannosaurid genera ships of the Tyrannosauridae Osborn, 1906, a family that has (including Albertosaurus Osborn, 1905, Daspletosaurus been well established as coelurosaurian (Holtz 1994, 2000, Russell, 1970, Gorgosaurus Lambe, 1914, Tarbosaurus 2001; Sereno 1997, 1999; Makovicky and Sues 1998). As Maleev, 1955, and Tyrannosaurus Osborn, 1905) have made pointed out by Holtz (2001), the relationship of tyrannosaurids it increasingly obvious that Aublysodon is based strictly on to other Coelurosauria von Huene, 1914 is unclear. Because characters that are morphologically and allometrically im− the intention of this paper was not to resolve this problem, mature. Aublysodon molnari Paul, 1988 (= Stygivenator mol− Acta Palaeontol. Pol. 48 (2): 227–234, 2003 http://app.pan.pl/acta48/app48−227.pdf 228 ACTA PALAEONTOLOGICA POLONICA 48 (2), 2003 nari Olshevsky, 1995) and Albertosaurus megagracilis Paul, for character 32. However, it is also evident that in Daspleto− 1988 (=Dinotyrannus megagracilis Olshevsky, 1995) are saurus the maxillary fenestra is small in juveniles (TMP probably juveniles of Tyrannosaurus rex Osborn, 1905 (see 94.143.1) but large in adults (NMC 8506, TMP 85.64.1). Be− Currie 2003). The Kirtland Shale aublysodontine (Lehman cause we do not know what the maxillary fenestra of an adult and Carpenter 1990) may be a juvenile Daspletosaurus Nanotyrannus looks like, character 32 is coded with a ×. (Holtz 2001). Nanotyrannus Bakker et al., 1988 was in− Some characters (such as character 40) were coded as seen in cluded in the Aublysodontinae by Sereno (1998), but most the only known specimens, are different in coding from other authors recognize it as a tyrannosaurid that may even Tarbosaurus and/or Tyrannosaurus, but may ultimately be congeneric with Tyrannosaurus (Carr 1999; Holtz 2001). prove to also be age−related and therefore invalid. Alectrosaurus has been placed in this subfamily because it has premaxillary teeth that lack denticles (Currie 2000), but Characters if Aublysodon is congeneric with tyrannosaurine genera, then the lack of denticles cannot be used to define the subfamily The characters used in this analysis were mostly derived Aublysodontinae. Shanshanosaurus Dong, 1977 has been from the research done by Hurum and Sabath (2003), and variously assigned to its own family (Shanshanosauridae Currie (2003). Work by earlier researchers (especially Rus− Dong, 1977), its own subfamily (Shanshanosaurinae), or to sell 1970; Carr 1999) provided a suitable framework for the Aublysodontinae (Paul 1988), but Currie and Dong character selection, but the most relevant research is that of (2001) suggested it is a tyrannosaurid that is possibly conge− Holtz (1994, 2000, and especially 2001). neric with Tarbosaurus. In short, there is no reason at present Holtz (2001) has done the most formal, phylogenetic anal− to assume that the Aublysodontinae exists. ysis of the intrafamily relationships of the Tyrannosauridae, This paper is based mostly on recent studies of the skull although he considered it as preliminary. In this paper, many anatomy of North American tyrannosaurids (Currie 2003), of his characters (Holtz 2001: appendix 7.1: 2, 3, 4, 15, 42, 48, and of Asian tyrannosaurids (Hurum and Sabath 2003); more 69, 70, 72, 74, 96, 103, 106, 108, 110) have been adopted. detailed anatomical data are given in these two papers. Others (9, 12, 56, 65, 66, 104+105, 109) have been modified Institutional abbreviations: AMNH, American Museum to accommodate the different style of presentation, and the of Natural History; NMC, National Museum of Canada, Ot− different suites of specimens and taxa. Some of the characters tawa; PIN, Palaeontological Institute, Russian Academy of used by Holtz (6, 7, 13, 42, 47, 49–55, 57, 59, 62, 63, 68, 93, Sciences, Moscow; TMP, Royal Tyrrell Museum of Palae− 99) were incorporated into this paper, but were completely re− ontology, Drumheller, Canada. worked or were included within character suites that bear little resemblance to his original wording. Anumber of characters used by Holtz (2001) were not Phylogenetic analysis used in our data matrix. As this paper examines only cranial anatomy of tyrannosaurids, none of his postcranial charac− ters were included. His characters numbered 1, 40, 41, 43, Selection of taxa and the problem of juvenile 44, 46, 58, 61, 64, 76, and 77 are considered to be allometric specimens (either within a growth series or between taxa in which there are absolute size differences at maturity). The relative size of Atotal of seven tyrannosaurid genera are included in this the prefrontals (character 5 of Holtz 2001) could not be quan− analysis. Albertosaurus, Alioramus, Daspletosaurus, Gorgo− tified in a meaningful way, and was therefore dropped from saurus, Nanotyrannus, Tarbosaurus, and Tyrannosaurus; the analysis. Prominent muscular fossae (character 10) were with Allosaurus as outgroup. More than 200 original cata− not recognized on the dorsal surfaces of palatines, and there− logued specimens of Albertosaurus, Daspletosaurus, fore the character was omitted pending further research. The Gorgosaurus, Tarbosaurus, and Tyrannosaurus have been angular is supposed to terminate posteriorly in front of the studied. Whenever possible, characters were coded from posterior surangular fenestra in Tarbosaurus bataar (Holtz
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