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GAIA Nº 15, LISBOA/, DEZEMBRO/DECEMBER 1998, pp. 1-3 (ISSN: 0871-5424)

THEROPOD , MORE THAN JUST ORIGINS

Thomas R. HOLTZ, Jr. Department of , University of Maryland. College Park, MARYLAND 20735. USA E-mail: [email protected]

BUCKLAND’s 1824 description of of theropod ichnology (e.g., FARLOW, 1981; THUL- initiated the study of the paleobiology of theropod di- BORN, 1990; LOCKLEY, 1991) help to reconstruct the nosaurs. Given the limited material upon which he locomotory ability and diversity within the carnivo- based his , Buckland misconceived of the rous , while functional morphological stud- “Great of ” as an enormous ies have examined theropod mastication varanid. The subsequent description of (ERICKSON et al. 1996), grasping ability (GALTON, by COPE (1866) revealed the bipedal 1971), and cursoriality (HOLTZ, 1995). posture and reduced of theropod dino- In this volume we examine new advances in the saurs, and the discovery of relatively complete study of the , , , ich- of longipes W AGNER, nology, functional , and distribution of 1859 (not initially recognized as a ) and of theropod dinosaurs, with emphasis on the non- nasicornis M , 1884 revealed ARSH avian members of this . We have organized the much of the basic anatomy of the carnivorous dino- twenty-nine papers in this volume around these ma- saurs. Theropod ichnology can be traced to the clas- jor themes in research on . sic work of HITCHCOCK (1858). The understanding of the phylogenetic relation- The early 1900s brought to light the highly spe- ships and global distribution of theropod dinosaurs cialized anatomy of some of the thero- has increased greatly in recent . A new phy- pods, such as the gigantic tyrannosaurids (O SBORN, logeny of theropods, incorporating these recent dis- 1905, 1906, 1916), ostrich-like ornithomimosaurs coveries, is presented by Holtz provides a new (O 1916), sail-backed conical toothed spino- SBORN, comprehensive analysis of the phylogeny of thero- saurids (S , 1915), and the various small- TROMER pod dinosaurs. Heckert & Lucas and Rauhut & Hun- bodied derived coelurosaurs of Mongolia (O , SBORN gerbühler examine aspects of theropod 1924). The discovery of by O in STROM diversity. Coria & Salgado describe for the first time the mid-1960s (1969a, b) revolutionized various a new theropod taxon from the Cretaceous of Ar- concepts of dinosaur , reviving late 19th Cen- gentina, which appears to represent a mem- tury hypotheses of the possibility of dinosaurian en- ber of the abelisaur radiation. Jones & Chure report dothermy and of the dinosaurian origin of . The on the use of systematic radiological surveying tech- latter concept was further expanded by phyloge- niques (a promising technology for future paleonto- netic analyses (e.g., GAUTHIER, 1986) that demon- logical exploration), which resulted in the recovery strated that Aves lies within the coelurosaurian of the of a new allosaurid from the Up- theropods, and thus that birds are the surviving part per Jurassic . Additional new of the theropod dinosaur radiation; bolstering this discoveries are also reported here, including new hypothesis is the discovery of in non-avian forms from the mid-Cretaceous of (Rich et theropod taxa (C ,DONG &ZHEN, 1998; JI et al. HEN al.; Calvo & Coria), and an unusual association of 1998). Discoveries of extremely primitive theropods troodontid teeth with baby hadrosaurids (Ryan et (S &NOVAS, 1992; SERENO et al. 1993) from ERENO al.). Argentina reveal the ancestral conditions of the car- nivorous dinosaurs. Bizarre new theropod taxa, The majority of the contributions concern the such as the therizinosauroids of (PERLE, 1979) functional anatomy and possible behaviors of vari- and the abelisaurids of (BONAPARTE & ous theropod dinosaurs. Carpenter describes two NOVAS, 1985), continued to be discovered around key specimens indicating active attempts the world in the latter part of the 20th Century: with the by theropod dinosaurs: the famous “fighting dino- discovery of HAMMER &HICKER- saur” pair of the dromaeosaurid and the SON, 1994, non-avian theropods have now been ceratopsian , and evidence of a failed found on every continent. New analyses in the field attack by a large theropod (almost certainly Tyran-

1 artigos/papers T.R. HOLTZ, JR.

nosaurus) on a hadrosaurid. The trophic apparatus ACKNOWLEDGEMENTS (jaws and teeth) of theropods has received consid- The guest editors of this special volume of Gaia erable attention: studies included herein examine would like to acknowledge our deep debt of thanks to the function of the jaws of allosaurids (Bakker), the Martin Lockley and Vanda Faria dos Santos for their of (Holtz, Brinkman & Chandler), contribution in reviewing the ichnological submis- the jaws (and hindlimbs) of (Mazzetta, sions to the journal. We would also like to acknowl- Fariña & Vizcaíno), and the skull of edge the patience of the contributing authors: due to (Molnar). Tanke & Currie review the abundant but a series of unforeseen delays this volume was re- larger unreported evidence of head-biting behavior leased in a less timely manner than anticipated. We in a number of large theropod dinosaurs. Hender- hope that the papers herein nevertheless serve to son compares the of various forms of the Mor- stimulate further examinations of the paleobiology rison Formation in terms of trophic and display of the carnivorous dinosaurs. apparatus, while Chure, Fiorillo & Jacobsen report on new examples of the prey utilization by REFERENCES theropods in the same stratigraphic unit. Moving posteriorly from the jaws, Chure discusses the diver- BONAPARTE, J.F. & NOVAS, F.E. (1985) - Abelisaurus comahuen- sis, n.g. et n.sp., del Cretácio Tardío de Patago- sity of form of the orbit between different theropod nia. Ameghiniana, 21: 259-265. taxa. The hindlimb anatomy of theropods, and the BUCKLAND, W. (1824) - Notice on Megalosaurus, or great fossil li- implications of these in terms of the possible cursor- zard of Stonesfield. Trans. Geol. Soc. , 21: 390-397. ial ability in some of these taxa, is the subject of pa- CHEN P.-J.; DONG Z.-M. & ZHEN S.-N. (1998) - An exceptionally pers by Christiansen and Paul. Concluding the well-preserved theropod dinosaur from the section on functional anatomy and behavior, Currie of . , 391: 147-152. presents a significant report of multiple individuals of COPE, E.D. (1866) - Discovery of a gigantic dinosaur in the Creta- ceous of New Jersey. Proc. Acad. Nat. Sci. Philadelphia, 18: tyrannosaurids form the same site in the Upper Cre- 275-279. taceous Horseshoe Canyon Formation of Alberta, ERICKSON, G.M.; VAN KIRK, S.D.; SU J.-T.; LEVENSTON, M.E.; CA- and its implication for possible gregarious behavior LER, W.E. & CARTER, D.R. (1996) - Bite-force estimation for between adult and juvenile individuals within this Tyrannosaurus rex from -marked . Nature, 382: group. 706-708. GALTON, P.M. (1971) - Manus movements of the coelurosaurian The section on theropod ichnology begins with a dinosaur Syntarsus and the opposability of the theropod personal philosophical examination of track mor- hallux. Arnoldia, 5(15): 1-8. phology studies by Lockley. Reports of tracks of car- GAUTHIER, J.A. (1986) - Saurischian monophyly and the , in PADIAN, K. (Ed.), and the Evoluti- nivorous dinosaurs from the Upper Triassic of on of . Mem. Calif. Acad. Sci., 8: 1-40. Queensland, (Thulborn), the Middle and HAMMER, W.R. & HICKERSON, W.J. (1994) - A crested theropod di- Upper Jurassic of Portugal, Germany, and Uzbeki- nosaur from Antarctica. , 264: 828-830. stan (Lockley, Meyer & Santos), the Upper Jurassic HITCHCOCK, E. (1858) - Ichnology of New England: A Report on and Lower Cretaceous of , Asia, and the Sandstone of the Connecticut Valley, Especially Its Fossil (Lockley, Meyer & Moratalla), the Lower and Footmarks. William Wight Press, Boston. lowermost Upper Cretaceous of and Croatia HOLTZ, T.R., Jr. (1995) - The arctometatarsalian , an unusual (Dalla Vecchia), and the Upper Cretaceous of Brazil structure of the metatarsus of Cretaceous Theropoda (Dino- : ). J. Paleontol., 14: 480-519. (Carvalho & Pedrão). Additionally, Loyal et al. pres- JI Q.; CURRIE, P.J., NORELL, M.A. & JI S.-A. (1998) - Two feathe- ent new information of the probably referable red dinosaurs from northeastern China. Nature, 393: 753- to carnivorous dinosaurs from India. 761. Finally, two papers speculate on struc- MARSH, O.C. (1884) - Principle characters of American Jurassic dinosaurs. Part VIII: the Theropoda. Am. J. Sci., 27: tures of the non-avian theropods: the reproductive 329-341. system (LARSON) and the earliest stages of OSBORN, H.F. (1905) - Tyrannosaurus and other Cretaceous car- (GRIFFITHS). nivorous dinosaurs. Bull. Am. Nat. Hist., 21: 259- 265. Even as this volume went to press, significant OSBORN, H.F. (1906) - Tyrannosaurus, Upper Cretaceous carni- new theropod specimens have been brought to light, vorous dinosaur (second communication). Bull. Am. Museum among the most dramatic of which are the first docu- Nat. Hist., 22: 281-296. mented feathered specimens of therizinosauroids OSBORN, H.F. (1916) - Skeletal of , (XU,TANG &WANG, 1999) and dromaeosaurids (XU, , Tyrannosaurus. Bull. Am. Museum Nat. Hist., 35: 733-771. WANG &WU, 1999). The Theropoda remain a great source of study for current and future paleontolo- OSBORN, H.F. (1924) - Three new Theropoda, Protoceratops Zone, central Mongolia. Am. Museum Novit., 144: 1-12. gists: we hope this volume serves as a useful source OSTROM, J.H. (1969) - A new theropod dinosaur from the Lower in these explorations. Cretaceous of . Postilla, 128: 1-17.

2 THEROPOD PALEOBIOLOGY, MORE THAN JUST BIRD ORIGINS

OSTROM, J.H. (1969b) - Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Bull. Peabody Museum Nat. Hist., 30: 1-165. PERLE, A. (1979) - Segnosauridae – a new family of theropods from the of Mongolia. Sovm. Sovetsko-Mon- gol’skaia Paleontol. Eksped. Trudy, 8: 45-55. [In Russian] SERENO, P.C. & NOVAS, F.E. (1992) - The complete skull and ske- leton of an early dinosaur. Science, 258: 1137-1140. SERENO, P.C., FORSTER, C.A., ROGERS, R.R. & MONETTA, A.M. (1993) - Primitive dinosaur from Argentina and the early evolution of Dinosauria. Nature, 361: 64-66. STROMER, E. (1915) - Wirbeltier-Reste der Baharije-Stufe (unters- tes Cenoman). 3. Das Original des Theropoden aegyptiacus nov. gen. nov. spec. Abh. K. Bayer. Akad. Wis- sensch., Math.-Phys. Kl., 28: 1-32. WAGNER, A. (1861) - Neue Beiträge zur Kenntnis der urweltlichen Fauna des lithographischen Schiefers. V. Compsognathus longipes Wagn. Abh. Bayer. Akad. Wiss., 9: 30-38. XU X.; TANG Z.-L. & WANG X.-L. (1999) - A therizinosauroid dino- saur with integumentary structures from China. Nature, 399: 350-354. XU X.; WANG X.-L. & WU X.-C. (1999) - A dromaeosaurid dinosaur with a filamentous integument from the Yixian Formation of China. Nature, 401: 262-266.

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