, Volume 95, Issue 3, pp 185–191 | Cite as

Behavioral and faunal implications of Early Cretaceous deinonychosaur trackways from China

  • Rihui Li
  • Martin G. LockleyEmail author
  • Peter J. Makovicky
  • Masaki Matsukawa
  • Mark A. Norell
  • Jerald D. Harris
  • Mingwei Liu
Original Paper


Deinonychosaurian theropods, the dinosaurian sister group of birds, are characterized by a large raptorial claw borne on a highly modified second digit that was thought to be held in a retracted position during locomotion. In this study, we present new trackway evidence for two coeval deinonychosaurian taxa from the Early Cretaceous of Shandong, China that indicate a hitherto unrecognized body size diversity for this period and continent. These fossil tracks confirm diversity and locomotory patterns implied by phylogeny and biogeography, but not yet manifested in the body fossil record. Multiple parallel and closely spaced trackways generated by the larger track maker provide the best evidence yet discovered for gregarious behavior in deinonychosaurian theropods.


Deinonychosaur Cretaceous Footprint Gregariousness China Locomotion 



This study was supported financially in part by National Natural Science Foundation of China grant 40572011 to R-H. L. Additional support was provided by the University and Society Collaboration of the Japanese Ministry of Education, grant-in-aid 1183303, 1999-2000 to M. M., and by the University of Colorado at Denver Dinosaur Tracks Museum to M. G. L. M.A.N. and P.J.M. were both supported by US National Science Foundation AToL EAR 0228607 and by the AMNH and FMNH, respectively. S. Harris produced the movie and M. Ellison photographed the seriema foot for Fig. 4.

Supplementary material (2.8 mb)
S1 Supplementary Movie of 3D views of Dromaeopodus paratype: a movie providing a 3D range of views of a digital rendering of the paratype Dromaeopodus shandongensis track (CU 214.112, replica of left pes) is available with the online version of this article at: (MOV 2.84 MB)
114_2007_310_MOESM2_ESM.doc (4 kb)
S2 Appendix (DOC 4.05kb)
114_2007_310_MOESM3_ESM.doc (30 kb)
S3 Table of comparative limb bone measurements (DOC 30.5kb)
114_2007_310_MOESM4_ESM.doc (26 kb)
S4 Supplementary References (DOC 26.0kb)


  1. Barsbold R (1998) Fighting dinosaurs—they really fought. In: de Carvalho AMG, Cachão MAP, Andrade AMPSAF, da Silva CAPFM, dos Santos VAF (eds) I Encontro Internacional sobre Paleobiologia dos Dinossáurios: Programa de Musealização para Pistas de Dinossáurios em Portugal. Museu Nacional de História Natural, Lisbon, pp 74–78Google Scholar
  2. Brinkman DL, Cifelli RL, Czaplewski NJ (1998) First occurrence of Deinonychus antirrhopus (Dinosauria: Theropoda) from the Antlers Formation (Lower Cretaceous: Aptian–Albian) of Oklahoma. Okla Geol Surv Bull 146:1–27Google Scholar
  3. Carpenter K (1998) Evidence of predatory behavior by carnivorous dinosaurs. Gaia 15:135–144Google Scholar
  4. Carpenter K, DiCroce T, Gilpin D, Kinneer B, Sanders F, Tidwell V, Shaw A (2002) Origins of the Early and “middle” Cretaceous dinosaurs of North America: implications for plate tectonics. Proceedings of the International Symposium on New Concepts in Global Tectonics, May 2002, Otero Junior College, La Junta, CO:289–308Google Scholar
  5. Chinnery BJ, Lipka TR, Kirkland JI, Parrish JM, Brett-Surman MK (1998) Neoceratopsian teeth from the Lower to middle Cretaceous of North America. N M Mus Nat Hist Sci Bull 14:297–302Google Scholar
  6. Cifelli R, Kirkland JI, Weil A, Deino AL, Kowallis BJ (1997) High-precision 40Ar/39Ar geochronology and the advent of North America’s Late Cretaceous terrestrial fauna. Proc Natl Acad Sci 94:11163–11167PubMedCrossRefGoogle Scholar
  7. Currie PJ, Dong Z (2001) New information on Cretaceous troodontids (Dinosauria, Theropoda) from the People’s Republic of China. Can J Earth Sci 38:1753–1766CrossRefGoogle Scholar
  8. Gaston R, Lockley MG, Lucas SG, Hunt AP (2003) Grallator-dominated fossil footprint assemblages and associated enigmatic footprints from the Chinle Group (Upper Triassic), Gateway area, Colorado. Ichnos 10:153–163CrossRefGoogle Scholar
  9. Gatesy SM, Middleton KM, Jenkins FA Jr, Shubin NH (1999) Three-dimensional preservation of foot movements in Triassic theropod dinosaurs. Nature 399:141–144CrossRefGoogle Scholar
  10. Kirkland JI (2005) Utah’s newly recognized dinosaur record from the Early Cretaceous Cedar Mountain Formation. Utah Geol Surv Notes 37:1–5Google Scholar
  11. Kirkland JI, Gaston R, Burge D (1993) A large dromaeosaur (Theropoda) from the Lower Cretaceous of eastern Utah. Hunteria 2:1–16Google Scholar
  12. Kirkland JI, Britt B, Burge DL, Carpenter K, Cifelli R, DeCourten F, Eaton J, Hasiotis S, Lawton T (1997) Lower to middle Cretaceous dinosaur faunas of the central Colorado Plateau: a key to understanding 35 million years of tectonics, sedimentology, evolution and biogeography. Brigham Young Univ Geol Stud 42:69–103Google Scholar
  13. Kirkland JI, Zanno LE, Sampson SD, Clark JM, DeBlieux DD (2005) A primitive therizinosauroid dinosaur from the Early Cretaceous of Utah. Nature 435:84–87PubMedCrossRefGoogle Scholar
  14. Li R, Lockley MG, Liu M (2005a) A new ichnotaxon of fossil bird track from the Early Cretaceous Tianjialou Formation (Barremian–Albian), Shandong Province, China. Chin Sci Bull 50:1149–1154CrossRefGoogle Scholar
  15. Li R-H, Liu M-W, Lockley MG (2005b) Early Cretaceous dinosaur tracks from the Houzuoshan Dinosaur Park in Junan County, Shandong Province, China. Geol Bull Chin 24:277–280 (in Chinese with English abstract)Google Scholar
  16. Li D, Azuma Y, Fugita M, Lee Y-N, Arakawa Y (2006) A preliminary report on two new vertebrate track sites including dinosaurs from the Early Cretaceous Hekou Group, Gansu Province, China. J Paleontol Soc Korea 22:29–49Google Scholar
  17. Lockley MG (1989) Tracks and traces: new perspectives on dinosaurian behavior, ecology, and biogeography. In: Padian K, Chure DJ (eds) The Age of Dinosaurs. Short courses in paleontology 2. The Paleontological Society, Knoxville, pp 134–145Google Scholar
  18. Lockley MG, Matsukawa M (1999) Some observations on trackway evidence for gregarious behavior among small bipedal dinosaurs. Palaeogeogr Palaeoclimatol Palaeoecol 150:25–31CrossRefGoogle Scholar
  19. Lockley MG, White D, Kirkland J, Santucci V (2004) Dinosaur tracks from the Cedar Mountain Formation (Lower Cretaceous), Arches National Park, Utah. Ichnos 11:285–293CrossRefGoogle Scholar
  20. Lockley MG, Kim SH, Kim JY, Matsukawa M, Li R, Li J, Yang S-Y (2007a) Minisauripus—the track of a diminutive dinosaur from the Cretaceous of China and Korea: implications for stratigraphic correlation and theropod foot morphodynamics. Cret Res, in pressGoogle Scholar
  21. Lockley MG, Li R, Harris JD, Matsukawa M, Liu M (2007b) Earliest zygodactyl bird feet: evidence from Early Cretaceous roadrunner-like tracks. Naturwissenschaften 94:657–665PubMedCrossRefGoogle Scholar
  22. Makovicky PJ, Norell MA (2004) Troodontidae. In: Weishampel DB, Dodson P, Osmólska H (eds) The Dinosauria, 2nd Edition. University of California Press, Berkeley, pp 184–195Google Scholar
  23. Makovicky PJ, Apesteguía S, Agnolin F (2005) The earliest dromaeosaurid theropod from South America. Nature 437:1007–1011PubMedCrossRefGoogle Scholar
  24. Maxwell WD, Ostrom JH (1995) Taphonomy and paleobiological implications of Tenontosaurus-Deinonychus associations. J Vertebr Paleontol 15:707–712CrossRefGoogle Scholar
  25. Norell MA, Makovicky PJ (1997) Important features of the dromaeosaur skeleton: information from a new specimen. Am Mus Novit 3215:1–28Google Scholar
  26. Norell MA, Xu X (2005) Feathered dinosaurs. Ann Rev Earth Planet Sci 33:277–299CrossRefGoogle Scholar
  27. Norell MA, Clark JM, Turner AH, Makovicky PJ, Barsbold R, Rowe T (2006) A new dromaeosaurid theropod from Ukhaa Tolgod (Ömnögov, Mongolia). Am Mus Novit 3545:1–51CrossRefGoogle Scholar
  28. Ostrom JH (1969) Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Bull Peabody Mus Nat Hist 30:1–165Google Scholar
  29. Ostrom JH (1972) Were some dinosaurs gregarious? Palaeogeogr Palaeoclimatol Palaeoecol 11:287–301CrossRefGoogle Scholar
  30. Ostrom JH (1990) Dromaeosauridae. In: Weishampel DB, Dodson P, Osmólska H (eds) The Dinosauria. University of California Press, Berkeley, pp 269–279Google Scholar
  31. Perle A, Norell M, Clark J (1999) A new maniraptoran theropod—Achillobator giganticus (Dromaeosauridae)—from the Upper Cretaceous of Burckhant, Mongolia. Contrib Mong-Am Mus Paleontol Proj 101:1–105Google Scholar
  32. Plafker G, Berg HC (1994) Overview of the geology and tectonic evolution of Alaska. In: Plafker G, Berg HC (eds) The Geology of Alaska. The Geology of North America G-1. Geological Society of America, Boulder, pp 989–1021Google Scholar
  33. Roach BT, Brinkman DB (2007) A reevaluation of cooperative pack hunting and gregariousness in Deinonychus antirrhopus and other nonavian theropod dinosaurs. Bull Peabody Mus Nat Hist 48:103–138CrossRefGoogle Scholar
  34. Russell DA, Dong Z (1993) The affinities of a new theropod from the Alxa Desert, Inner Mongolia, People’s Republic of China. Can J Earth Sci 30:2107–2127CrossRefGoogle Scholar
  35. Senter P (2007) A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). J Syst Palaeontol doi: 10.1017/S1477201907002143
  36. Senter P, Barsbold R, Britt BB, Burnham DA (2004) Systematics and evolution of Dromaeosauridae (Dinosauria, Theropoda). Bull Gunma Mus Nat Hist 8:1–20Google Scholar
  37. Turner AH, Hwang SH, Norell MA (2007a) A small derived theropod from Öösh, Early Cretaceous, Baykhangor Mongolia. Am Mus Novit 3557:1–27CrossRefGoogle Scholar
  38. Turner AH, Pol D, Clarke JA, Erickson GM, Norell MA (2007b) A basal dromaeosaurid and size evolution preceding avian flight. Science 317:1378–1381CrossRefGoogle Scholar
  39. Xu X, Norell MA (2004) A new troodontid dinosaur from China with avian-like sleeping posture. Nature 431:838–841PubMedCrossRefGoogle Scholar
  40. Xu X, Norell MA, Wang X-L, Makovicky PJ, Wu X-C (2002) A basal troodontid from the Early Cretaceous of China. Nature 415:780–784PubMedGoogle Scholar
  41. Zhen S, Li J, Zhang B (1994) Dinosaur and bird footprints from the Lower Cretaceous of Emei County, Sichuan, China. Mem Beijing Nat Hist Mus 54:105–120Google Scholar

Copyright information

© Springer-Verlag 2007

Authors and Affiliations

  • Rihui Li
    • 1
  • Martin G. Lockley
    • 2
    Email author
  • Peter J. Makovicky
    • 3
  • Masaki Matsukawa
    • 4
  • Mark A. Norell
    • 5
  • Jerald D. Harris
    • 6
  • Mingwei Liu
    • 7
  1. 1.Qingdao Institute of Marine GeologyChina Geological SurveyQingdaoPeople’s Republic of China
  2. 2.Dinosaur Tracks MuseumUniversity of Colorado at DenverDenverUSA
  3. 3.Department of GeologyField Museum of Natural HistoryChicagoUSA
  4. 4.Department of Environmental SciencesTokyo Gakugei UniversityKoganeiJapan
  5. 5.Department of Vertebrate PaleontologyAmerican Museum of Natural HistoryNew YorkUSA
  6. 6.Science DepartmentDixie State CollegeSt. GeorgeUSA
  7. 7.Fourth Geological and Mineral Resources Survey of ShandongWei-fangPeople’s Republic of China

Personalised recommendations