Hostname: page-component-8448b6f56d-cfpbc Total loading time: 0 Render date: 2024-04-23T23:36:52.665Z Has data issue: false hasContentIssue false

Crouching theropod at the seaside. Matching footprints with metatarsal impressions and theropod authopods: a morphometric approach

Published online by Cambridge University Press:  11 July 2016

M. ROMANO
Affiliation:
Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Invalidenstr 43, Berlin, Germany Dipartimento di Scienze della Terra, ‘Sapienza’ Università di Roma, Piazzale Aldo Moro, 5, 00185, Rome, Italy
P. CITTON*
Affiliation:
Dipartimento di Scienze della Terra, ‘Sapienza’ Università di Roma, Piazzale Aldo Moro, 5, 00185, Rome, Italy
*
Author for correspondence: paolo.citton@uniroma1.it

Abstract

We compare theropod footprints with elongate metatarsal prints from central Italy with known autopod structures in major theropod groups, in order to more accurately define the trackmaker attribution. Our work, using morphometric analysis, shows the considerable potential of explorative methods such as PCA (principal component analysis) and cluster analysis when describing important characters for a given taxonomic group (body and ichnofossils) and identifying important anatomical regions. Moreover, the results of the analysis suggest that the putative trackmaker is likely a member of Ornithomimosauria, with significant affinities in the posterior autopod structure with the genus Struthiomimus. The fundamental importance of integrating both osteological and ichnological data, when investigating locomotor and behavioural hypotheses, is highlighted. This approach could also contribute positively to the complex cognitive process of trackmaker identification and be favourable for the attainment of a more natural definition of ichnotaxa.

Type
Original Articles
Copyright
Copyright © Cambridge University Press 2016 

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Aubouin, J. 1959. Contribution à l’étude géologique de la Grèce Septentrional: les confines de l'Epire et de la Thessalie. Annales Géologiques des Pays Helléniques 10, 1525.Google Scholar
Accordi, G., Carbone, F., Civitelli, G., Corda, L., De Rita, D., Esu, D., Funiciello, R., Kotsakis, T., Mariotti, G. & Sposato, A. 1988. Note illustrative della carta delle litofacies del Lazio-Abruzzo ed aree limitrofe. C.N.R.-P.F. Geodinamica: sottoprogetto 4. Quaderni della Ricerca Scientifica 114, 1223.Google Scholar
Arnaud Vanneau, A. & Premoli Silva, I. 1995. Biostratigraphy and systematic description of benthic foraminifers from mid-Cretaceous shallow-water carbonate platform sediments at sites 878 and 879 (Mit and Takuyo-Daisan Guyots). In Proceedings of the Ocean Drilling Program, Scientific Results, vol. 144 (eds Haggerty, J. A., Premoli Silva, I., Rack, F. & McNutt, M. K.), pp. 199219. College Station, Texas.Google Scholar
Avanzini, M. 1998. Anatomy of a footprint: bioturbation as a key to understanding dinosaur walk dynamics. Ichnos 6, 129–39.Google Scholar
Avanzini, M., Gierliński, G. & Leonardi, G. 2001. First report of sitting Anomepus tracks in European Lower Jurassic (Lavini di Marco site – Northern Italy). Rivista Italiana di Paleontologia e Stratigrafia 107, 131–6.Google Scholar
Baird, D. 1980. A prosauropod dinosaur trackway from the Navajo Sandstone (Lower Jurassic) of Arizona. In Aspect of Vertebrate History: Essay in Honor of Harris Colbert (ed. Jacobs, L. L.), pp. 219230. Flagstaff, Arizona: Museum of Northern Arizona Press.Google Scholar
Burnham, D. A., Derstler, K. L., Currie, P. J., Bakker, R. T., Zhou, Z. & Ostrom, J. H. 2000. Remarkable new bird-like dinosaur (Theropoda: Maniraptora) from the Upper Cretaceous of Montana. University of Kansas Paleontological Contributions 13, 114.Google Scholar
Canudo, J. I., Barco, J. L., Pereda-Suberbiola, X., Ruiz-Omeñaca, J. I., Salgado, L., Fernández- Baldor, F. T. & Gasulla, J. M. 2009. What Iberian dinosaurs reveal about the bridge said to exist between Gondwana and Laurasia in the Early Cretaceous. Bulletin de la Société géologique de France 180, 511.Google Scholar
Carr, T. D. & Williamson, T. E. 2000. A review of Tyrannosauridae (Dinosauria, Coelurosauria) from New Mexico. New Mexico Museum of Natural History and Science Bulletin 17, 113–45.Google Scholar
Castanera, D., Colmenar, J., Sauqué, V. & Canudo, J. I. 2015. Geometric morphometric analysis applied to theropod tracks from the Lower Cretaceous (Berriasian) of Spain. Palaeontology 58, 183200.Google Scholar
Centamore, E., Di Manna, P. & Rossi, D. 2007. Kinematic evolution of the Volsci Range: a new overview. Italian Journal of Geosciences 126, 159–72.Google Scholar
Carrano, M. T. 2007. The appendicular skeleton of Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology 27, 163–79.Google Scholar
Channel, J. E. T. & Kozur, H. W. 1997. How many oceans? Meliata, Vardar and Pindos oceans in Mesozoic Alpine paleogeography. Geology 25, 183–86.2.3.CO;2>CrossRefGoogle Scholar
Cheng, Y. N., Holmes, R., WU, X. C. & Alfonso, N. 2009. Sexual dimorphism and life history of Keichosaurus hui (Reptilia: Sauropterygia). Journal of Vertebrate Paleontology 29, 401–8.Google Scholar
Chinnery, B. 2004. Morphometric analysis of evolutionary trends in the ceratopsian postcranial skeleton. Journal of Vertebrate Paleontology 24, 591609.CrossRefGoogle Scholar
Chiocchini, M., Farinacci, A., Mancinelli, A., Molinari, V. & Potetti, M. 1994. Biostratigrafia a foraminiferi, dasicladali e calpionelle delle successioni carbonati che mesozoiche dell'Appennino centrale. Studi Geologici Camerti, Volume Speciale 1994A, 9130.Google Scholar
Chiocchini, M., Pampaloni, M. L. & Pichezzi, R. M. 2012. Microfacies e Microfossili delle Successioni Carbonatiche Mesozoiche del Lazio e dell'Abruzzo (Italia Centrale) – Cretacico. Memorie per servire alla descrizione della Carta Geologica d'Italia. Volume 17. Roma: ISPRA, Servizio Geologico d'Italia, Dipartimento Difesa del Suolo, 269 pp.Google Scholar
Choiniere, J. N., Forster, C. A. & de Klerk, W. J. 2012. New information on Nqwebasaurus thwazi, a coelurosaurian theropod from the Early Cretaceous Kirkwood Formation in South Africa. Journal of African Earth Sciences 71, 117.Google Scholar
Citton, P., Nicosia, U., Nicolosi, I., Carluccio, R. & Romano, M. 2015. Elongated theropod tracks from the Cretaceous Apenninic Carbonate Platform of southern Latium (central Italy). Palaeontologia Electronica 18.3.49a, 1–12.Google Scholar
Citton, P., Nicosia, U. & Sacchi, E. 2015. Updating and reinterpreting the dinosaur track record of Italy. Palaeogeography, Palaeoclimatology, Palaeoecology 439, 117125.CrossRefGoogle Scholar
Clark, J. M., Norell, M. A. & Barsbold, R. 2001. Two new oviraptorids (Theropoda: Oviraptorosauria), Upper Cretaceous Djadokhta Formation, Ukhaa Tolgod, Mongolia. Journal of Vertebrate Paleontology 21, 209–13.Google Scholar
Conti, M. A., Morsilli, M., Nicosia, U., Sacchi, E., Savino, V., Wagensommer, A., Di maggio, L. & Gianolla, P. 2005. Jurassic dinosaur footprints from Southern Italy: footprints as indicators of constraints in paleogeographic interpretation. Palaios 20, 534–50.Google Scholar
Coria, R. A., Chiappe, L. M. & Dingus, L. 2002. A new close relative of Carnotaurus sastrei Bonaparte 1985 (Theropoda: Abelisauridae) from the Late Cretaceous of Patagonia. Journal of Vertebrate Paleontology 22, 460–5.Google Scholar
Cosentino, D., Cipollari, P., Di Donato, V., Sgrosso, I. & Sgrosso, M. 2002. The Volsci Range in the kinematic evolution of the northern and southern Apennine orogenic system. Bollettino della Società Geologica Italiana, Special Issue 1, 209–18.Google Scholar
Cuesta, E., Díaz-Martínez, I., Ortega, F. & Sanz, J. L. 2015. Did all theropods have chickenlike feet? First evidence of a non-avian dinosaur podotheca. Cretaceous Research 56, 53–9.Google Scholar
Dalla vecchia, M. & Tarlao, A. 2000. New dinosaur track sites in the Albian (Early Cretaceous) of the Istrian peninsula (Croatia). Part II – Paleontology. Memorie di Scienze 52, 227–93.Google Scholar
ELŻAnowski, A. 2001. A new genus and species for the largest specimen of Archaeopteryx . Acta Palaeontologica Polonica 46, 519–32.Google Scholar
Falkingham, P. L. & Gatesy, S. M. 2014. The birth of a dinosaur footprint: subsurface 3D motion reconstruction and discrete element simulation reveal track ontogeny. Proceedings of the National Academy of Sciences (PNAS) 111, 18279–84.Google Scholar
Farlow, J. O., Bates, K. T., Bonem, R. M., Dattilo, B. F., Falkingham, P. L., Gildner, R., Jacen, J., Kuban, G. J., Martin, A. J., O'Brien, M. & Whitcraft, J. 2015. Dinosaur footprints from the Glen Rose Formation (Paluxy River, Dinosaur Valley State Park, Somervell County, Texas). In Early- and Mid-Cretaceous Archosaur Localities of North-Central Texas, Field Trip Guidebook (ed. Noto, C.), pp. 1437. Dallas, Texas: Society of Vertebrate Paleontology.Google Scholar
Farlow, J. O., Holtz, T. R., Worthy, T. H. Jr. & Chapman, R. E. 2013. Feet of the fierce (and not so fierce): pedal proportions in large theropods, other non-avian dinosaurs, and large ground birds. In Tyrannosaurid Biology (eds Parrish, J. M., Molnar, R. E., Currie, P. J. & Koppelhaus, E. B.), pp. 88132. Bloomington, Indiana: Indiana University Press.Google Scholar
Farlow, J. O. & Lockley, M. G. 1993. An osteometric approach to the identification of the makers of early Mesozoic tridactyl dinosaur footprints. New Mexico Museum of Natural History and Science Bulletin 3, 123–31.Google Scholar
Farlow, J. O., Schachner, E. R., Sarrazin, J. C., Klein, H. & Currie, P. J. 2014. Pedal proportions of Poposaurus gracilis: convergence and divergence in the feet of archosaurs. The Anatomical Record 297, 1022–46.Google Scholar
Gatesy, S. M., Middleton, K. M., Jenkins, F. A. Jr. & Shubin, N. H. 1999. Three-dimensional preservation of foot movements in Triassic theropod dinosaurs. Nature 399, 141–4.Google Scholar
Gierliński, G. 1994. Early Jurassic theropod tracks with the metatarsal impressions. Przeglad Geologiczny 42, 280–4.Google Scholar
Gierliński, G. 1996. Feather-like impressions in a theropod resting trace from the Lower Jurassic of Massachusetts. In The Continental Jurassic (ed. Morales, M.), pp. 179–84. Museum of Northern Arizona Bulletin Volume 60.Google Scholar
Gierliński, G. D., Lockley, M. G. & Niedzwiedzki, G. 2009. A distinctive crouching theropod trace from the Lower Jurassic of Poland. Geological Quarterly 53, 471–6.Google Scholar
Godefroit, P., Cau, A., Dong-YU, H., Escuillié, F., Wenhao, W. & Dyke, G. 2013. A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds. Nature 498, 359–62.Google Scholar
Hammer, Ø. 2015. Paleontological Statistics (PAST) 2.06. Oslo, Norway: University of Oslo. http://folk.uio.no/ohammer/past/.Google Scholar
Hammer, Ø. & Harper, D. A. T. 2006. Paleontological Data Analysis. Oxford: Blackwell Publishing Ltd, 351 pp.Google Scholar
Holtz, T. R. 1994. The arctometatarsalian pes, an unusual structure of the metatarsus of Cretaceous Theropoda (Dinosauria: Saurischia). Journal of Vertebrate Paleontology 14, 480519.Google Scholar
HU, D., Hou, L., Zhang, L. & XU, X. 2009. A pre-Archaeopteryx troodontid theropod from China with long feathers on the metatarsus. Nature 461, 640–3.Google Scholar
Kim, J. Y., Kim, K. S., Lockley, M. G., Yang, S. Y., Seo, S. J., Choi, H. I. & Lim, J. D. 2008. New didactyl dinosaur footprints (Dromaeosauripus hamanensis ichnogen. et ichnosp. nov.) from the Early Cretaceous Haman Formation, south coast of Korea. Palaeogeography. Palaeoclimatology, Palaeoecology 262, 72–8.Google Scholar
Kim, J. Y., Lockley, M. G., Woo, J. O. & Kim, S. H. 2012. Unusual didactyl traces from the Jinju Formation (Early Cretaceous, South Korea) indicate a new ichnospecies of Dromaeosauripus . Ichnos 19, 7583.Google Scholar
Kirkland, J. I., Britt, B. B., Whittle, C. H., Madsen, S. K. & Burge, D. L. 1998. A small coelurosaurian theropod from the Yellow Cat Memeber of the Cedar Mountain Formation (Lower Cretaceous, Barremian) of Eartern Utah. In Lower and Middle Cretaceous Terrestrial Ecosystems (eds Lucas, S. G., Kirkland, J. I. & Estep, J. W.), pp. 239–48. New Mexico Museum of Natural History and Science Bulletin no. 14.Google Scholar
Kuban, G. 1989. Elongate dinosaur tracks. In Dinosaur Tracks and Traces (eds Gillette, D. D. & Lockley, M. G.), pp. 5779. Cambridge: Cambridge University Press.Google Scholar
Lee, Y. N., Barsbold, R., Currie, P. J., Kobayashi, Y., Lee, H. J., Godefroit, P., Escuillie, F. & Chinzorig, T. 2014. Resolving the long-standing enigmas of a giant ornithomimosaurs Deinocheirus mirificus . Nature 515, 257–60.Google Scholar
LI, D., Azuma, Y., Fujita, 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. Journal of the Paleontological Society of Korea 22, 2949.Google Scholar
LI, R. & Lockley, M. G. 2005. Dromaeosaurid trackways from Shandong Province China. Journal of Vertebrate Paleontology 25, 84A.Google Scholar
LI, R., Lockley, M. G., Makovicky, P. J., Matsukawa, M., Norell, M. A., Harris, J. D. & Liu, M. 2008. Behavioral and faunal implications of Early Cretaceous deinonychosaur trackways from China. Naturwissenschaften 95, 185–91.Google Scholar
Lockley, M. G. 1991. Tracking Dinosaurs: A New Look at an Ancient World. Cambridge: Cambridge University Press, 238 pp.Google Scholar
Lockley, M. G., Hunt, A. P., Meyer, C., Rainforth, E. C. & Schultz, R. J. 1998. A survey of fossil footprint sites at Glen Canyon National Recreation Area (western USA): a case study in documentation of trace fossil resources at a national preserve. Ichnos 5, 177211.Google Scholar
Lockley, M. G., Matsukawa, M. & LI, J. 2003. Crouching theropods in taxonomic jungles: ichnological and ichnotaxonomic investigations of footprints with metatarsal and ischial impressions. Ichnos 10, 169–77.Google Scholar
Lockley, M. G., Matsukawa, M. & Witt, D. 2006. Giant theropod tracks from the Cretaceous Dakota Group northeastern New Mexico. New Mexico Museum of Natural History & Science Bulletin 35, 83–7.Google Scholar
Lockley, M. G., Meyer, C. A., Faria Dos Santos, V. 1998. Megalosauripus and the problematic concept of Megalosaur footprints. Gaia 15, 313–37.Google Scholar
Lockley, M. G., White, D., Kirkland, J. & Santucci, V. 2004. Dinosaur tracks from the Cedar Mountain Formation (Lower Cretaceous), Arches National Park, Utah. Ichnos 11, 285–93.Google Scholar
Longrich, N. 2008. A new, large ornithomimid from the Cretaceous Dinosaur Park Formation of Alberta, Canada: implications for the study of dissociated dinosaur remains. Palaeontology 51, 983–97.Google Scholar
Lucas, S. G., Klein, H., Lockley, M. G., Spielmann, J. A., Gierliński, G. D., Hunt, A. P. & Tanner, L. H. 2006. Triassic–Jurassic stratigraphic distribution of the theropod footprint ichnogenus Eubrontes . In The Triassic–Jurassic Terrestrial Transition (eds Harris, J. D., Lucas, S. G., Spielmann, J. A., Lockley, M. G., Milner, A. R. C. & Kirkland, J. I.), pp. 8693. New Mexico Museum of Natural History and Science Bulletin Volume 37.Google Scholar
Madsen, J. H. 1993. Allosaurus fragilis: a revised osteology. Utah Geological Survey Bulletin 109, 1163.Google Scholar
Makovicky, P. J., Kobayashi, Y. & Currie, P. J. 2004. Ornithomimosauria. In The Dinosauria. Second Edition (eds Weishampel, D. B., Dodson, P. & Osmólska, H.), pp. 137–50. Berkeley: University of California Press.Google Scholar
Manning, P. L., Margetts, L., Johnson, M. R., Withers, P. J., Sellers, W. I., Falkingham, P. L., Mummery, P. M., Barrett, P. M. & Raymont, D. R. 2009. Biomechanics of dromaeosaurid dinosaur claws: application of X-ray microtomography, nano-indentation, and finite element analysis. The Anatomical Record 292, 1397–405.Google Scholar
Martin, L. D. & Zhou, Z. 1998. Confuciusornis sanctus compared to Archaeopteryx lithographica . Naturwissenschaften 85, 286–89.Google Scholar
Marty, D., Strasser, A. & Meyer, C. A. 2009. Formation and taphonomy of human footprints in microbial mats of present-day tidal-flat environments: implications for the study of fossil footprints. Ichnos 16, 127–42.Google Scholar
Milàn, J. 2006. Variations in the morphology of emu (Dromaius novaehollandiae) tracks reflecting differences in walking pattern and substrate consistency: ichnotaxonomic implications. Palaeontology 49, 405–20.Google Scholar
Milán, J., Loope, D. B. & Bromley, R. G. 2008. Crouching theropod and Navahopus sauropodomorph tracks from the Early Jurassic Navajo Sandstone of USA. Acta Palaeontologica Polonica 53, 197205.Google Scholar
Milner, A. R. C., Harris, J. D., Lockley, M. G., Kirkland, J. I. & Matthews, N. A. 2009. Birdlike anatomy, posture, and behavior revealed by an Early Jurassic theropod dinosaur resting trace. PLoS ONE 4, e4591. doi: 10.1371/journal.pone.0004591.Google Scholar
Mostardini, F. & Merlini, S. 1986. Appennino centro-meridionale: sezione geologiche e proposta di modello strutturale. AGIP, 73rd Congresso della Societa Geologica Italiana, Roma, 59 pp.Google Scholar
Mudroch, A., Richter, U., Joger, U., Kosma, R., Idé, O. & Maga, A. 2011. Didactyl tracks of paravian theropods (Maniraptora) from the ?Middle Jurassic of Africa. PLoS ONE 6, e14642. doi: 10.1371/journal.pone.0014642.Google Scholar
Nicosia, U., Petti, F. M., Perugini, G., D'Orazi porchetti, S., Sacchi, E., Conti, M. A., Mariotti, N. & Zarattini, A. 2007. Dinosaur tracks as paleogeographic constraints: new scenarios for the Cretaceous geography of the periadriatic region. Ichnos 14, 6990.Google Scholar
Osmólska, H. 1981. Coossified tarsometatarsi in theropod dinosaurs and their bearing on the problem of bird origins. In Results of the Polish−Mongolian Palaeontological Expeditions—Part IX (ed. Kielan-Jaworowska, Z.), pp. 7995. Palaeontologia Polonica 42.Google Scholar
Petti, F. M., Conti, M. A., D'orazi Porchetti, S., Morsilli, M., Nicosia, U. & Gianolla, P. 2008. A theropod dominated ichnocoenosis from the Late Hauterivian–Early Barremian of Borgo Celano (Gargano promontory, Apulia, Southern Italy). Rivista Italiana di Paleontologia e Stratigrafia 114, 317.Google Scholar
Pescatore, T., Renda, P., Schiattarella, M. & Tramutoli, M. 1999. Stratigraphic and structural relationship between Meso-Cenozoic Lagonegro basin and coeval carbonate platforms in Southern Apennines, Italy. Tectonophysics 315, 269–86.Google Scholar
PU, H., Chang, H., , J., WU, Y., XU, L., Zhang, J. & Jia, S. 2013. A new juvenile specimen of Sapeornis (Pygostylia: Aves) from the Lower Cretaceous of Northeast China and allometric scaling of this basal bird. Paleontological Research 17, 2738.Google Scholar
Qiang, J., Chiappe, L. M. & Shu'AN, J. 1999. A new Late Mesozoic confuciusornithid bird from China. Journal of Vertebrate Paleontology 19, 17.Google Scholar
Qiang, J., Currie, P. J., Norell, M. A. & Shu-An, J. 1998. Two feathered dinosaurs from northeastern China. Nature 393, 753–61.Google Scholar
Romano, M. & Citton, P. 2015. Reliability of digit length impression as a character of tetrapod ichnotaxobase: considerations from the Carboniferous–Permian ichnogenus Ichniotherium . Geological Journal 50, 827–38.Google Scholar
Romano, M., Citton, P. & Nicosia, U. 2016. Corroborating trackmaker identification through footprint functional analysis: the case study of Ichniotherium and Dimetropus . Lethaia 49, 102–16.Google Scholar
Romero-Molina, M., Sarjeant, W., Pérez-Lorente, F., Lopez, A. & Requeta, E. 2003. Orientation and characteristics of theropod trackways from the Las Losas Palaeoichnological Site (La Rioja, Spain). Ichnos 10, 241–54.Google Scholar
Sacchi, E., Conti, M. A., D'Orazi Porchetti, S., Logoluso, A., Nicosia, U., Perugini, G. & Petti, F. M. 2009. Aptian dinosaur footprints from the Apulian platform (Bisceglie, Southern Italy) in the framework of periadriatic ichnosites. Palaeogeography, Palaeoclimatology. Palaeoecology 271, 104–16.Google Scholar
Sarjeant, W. A. S. 1971. Vertebrate tracks from the Texas of Castle Peak, Texas. The Texas Journal of Science 22, 343–66.Google Scholar
Senter, P. 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology 5, 429–63.Google Scholar
Smith, J. B., Farlow, J. O. 2003. Osteometric approaches to trackmaker assignment for the Newark Supergroup ichnogenera Grallator, Anchisauripus, and Eubrontes . In The Great Rift Valley of Pangea in Eastern North America. Sedimentology, Stratigraphy and Paleontology Volume 2 (eds LeTourneau, P. M. & Olsen, P. E.), pp. 273–92. New York: Columbia University Press.Google Scholar
Thulborn, R. A. 1990. Dinosaur Tracks. London: Chapman and Hall, 410 pp.Google Scholar
Thulborn, R. A. 2013. Lark Quarry revisited: a critique of methods used to identify a large dinosaurian track-maker in the Winton Formation (Albian–Cenomanian), western Queensland, Australia. Alcheringa 37, 312–30.Google Scholar
Thulborn, R. A. & Wade, M. 1989. A footprint as a history of movement. In Dinosaur Tracks and Traces (eds Gillette, D. D. & Lockley, M. G.), pp. 51–6. Cambridge: Cambridge University Press.Google Scholar
Turco, E., Schettino, A., Nicosia, U., Santantonio, M., Di stefano, P., Iannace, A., Cannata, D., Conti, M. A., Deiana, G., D'Orazi Porchetti, S., Felici, F., Liotta, D., Mariotti, N., Milia, A., Petti, F. M., Pierantoni, P. P., Sacchi, E., Sbrescia, V., Tommasetti, K., Valentini, M., Zamparelli, V. & Zarcone, G. 2007. Mesozoic paleogeography of the Central Mediterranean Region. Geoitalia 2007, VI Forum Italiano di Scienze della Terra. Epitome 2, 108.Google Scholar
Vickers-Rich, P., Chiappe, L. M. & Kurzanov, S. 2002. The enigmatic bird-like dinosaur Avimimus portentosus . In Mesozoic Birds (eds Chiappe, L. M. & Witmer, L. M.), pp. 6586. Berkeley: University of California Press.Google Scholar
Weishampel, D. B., Dodson, P. & Osmólska, H. (eds) 1990. The Dinosauria. Berkeley: University of California Press, 733 pp.Google Scholar
Weishampel, D. B., Dodson, P. & Osmólska, H. (eds) 2004. The Dinosauria. Second Edition. Berkeley: University of California Press, 861 pp.Google Scholar
White, M. A. 2009. The subarctometatarsus: intermediate metatarsus architecture demonstrating the evolution of the arctometatarsus and advanced agility in theropod dinosaurs. Alcheringa 33, 121.Google Scholar
Wilson, J. A., Marsicano, C. A. & Smith, R. M. H. 2009. Dynamic locomotor capabilities revealed by early dinosaur trackmakers from Southern Africa. PLoS ONE 4, e7331. doi: 10.1371/journal.pone.0007331.Google Scholar
XU, X., Wang, X. L., WU, X. C. 1999. A dromaeosaurid dinosaur with a filamentous integument from the Yixian Formation of China. Nature 401, 262–6.Google Scholar
Zarcone, G., Petti, F. M., Cillari, A., Di stefano, P., Guzzetta, D. & Nicosia, U. 2010. A possible bridge between Adria and Africa: new palaeobiogeographic and stratigraphic constraints on the Mesozoic palaeogeography of the Central Mediterranean area. Earth-Science Reviews 103, 154–62.Google Scholar
Zhen, S., LI, J., Zhang, B., Chen, W. & Zhu, S. 1995. Dinosaur and bird footprints from the Lower Cretaceous of Emei County, Sichuan. Memoirs of Beijing Natural History Museum 54, 105–20.Google Scholar
Supplementary material: File

Romano and Citton supplementary material

Figures S1-S3

Download Romano and Citton supplementary material(File)
File 547.8 KB