Scampering, trotting, walking tridactyl bipedal dinosaurs in southern Africa: ichnological account of a Lower Jurassic palaeosurface (upper Elliot Formation, Roma Valley) in Lesotho

References

• Alexander RMcN. 1976. Estimates of speeds of dinosaurs. Nature. 261:129–130.10.1038/261129a0
   Web of Science ®Google Scholar
• Ambrose D. 2003. A note on fossil trackways at Roma. Lesotho Miscellaneous Documents No. 4, House 9 Publications Roma, iv+ 14 pp.
   Google Scholar
• Avnimelech MA. 1966. Dinosaur tracks in the Judean Hills. Proceedings of the Israel Academy of Sciences and Humanities, Section of Sciences. 1:1–19.
   Google Scholar
• Avanzini M, Pinuela L, Garcia-Ramos JC. 2012. Late Jurassic footprints reveal walking kinematics of theropod dinosaurs. Lethaia. 45:238–252.
   Web of Science ®Google Scholar
• Belvedere M, Dyke G, Hadri M, Ishigaki S. 2011. The oldest evidence for birds in Northern Gondwana? Small tridactyl footprints from the Middle Jurassic of Msemrir (Morocco). Gondwana Res. 19:542–549.10.1016/j.gr.2010.08.004
   Web of Science ®Google Scholar
• Blackburn TJ, Olsen PE, Bowring SA, McLean NM, Kent DV, Puffer J, McHone G, Rasbury ET, Touhami M. 2013. Zircon U-Pb geochronology links the end-Triassic extinction with the Central Atlantic Magmatic Province. Science. 340:941–945.10.1126/science.1234204
   PubMed Web of Science ®Google Scholar
• Bordy EM, Eriksson P. 2015. Lithostratigraphy of the Elliot Formation (Karoo Supergroup), South Africa. S Afr J Geol. 118:311–316.10.2113/gssajg.118.3.311
   Web of Science ®Google Scholar
• Bordy EM, Hancox PJ, Rubidge BS. 2004a. Fluvial style variations in the Late Triassic–Early Jurassic Elliot formation, main Karoo Basin, South Africa. J Afr Earth Sci. 38:383–400.10.1016/j.jafrearsci.2004.02.004
   Web of Science ®Google Scholar
• Bordy EM, Hancox PJ, Rubidge BS. 2004b. Basin development during the deposition of the Elliot Formation (Late Triassic – Early Jurassic), Karoo Supergroup, South Africa. S Afr J Geol. 107:395–410.
   Web of Science ®Google Scholar
• Bordy EM, Hancox PJ, Rubidge BS. 2004c. Provenance Study of the Late Triassic – Early Jurassic Elliot Formation, main Karoo Basin, South Africa. S Afr J Geol. 107:587–602.10.2113/gssajg.107.4.587
   Web of Science ®Google Scholar
• Bordy EM, Hancox PJ, Rubidge BS. 2004d. A description of the sedimentology and palaeontology of the Upper Triassic – Lower Jurassic Elliot Formation in Lesotho. Palaeontol Afr. 40:43–58.
   Google Scholar
• Bordy EM, Hancox PJ, Rubidge BS. 2005. The contact of the Molteno and Elliot formations through the main Karoo Basin, South Africa: a second-order sequence boundary. S Afr J Geol. 108:349–362.
   Web of Science ®Google Scholar
• Bordy EM, Sciscio L, Abdala F, McPhee B, Choiniere J. 2016. First Lower Jurassic vertebrate burrow from southern Africa (upper Elliot Formation, Karoo Basin, South Africa). Palaeogeogr Palaeoecol. doi:10.1016/j.palaeo.2016.12.024.
   Web of Science ®Google Scholar
• Botha BJV. 1967. The stratigraphy of the Red Beds Stage, Karoo System at Elliot, Cape Province. Trans Geol Soc S Afr. 71:101–117.
   Google Scholar
• Bristowe A, Raath M. 2004. A juvenile coelophysoid skull from the Early Jurassic of Zimbabwe, and the synonymy of Coelophysis and Syntarsus. Palaeontol Afr. 40:31–41.
   Google Scholar
• Dalman SG, Weems RE. 2013. A new look at morphological variation in the ichnogenus Anomoepus, with special reference to material from the Lower Jurassic Newark Supergroup: implications for ichnotaxonomy and ichnodiversity. Bull Peabody Mus Nat Hist. 54:67–124.10.3374/014.054.0104
   Web of Science ®Google Scholar
• Day JJ, Norman DB, Gale AS, Upchurch P, Powell HP. 2004. A middle Jurassic dinosaur trackway site from Oxfordshire, UK. Palaeontology. 47:319–348.10.1111/pala.2004.47.issue-2
   Web of Science ®Google Scholar
• de Valais S. 2011. Revision of dinosaur ichnotaxa from the La Matilde formation (middle Jurassic), Santa Cruz province, Argentina. Ameghiniana. 48:28–42.10.5710/AMGH.v48i1(244)
   Web of Science ®Google Scholar
• de Valais S, Melchor RN. 2008. Ichnotaxonomy of bird-like footprints s: an example from the Late Triassic-Early Jurassic of Northwest Argentina. J Vertebr Paleontol. 28:14–159.
   Web of Science ®Google Scholar
• D’orazi Porchetti S, Nicosia U. 2007. Re-examination of some large early Mesozoic tetrapod footprints from the African collection of Paul Ellenberger. Ichnos. 14:219–245.10.1080/10420940601049990
   Google Scholar
• Duncan RA, Hooper PR, Rehacek J, Marsh JS, Duncan AR. 1997. The timing and duration of the Karoo igneous event, southern Gondwana. J Geophys Res-Sol Earth. 102:18127–18138.10.1029/97JB00972
   Web of Science ®Google Scholar
• Ellenberger F, Ellenberger P, Fabre J, Ginsburg L, Mendrez C. 1964. The Stormberg series of Basutoland (South Africa). In: Proceedings of the XXII International Geological Congress. New Delhi; p. 320–330.
   Google Scholar
• Ellenberger P. 1970. The fossil-bearing strata associated with the earliest appearance of mammals in South Africa and their ichnology: establishment of detailed stratigraphic zones in the Stormberg of Lesotho (South Africa)(Upper Triassic to Jurassic). Proceedings and Papers of the second Gondwana Symposium; p. 343–370.
   Google Scholar
• Ellenberger P. 1972. Contribution to the classification of Triassic vertebrate trackways: types found in the Stormberg Series of South Africa (I). Paleovertebrata, Memoire Extraordinaire 1972, Montpellier; p. 152.
   Google Scholar
• Ellenberger P. 1974. Contribution to the classification of vertebrate tracks of the Triassic: the types of the Stormberg of South Africa, (2). Palaeovertebrata, Memoire Extraordinaire 1974, Montpellier; p. 170.
   Google Scholar
• Ellenberger P. 1978. The origin of birds: history and new techniques; the problem of the Archaeornithes; the coming to light of Cosesaurus aviceps (Upper Muschelkalk). Mém. Trav. Ecole prat. Hautes Etud., Inst. 4:91–117.
   Google Scholar
• Ellenberger P. 1993. Cosesaurus aviceps: Aviform vertebrate of the middle Triassic of Catalonia. Mém: Univ. Sci. Tech. Languedoc; p. 741.
   Google Scholar
• Fastovsky DE, Smith JB. 2004. Dinosaur paleoecology. In: Weishample P, Dodson P, Osmolska H, editors. The Dinosauria. 2nd ed. Berkeley: University of California Press; p. 614–626.
   Google Scholar
• Gand G, Demathieu G, Grancier M, Sciau J. 2005. Dinosauroid tracks of French Upper Triassic. Discrimination, interpretation and comparison. Bulle Soc Geol Fr. 176:69–79.10.2113/176.1.69
   Web of Science ®Google Scholar
• Gierlínski G. 1996. Avialian theropod tracks from the early Jurassic strata of Poland. Zubia. 14:79–87.
   Google Scholar
• Gierlínski G. 2016. Middle Jurassic avialan footprints from Imilchil in Morocco. In: Baucon A, Neto de Carvalho C, Rodrigues J. Ichnia. 2016: Abstract book. UNESCO Geopark Naturtejo/International Ichnological Association, Castelo Branco; p. 98–99.
   Google Scholar
• Gierlínski G, Sabbath K. 2008. Stegosaurian footprints from the Morrison formation of Utah and their implications for interpreting other ornithischian tracks. Oryctos. 8:29–46.
   Google Scholar
• Henderson D. 2003. Tracks, trackways, and hip heights of bipedal dinosaurs – testing hip height predictions with computer models. Ichnos. 10:99–114.10.1080/10420940390257914
   Google Scholar
• Klein H, Lucas SG. 2010. Tetrapod footprints – their use in biostratigraphy and biochronology of the Triassic. In: Lucas SG, editor. The Triassic timescale. Geological Society of London: Special publications; p. 419–446.
   Google Scholar
• Kim BS, Huh M. 2010. Analysis of the acceleration phase of a theropod dinosaur based on a Cretaceous trackway from Korea. Palaeogeogr Palaeoecol. 293:1–8.10.1016/j.palaeo.2010.04.020
   Web of Science ®Google Scholar
• Kitching JW, Raath MA. 1984. Fossils from the Elliot and Clarens formations (Karoo sequence) of the northeastern Cape, Orange Free State and Lesotho, and a suggested biozonation based on tetrapods. Palaeontol Afr. 25:111–125.
   Google Scholar
• Knoll F. 2004. Review of the tetrapod fauna of the “Lower Stormberg Group” of the main Karoo Basin (southern Africa): implication for the age of the Lower Elliot Formation. Bull Soci Géol Fr. 175:73–83.10.2113/175.1.73
   Web of Science ®Google Scholar
• Knoll F. 2005. The tetrapod fauna of the Upper Elliot and Clarens formations in the main Karoo Basin (South Africa and Lesotho). Bull Soc Géol Fr. 176:81–91.10.2113/176.1.81
   Web of Science ®Google Scholar
• Knoll F, Padian K, de Ricqlès A. 2010. Ontogenetic change and adult body size of the early ornithischian dinosaur Lesothosaurus diagnosticus: implications for basal ornithischian taxonomy. Gondwana Res. 17:171–179.10.1016/j.gr.2009.03.010
   Web of Science ®Google Scholar
• Lockley MG. 2009. New perspectives on morphological variation in tridactyl footprints: clues to widespread convergence in developmental dynamics. Geol Q. 53:415–432.
   Web of Science ®Google Scholar
• Lockley MG, Cart K, Martin J, Milner A. 2011. New theropod tracksites from the Upper Cretaceous “Mesaverde” Group, Western Colorado: implications for ornithomimosaur track morphology. New Mex Mus Nat Hist Sci Bul. 53:231–329.
   Google Scholar
• Lockley MG, Gierlínski G. 2006. Diverse vertebrate ichnofaunas containing Anomoepus and other unusual trace fossils from the Lower Jurassic of the Western United States: implication from paleoecology and palichnostratigraphy. New Mex Mus Nat Hist Sci Bul. 37:171–191.
   Google Scholar
• Lockley MG, Hunt AP. 1995. Dinosaur tracks and other fossil footprints of the Western United States. New York, NY: Columbia University Press; p. 338.
   Google Scholar
• Lockley MG, Hunt AP, Paquette M, Bilbey S, Hamblin A. 1998. Dinosaur tracks from the Carmel formation, northeastern Utah: implications from middle Jurassic paleoecology. Ichnos. 5:255–267.10.1080/10420949809386424
   Google Scholar
• Lockley MG, Meyer CA. 1999. Dinosaur tracks and other fossil footprints of Europe. New York (NY): Columbia University Press.
   Google Scholar
• Lockley MG, Rainforth EC. 2002. The track record of Mesozoic birds and pterosaurs: An ichnological and paleoecological perspective. In: Chiappe LM, Witmer LM, editors. Mesozoic birds above the heads of dinosaurs. Berkeley: University of California Press; p. 405–418.
   Google Scholar
• Lockley MG, Wright JL, Matsukawa M. 2001. A new look at Magnoavipes and so-called ‘Big Bird’ tracks from Dinosaur Ridge (Cretaceous, Colorado). Mountain Geol. 38:137–146.
   Google Scholar
• Lockley MG, Yang SY, Mutsukawa M, Fleming F, Lim SK. 1992. The track record of Mesozoic birds: evidence and implications. Philos Trans R Soc. B. 336:113–134.10.1098/rstb.1992.0051
   Web of Science ®Google Scholar
• Lockley MG, Young BH, Carpenter K. 1983. Hadrosaur locomotion and herding behavior: evidence from footprints in the Mesaverde Formation, Grand Mesa coal field, Colorado. Mountain Geol. 2:5–14.
   Google Scholar
• Lucas SG, Hancox PJ. 2001. Tetrapod-based correlation of the nonmarine Upper Triassic of southern Africa. Albertiana. 25:5–9.
   Google Scholar
• Lucas SG, Klein H, Lockley MG, Spielmann JA, Gierlínski GD, Hunt AP, Tanner LH. 2006. Triassic-Jurassic stratigraphic distribution of the theropod footprint ichnogenus Eubrontes. New Mex MusNat Hist Sci Bul. 37:86–93.
   Google Scholar
• Lucas SG, Tanner LH. 2015. End-Triassic nonmarine biotic events. J Palaeogeogr. 4:331–348.
   Web of Science ®Google Scholar
• Mallison H, Wings O. 2014. Photogrammetry in paleontology – a practical guide. J Pal Tech. 12:1e31.
   Google Scholar
• Mariotti G, Pruss B, Perron JT, Bosak T. 2014. Microbial shaping of sedimentary wrinkle structures. Nat Geosci. 7:736–740.10.1038/ngeo2229
   Web of Science ®Google Scholar
• Melchor RN, Buchwaldt R, Bowring SA. 2013. A Late Eocene date for Late Triassic bird tracks. Nature. 495:E1–E2.10.1038/nature11931
   PubMed Web of Science ®Google Scholar
• Melchor RN, de Valais S, Genise JF. 2002. Bird-like fossil footprints from the Late Triassic. Nature. 417:936–938.10.1038/nature00818
   PubMed Web of Science ®Google Scholar
• Munyikwa D, Raath MA. 1999. Further material of the ceratosaurian dinosaur Syntarsus from the Elliot Formation (Early Jurassic) of South Africa. Palaeontol Afr. 35:55–59.
   Google Scholar
• Olsen PE, Galton PM. 1984. A review of the reptile and amphibian assemblages from the Stormberg of southern Africa, with special emphasis on the footprints and the age of the Stormberg. Palaeontol Afr. 25:87–110.
   Google Scholar
• Olsen PE, Kent DV, Whiteside JH. 2011. Implications of the Newark Supergroup-based astrochronology and geomagnetic polarity time scale (Newark-APTS) for the tempo and mode of the early diversification of the Dinosauria. Earth Environ. Sci. Trans. R. Soc. Edinburgh. 101:201–229.
   Web of Science ®Google Scholar
• Olsen PE, Rainforth EC. 2003. The Early Jurassic Ornithischian sinosaurian ichnogenus Anomoepus. In: Le Tourneau RM, Olsen PE, editors. The great rift valleys of Pangea in Eastern North America. New York, NY: Columbia University Press; p. 314–367.
   Google Scholar
• Olsen PE, Smith JB, McDonald NG. 1998. Type material of the type species of the classic theropod footprint genera Eubrontes, Anchisauripus, and Grallator (Early Jurassic, Hartford and Deerfield basins, Connecticut and Massachusetts, USA). J Vertebr Paleontol. 18:586–601.10.1080/02724634.1998.10011086
   Web of Science ®Google Scholar
• Parizot M, Eriksson PG, Aifa T, Sarkar S, Banerjee S, Catuneanu O, Alterman W, Bumby AJ, Bordy EM, Van Rooy JL, Boshoff AJ. 2005. Suspected microbial mat-related crack-like sedimentary structures in the Palaeoproterozoic Magaliesberg Formation sandstones,South Africa. Precambrian Res. 138:274–296.10.1016/j.precamres.2005.05.007
   Web of Science ®Google Scholar
• Raath MA. 1977. The anatomy of the Triassic theropod Syntarsus rhodesiensis (Saurischia: Podokesauridae) and a consideration of its biology [ PhD thesis]. Grahamstown: Rhodes University.
   Google Scholar
• Raath MA. 1980. The theropod dinosaur Syntarsus (Saurischia: Podokesauridae) discovered in South Africa. S Afr J Sci. 76:375–376.
   Web of Science ®Google Scholar
• Raath MA, Yates AM. 2005. Preliminary report of a large theropod dinosaur trackway in Clarens Formation sandstone (Early Jurassic) in the Paul Roux district, northeastern Free State, South Africa. Palaeontol Afr. 41:101–104.
   Google Scholar
• Rainforth EC. 1997. Vertebrate ichnological diversity and census studies Lower Jurassic Navajo Sandstone. Boulder: University of Colorado; p. 48.
   Google Scholar
• Rainforth EC. 2001. Late Triassic – Early Jurassic dinosaur ichnofaunas, eastern North America and Southern Africa. J Vertebr Palaeontol. 21:91A.
   Google Scholar
• Rainforth EC. 2005. Ichnotaxonomy of the fossil footprints of the Connecticut Valley (Early Jurassic, Newark Supergroup, Connecticut and Massachusetts) [ PhD thesis]. New York (NY): Columbia University.
   Google Scholar
• Rainforth EC, Manzella M. 2007. Estimating speeds of dinosaurs from trackways: a re-evaluation of assumptions. Contributions to the Paleontology of New Jersey (II): Field Guide and Proceedings, Geological Association of New Jersey 24th Annual Conference and Field Trip, 41–48.
   Google Scholar
• Ruiz J, Torices A. 2013. Humans running at stadiums and beaches and the accuracy of speed estimations from fossil trackways. Ichnos. 20:31–35.10.1080/10420940.2012.759115
   Web of Science ®Google Scholar
• Schieber J, Bose PK, Eriksson PG, Banerjee S, Sakar S, Altermann W, Catuneanu O. 2007. Atlas of microbial mat features preserved within the siliciclastic rock record. Italy: Elsevier.
   Google Scholar
• Schneider CA, Rasband WS, Eliceiri KW. 2012. NIH Image to ImageJ: 25 years of image analysis. Nature Methods. 9:671–675.10.1038/nmeth.2089
   PubMed Web of Science ®Google Scholar
• Sciscio L, Bordy EM. 2016. Palaeoclimatic conditions in the Late Triassic-Early Jurassic of southern Africa: a geochemical assessment of the Elliot Formation. J Afr Earth Sci. 119:102–119.10.1016/j.jafrearsci.2016.03.014
   Web of Science ®Google Scholar
• Sciscio L, Bordy EM, Reid M, Abrahams M. 2016. Sedimentology and ichnology of the Mafube dinosaur track site (Lower Jurassic, eastern Free State, South Africa): a report on footprint preservation and palaeoenvironment. Peer J. p.e2285. doi:10.7717/peerj.2285.
   PubMed Web of Science ®Google Scholar
• Sciscio L, de Kock MO, Bordy EM, Knoll F. Forthcoming. Magnetostratigraphy over the Triassic – Jurassic lower Elliot – upper Elliot Formation, Stormberg Group, Karoo Basin. Gondwana Res.
   Google Scholar
• Smith RM, Kitching J. 1997. Sedimentology and vertebrate taphonomy of the Tritylodon Acme Zone: a reworked palaeosol in the Lower Jurassic Elliot Formation, Karoo Supergroup, South Africa. Palaeogeogr Palaeoecol. 131:29–50.10.1016/S0031-0182(96)00143-5
   Web of Science ®Google Scholar
• Svensen H, Corfu F, Polteau S, Hammer Ø, Planke S. 2012. Rapid magma emplacement in the Karoo Large Igneous Province. Earth Planet Sci Lett. 325–326:1–9.10.1016/j.epsl.2012.01.015
   Web of Science ®Google Scholar
• Thulborn RA. 1981. Estimated speed of a giant bipedal dinosaur. Nature. 292:273–274.10.1038/292273c0
   Web of Science ®Google Scholar
• Thulborn RA. 1982. Speeds and gaits of dinosaurs. Palaeogrogr Palaeoecol. 38:227–256.10.1016/0031-0182(82)90005-0
   Web of Science ®Google Scholar
• Thulborn RA. 1984. Preferred gaits of bipedal dinosaurs. Alcheringa. 8:243–252.10.1080/03115518408618947
   Web of Science ®Google Scholar
• Thulborn RA. 1989. The gaits of dinosaurs. In: Gillete DO, Lockley MG, editors. Dinosaur tracks and traces. Cambridge: Cambridge University Press; p. 39–50.
   Google Scholar
• Thulborn RA. 1990. Dinosaur tracks. London: Chapman and Hall.10.1007/978-94-009-0409-5
   Google Scholar
• Thulborn RA. 1998. Australia’s earliest theropods: footprint evidence in the Ipswich Coal Measures (Upper Triassic) of Queensland. Gaia. 1:301–311.
   Google Scholar
• Thulborn RA, Wade M. 1984. Dinosaur trackways in the Winton Formation (mid Cretaceous) of Queensland. Mem Qd Mus. 21:413–517.
   Google Scholar
• Wilson JA, Marsicano CA, Smith RMH. 2009. Dynamic locomotor capabilities revealed by early dinosaur trackmakers from southern Africa. PLoS ONE. 4:e7331. doi:10.1371/journal.pone.0007331.
   PubMed Web of Science ®Google Scholar
• Yates AM. 2005. A new theropod dinosaur from the Early Jurassic of South Africa and its implications for the early evolution of theropods. Palaeontol Afr. 41:105–122.
   Google Scholar
• Xing LD, Lockley MG, Klein H, Falkingham PL, Kim JY, Mccrea RT, Zhang J, Persons WS IV, Wang T, Wang Z. 2016. First Early Jurassic small ornithischian tracks from the Yunnan province, south western China. Palaios. 31:516–524.10.2110/palo.2015.074
   Web of Science ®Google Scholar
• Xing LD, Lockley MG, Klein H, Gierlínski GD, Divay JD, Hu S-M, Zhang J-P, Ye Y, He Y-P. 2014. The non-avian theropod track Jialingpus from the Cretaceous of the Ordos Basin, China, with a revision of the type material: Implications for ichnotaxonomy and trackmaker morphology. Palaeoworld. 23:197–199.
   Web of Science ®Google Scholar