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The Widespread Distribution of a Late Jurassic Theropod with Well-Padded Feet

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The Widespread Distribution of a Late Jurassic Theropod with Well-Padded Feet GAIA N°15, LlSBOAlLISBON, DEZEMBRO/DECEMBER 1998, pp. 339-353 (ISSN: 0871-5424) THERANGOSPODUS: TRACKWAY EVIDENCE FOR THE WIDESPREAD DISTRIBUTION OF A LATE JURASSIC THEROPOD WITH WELL-PADDED FEET Martin G. LOCKLEY Geology Department, Campus Box 172, University of Colorado at Denver. P.O. Box 173364, DENVER, COLORADO 80217-3364. USA E-mail: [email protected] Christian A. MEYER Universitat Basel, Geologisch-palaontologisches Institut. Bernoullistrasse, 32, BASEL, CH-4056. SWITZERLAND E-mail: [email protected] Joaquin J. MORATALLA Unidad de Paleontologia, Departamento Biologia, Universidad Aut6noma de Madrid. CANTOBLANCO 28049, MADRID. SPAIN ABSTRACT: Assemblages of distinctive, medium·sized theropod tracks indicative of animals with well-padded feet are known from large Upper Jurassic samples of well-preserved mate­ rial from North America and Asia. These tracks, herein named Therangospodus pandemi­ cus, always reveal a lack of distinct, separate digital pads, regardless of whether they are preserved as casts or molds. We interpret this as consistent ichnological evidence of a fleshy fool. Similar tracks from the ?Upper Jurassic-?Lower Cretaceous of Spain, first na­ med Therangospodus onca/ensis, are also formally described. Although the fleshy nature of the foot of this trackmaker, makes it's individual digits appear comparable with those of ornithopod trackmakers, the elongate track and asymmetric postero-medial indentation and narrow trackway, indicate that it is probably of theropod an affinity. Tracks of this type provide an instructive lesson for ichnologists and paleontologist in general because they reveal that tracks are a record of flesh on foot bones, and need not necessarily be an accura­ te reflection of the morphology of foot skeletons. In this case to the best of our knowledge, the trackmaker is not known from the skeletal record. In North America and Asia the tracks are found in strata that has been dated close to the Oxfordian-Kimmeridgian boundary. In Europe the strata in which some ofthe tracks are found is less precisely dated, though some tracks but may be of the same age. Especially in North America and Asia, the tracks are as­ sociated with larger theropod tracks that have been variously referred to as "megalosaur tracks" or Mega/osauripus LESSERTISSEUR. This widespread co-occurrence of two theropod track ichnotaxa (i.e., the Mega/osauripus-Therangopodus assemblage) evidently has po­ tential utility for biostratigraphic correlation on a global scale. INTRODUCTION are quite distinct from others on the basis of morphology as well as size. For example no one Theropod tracks are traditionally regarded as would confuse classic Grallator cursorius somewhat conservative morphologically, and (HITCHCOCK, 1858) with Tyranosauripus pillmorei therefore hard to distinguish. This may be true, to (LOCKLEY & HUNT, 1994, 1995a). Similarly, the some extent, as reflected by proposals to regard case has recently been made that the Middle the ichnogenera Grallator-Anchisauripus and Jurassic theropod track Carme/opdus (LOCKLEY et Eubrontes as varieties (subgenera) of the same al., 1998) is a unique morphotype. As discussed ichnogenus Grallator (OLSEN, 1980). See LOCKLEY elsewhere in this volume, Upper Jurassic & HUNT (1995a) for a dissenting view. However Megalosaurid tracks (i.e., Megalosauripus, sensu recent studies suggest that some theropod tracks LOCKLEY, MEYER & SANTOS, 1996; 1998) are also 339 artigos/papers M.G. LOCKLEY; C.A. MEYER & J.J. MORATALLA distinct from named theropod track ichnotaxa from 1971; WEEMS, 1992), and the many other new earlier and later epochs. names introduced by ELLENBERGER (1972, 1974), have been valid and useful descriptions of distinct Thus, as more is learned about theropod tracks it morphology. On the contrary, these names have appears that their morphologies are much more generally been regarded with caution and variable through time than previously supposed skepticism (OLSEN & GALTON, 1984; HAUBOLD, (LOCKLEY & HUNT, 1995b), and that the perception 1986; LOCKLEY, 1986, 1991a; FARLOW, 1987; of morphological conservatism and uniformity THULBORN, 1990; LOCKLEY & HUNT, 1995a, through time is, in part, a function of lack of study of LOCKLEY & MEYER, 1999) especially when authors adequately large samples of well-preserved make little or no mention of, or comparison with, material (BAIRD, 1957) . In this regard , similar ichnotaxa elsewhere in deposits of the paleontologists can not point to one comprehensive same age. study of theropod tracks since the time of HITCHCOCK (1858) and LULL (1953), and sadly Recent studies in the western USA have re­ these studies are fraught with problems (personal vealed that theropod tracks are abundant in Middle communications from Baird, Farlow, Gierlinski, and Upper Jurassic formations, as well as through­ Olsen and others). James Farlow has recently out the Cretaceous (LOCKLEY & HUNT, 1995a). Ade­ initiated an attempt to standardize an osteometric quate material now exists to begin to describe some approach to the measurement offoot (pes) bones of of the more distinctive forms, that can be shown to tridactyl and tetradactyl dinosaurs (FARLOW & differ from the GAE plexus. This does not mean that LOCKLEY, 1993; FARLOW & CHAPMAN , 1997), but other theropod tracks, especially those more closely little of what has been published to date deals resembling GAE forms, or those from small, poorly explicitly with morphometric comparisons of actual preserved assemblages, may not continue to be re­ ichnofaunas from discrete assemblages at specific ferred to ct. Grallator sp. , indet. theropod track, or stratigraphic levels. An attempt to review these some other general category. Having recently rede­ problems from a systemic or philosophical fined megalosaurid tracks (Mega/osauripus) from a perspective is presented elsewhere in this volume well-defined stratigraphic interval in the Upper Ju­ (LOCKLEY, 1998). rassic (LOCKLEY, MEYER & SANTOS, 1996; 1998), we herein formally describe another distinctive thero­ The proliferation of names in the Grallator­ pod ichnite that can be assigned to the ichnogenus Anchisauripus-Eubrontes plexus (GAE) that began Therangospodus. As noted below, this ichnogenus with the work of HITCHCOCK (1858) and LULL, (1953) co-occurs with Mega/osauripus in many assem­ has mainly been extended to other Lower Jurassic blages. ichnofaunas through introduction of new ichnospecies names, for material from new sites STUDY MATERIAL (e.g. , LAPPARENT & MONTENAT, 1967; DEMATHIEU, 1990, 1993; DEMATHIEU & SCIAU , 1995; GIERLINSKI, Tracks herein assigned to Therangospodus pan­ 1991; GIERLINSKI & ALHBERG, 1994; IRBY, 1996). demicus were first discovered and reported from the This is notlo say that new ichnogenus names such ?Middle to Upper Jurassic Entrada-Summerville as Kayentapus and Dilophosauripus (WELLES, transition zone in eastern Utah (LOCKLEY, 1989, 20 em Therangospodus Fig. 1 - Comparison of line drawings shows the similarity. A - Therangospodus track from Utah (CU-MWC 188.14). B, e - Therangospodus tracks from Spain (MORATALLA, 1993). 340 THERANGOSPODUS: TRACK EVIDENCE OF A JURASSIC THEROPOD WITH WELL-PADDED FEET TABLE I Basic trackway measurements/parameters for Therangospodus trackways from North America and Asia. TRACKWAY LENGTH WITH DEPTH STEP STRIDE TW NUMBER P.A. ORIENT. KPA5 (22) 19 3 91-101 200 (180) 20 50° KPA 16 26 22 4.5 96-102 198 176 28 355° KPA25 27 22 3 98 198 170 28 260° KPA27 (28) 22 4 110 212 152 43 255° KPA30 27 22 2.5 90 181 170 35 60° FTS R1 21.7 19.6 53.4 107.5 168 (115")- FTS 2 (31 ) 22 (270°) MMA-3 32.6 22.3 3-7 106.5 213 36.5 25° MMA-5 31 21 (3) 98.5 193 45 320° MMA-7 30 21 3.5 102.5 211 22 25° MMB-2 26 19 2-4 85.5 172 24 300° MMB-3 26 20.5 4 104 202 38.5 320° MMC-1 26 (16) 5 92 183 30 30° MMC-4 24 (13.5) 79 163.5 23 320° MMC-7 28 (17) 2-3 95.5 188 145° MMO-1 31 22 1-4 116 230 46 352° MMO-2 27 18 2-3 106 225 29 120° MMO-3 22 14 3 91 180 49 225° MME-1 27 19 3-8 85 170 40.5 210° MME-2 33 20 2-5 89 184 39 50° MME-3 27 (17) 3-5 74 161 34.5 30° MME-4 30 20 1-2 109 212 40 10° MME-5 31 22 4-5 101.5 196 34 30° MMG-1 32 25 3-7 92 182 54 75° MMG-2 32 18.5 2-3 109 225 36.5 325° MMG-3 23 16 3 69 140 215° MMG-4 35 25 2-4 110 42 145° MMG-5 30 21 2-3 92 186 33 195° MMG-8 27 21 1-3 79 158 20° MMG-9 33 24 1-2 110 221 150° MMG-10 28 19 1-2 90 180 190° MMG-12 31 19.5 3-6 92 178 195 MMH-1 26 18 3 76.5 162 39 210° MMH-2 33 24 1.5 106.5 208 30.5 350° MMH-4 (26) 20 86 171 50.5 145° MMH-5 25 18 2-3 100 196 43 185 MMH-E 27 21.5 1.5-2 101 202 30 350 1991 a, b), in beds that are considered to be close to now known to extend over an area of more than 1000 the Oxfordian Kimmeridgian boundary (LOCKLEY, km' (LOCKLEY & HUNT, 1995a). These and other 1998: fig. 3). Subsequent studies revealed that studies have resulted in an extensive photographic these tracks are very abundant (TABLE I) in associa­ record ofthe site, as indicated by the number of ill us­ tion with a single surface that extends over a large trations detailed in the list of referred material. area known as the Moab Megatracksite (LOCKLEY & All authors and other ichnologists who have vis­ PITTMAN, 1989; LOCKLEY, 1991 a, b). This area is ited this megatracksite and been involved in collect- 341 M.G.
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