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. LOCKLEY; C.A. MEYER & J.J. MORATALLA

homa, we have recognized that the tracks fall into two distinct categories as follows: medium sized, narrow-gauge trackways, with moderately long steps (low foot length/step ratio) and impressions that indicate well-padded feet, without discrete pad impressions (=Therangospodus MORATALLA, 1993, as defined herein). By contrast other tracks (Mega­ fosauripus) are consistently much larger, with wide­ gauge and somewhat irregular trackways, short steps (high foot length/step ratios) and somewhat belter-defined pad impressions. 1 m One of the keys to recognizing the importance of these differences came in a recent study of tracli.s from the Khodja Pil Ata tracksite in Turkmenistan, Central Asia. At this large and well-exposed site two of us M.G.L. and CAM were able to observe multi­ ple trackways of both the small narrow-gauge form, and the large wide-gauge form , in sediments that have been dated as close to the O xfordian­ Kimmeridgian boundary. The large tracks from this area (Turkmenistan, Uzbekistan and Tadjikistan) have been given many different names as dis­ cussed elsewhere (LOCKLEY, MEYER & SANTOS , 1996, 1998). This intriguing evidence of an intercontinental correlation of not one but two track types, led us to examine the Upper Jurassic european track record. We established that very large Megafosauripus trackways have been described from well-dated Up­ per Jurassic (Oxfordian-Kimmeridgian) deposits in Portugal, and Spain. In Portugal we have not yet confidently established the presence of the smaller morphotype Therangospodus, though one possible Trackway no. 16 example is reported from rocks of the same age. We believe however, that tracks that show a remarkable resemblance to this smaller, well-padded variety had been described from the somewhat imprecisely Therangospodus dated Upper Jurassic-Lower Cretaceous Oncala Turkmenistan Group of the Cameros Basin, in Spain. These tracks, which had been named "Therangospodus onca/en­ sis" in an unpublished doctoral thesis (MORATALLA, Fig. 2 - Therangospodus tracks from the late Jurassic 1993), give us the ichnogenus name that we apply to of Central Asia. Top and center left (5/849 and 41849) trac­ the Upper Jurassic tracks from North America, Asia ings of specimens in the Tashkent Geological Museum, and Europe. Uzbekistan. Lower left and right tracing of individual foot­ print and trackway 16 from which it originates, from Khodja Thus a label now exists for both the large and the Pi! Ata site in Turkmenistan. The trackway is typical in hav­ small morphotypes, and we are able to speak of a ing elongate cigar- shaped pads with no discrete digital "Mega/osauripus-Therangospodus" track assem­ pad impressions, but is atypical in showing that the animal blage that can be correlated across three continents stopped during normal walking progression. (LOCKLEY, 1998). The exact age of the Therango­ spodus onca/ensis assemblage in Spain requires further investigation. In the sections that follow we ing material and data at various times have noted formally describe the material labeled Therango­ that the assemblage is unusual because it is entirely spodus (MORATALLA, 1993) and show that, based on dominated by theropod tracks. During the course of a substantial sample of well-preserved tracks that ongoing studies of the site, and other sites at the are morphologically quite distinct from any other same stratigraphic level, elsewhere in eastern Utah known theropod tracks, it is an ichnite that requires and the American west, including Arizona, and Okla-

342 THERANGOSPODUS: TRACK EVIDENCE OF A JURASSIC THEROPOD WITH WELL-PADDED FEET

accommodation in its own discrete ichnogenus and that is morphologically distinct (SARJEANT, 1989: ichnospecies. ICZN code). In addition PEABODY (1955) and BAIRD (1957), respectively have stressed the importance SYSTEMATIC DISCUSSION of adequate supplies of well-preserved material, in­ ctuding adequate trackway segments. We believe There are guidelines regarding the practice of that these latter criteria are met in the American, naming new fossil footprints. First and foremost is European and Asian samples described herein. For the standard practice of only naming new material historical consistency we adopt the name Therango-

Fig. 3 - Photograph of frequently-illustrated trackway from the Moab Megatracksite now Therangospodus pandemi­ GUS, consisting of 13 consecutive footprints (view from distal/youngestllast-formed track, towards proximal/oldestlfirst­ formed track). The last (youngest) five are preserved as a rubber mold and fibreglass replica (~holotype CU-MWC 186.5), see Figure 4.

343 M.G. LOCKLEY; C.A. MEYER & J.J. MORATALLA

but never named (LOCKLEY, 1989, 1991a, b). They I . were first illustrated using line drawings (LOCKLEY, ~-:'.. : . ; .- 1989: fig . 50.2), that reveal how the tracks bear a re ­ markable resemblance to line drawings in the doc­ toral thes.is of MORATALLA (1993: fig. 8.5.1); see Figures 1 and 6 herein. Similar tracks from nearby localities within the Moab megatracksite were sub­ sequently illustrated photographically (LOCKLEY, 1991a: pI. 4, 6). MORATALLA (1993: fig. 8.5.2) also provided photographic illustrations, and a system­ atic description, of Lower Cretaceous footprints, us­ ing the name Therangospodus oncalensis. Owing to the photogenic nature of one of the trackways from the Moab megatracksite that con­ sists of an exposure re vealing 13 consecutive steps (LOCKLEY, 1991 a: pI. 4, top left), this particular track­ way (Fig . 3 within) has been photographed many times, and has figured in a number of television documentaries. Excluding the identical photograph reprinted in LOCKLEY (1993a, b), the same trackway has been re-photographed from different angles (LOCKLEY, MATSUKAWA & OBATA, 1991: fig 12, 16;

Fig. 4 - Photograph of holotype trackway from proxi­ mal/oldesUfirst-formed to distal/youngesUlast-formed, with fibreglass replica (= holotype CU-MWC 186.5) corre­ sponding to same five tracks. Compare with Figure 3. spodus, first informally applied of tracks from Spain by MORATALLA (1993), and, for the first time provide a formal description or this ichnogenus. It is instructive to compare the first published de­ tailed line drawing of what we herein call Therango­ spodus (LOCKLEY, 1989: fig. 50.2; 1998: fig 3) with the first ichnite so named (Therangospodus sensu MORATALLA, 1993: fig. 8.5.1) as done in Figures 1 and 6 and TABLE I. Two of the authors (M.G.L. and J.J.M.) have examined the original tracks in both Spain and Utah, and M.G.L. and CAM. have com­ pared the original Jurassic tracks from Utah with those found in Turkmenistan. The former compari­ son clearly shows that the tracks are similar both in size, pad morphology and trackway parameters. Trackways from Turkmenistan also reveal tracks that are similar with respect to all these fundamental parameters (Fig. 2, TABLE I). Fig . 5 - Photograph of natural cast (CU-MWC 186.2) of a paratype, an original specimen (see Fig . 5 - G for line Therangospodus tracks have been known from drawing. Measurements given in TABLE I (see MME-1). the "Moab Megatracksite" area for almost a decade

344 THERANGOSPODUS: TRACK EVIDENCE OF A JURASSIC THEROPOD WITH WELL-PADDED FEET

.' / '. \ , I / '"'?, ~_./ ..... '-.... ~I "- .... A B C '<.,.

u... \ • o F

20cm

G H

Fig. 6 - Line drawings of Therangospodus footprints from the Upper Jurassic Summerville Formation of Utah (A-G) and a comparable footprintfrom Portugal (H). A. B - CU-MWC 188.10. These two natural casts occur side by side on the same block, facing in opposite directions, and overlap at the point shown by the semi-circular dotted line. C - CU-MWC 188.13, natural cast. D - CU-MWC 188.14, replica. E - Tracing of track from Butler Wash, Summerville Formation, near Blanding Utah. F - Tracing of tracks from Moab Megatracksite (site D, see TABLE I). G - CU-MWC, 186.2 (see Fig. 4). H - tridactyl track, from the Upper Jurassic of Sao Martinho do Porto (Portugal), that is comparable in size and general morphology to North American footprints. See text for details, and ApPENDIX for locality stratigraphic and information per­ taining to numbered specimens.

345 M.G. LOCKLEY; C.A. MEYER & J.J. MORATALLA

LOCKLEyet al. , 1991: fig. 4; LOCKLEY & HUNT , 1995: lished illustrations is given below in the section on fig. 4.37). The last five consecutive tracks (four referred material. steps) in the trackway have also been replicated and reposited in the joint University of Colorado at SYSTEMATIC DESCRIPTION Denver-Museum of Western Colorado collections as CU-MWC 186.5. Three consecutive tracks from Therangospodus ichnogen. nov. this specimen were illustrated by LOCKLEY et al. Fig. 1-7 (1992: fig. 13) and MORITA& MATSUKAWA(1996: fig. 57) and formed part of an international tracks ex­ Diagnosis: Medium sized, elongate, asymmet­ hibit. In all we estimate that trackways from three dif­ ric theropod track with coalesced, elongate, oval ferent but adjacent locations in the Moab mega­ digital pads, not separated into discrete phalangeal tracksite have been photographed or drawn for at pads. Trackway narrow with little or no rotation of least 20 separate published illustrations. Tracks digit III long axis from trackway axis. from a single site in Spain have appeared in two illus­ Etymology: Referring to a narrow footed thero­ trations in the same thesis. A complete list of pub- pod tracks (MORATALLA, 1993).

Fig. 7 - Type specimen of Therangospodus oncalensis, after MORATALLA (1993). Measurements given in TABLE I (see FTS-1).

346 THERANGOSPODUS: TRACK EVIDENCE OF A JURASSIC THEROPOD WITH WELL-PADDED FEET

Type ichnospecies: Therangospodus pande­ Etymology: Meaning pandemic, widespread or micus ichnosp. nov. (hoc opus). cosmopolitan in distribution. Other ichnospecies: The rangospodus on­ Type locality: Horizon and distribution as for ich­ calensis MORATALLA, 1993. npgenus. Type locality: Moab megatracksite site E (sensu Referred material: LOCKLEY, 1991a, b), west side of Arches National Monument, Utah. LOCKLEY, 1989: fig. 50.2. LOCKL EY, 1991a: pI. 4 top left, pi. 6 top left and right. Type horizon: Entrada (Moab Tongue Member)­ LOCKLEY, 1991b: fig. 2,3 and 5 in part. Summerville (upper tongue), transition zone, proba­ GOODKNIGHT & ERTEL, 1991: fig. 19.8d. biyearly Upper Jurassic (Oxfordian-Lower Kimme­ LOCKLEY et al., 1991: fig. 12, and fig.16 top left and lower left. ridg ian). LOCKlEY, 1993a, b: p1.4, 6(samefiguresasLOCKLEY, 1991a). LOCKlEyet aI, 1991: 4 (lower; same as LOCKLEyel al., 1991 ~ fig. Type specimen: CU-MWC 186.5, rubber mold 12), also p. 10 (upper same as LOCKLEY, 1991: pI. 6 upper and fibreglass replica of five consecutive footprints, right). and intervening rock surface from a single trackway LOCKLEY et al., 1992: fig . 13. of 13 consecutive steps, (Fig. 3-4), from Moab LOCKLEY & HUNT, 1995: fig. 4.37. megatracksite, site E. MORITA & MATSUKAWA. 1996: fig. 57. LOCKLEY, HUNT & LUCAS, 1996: 252, fig. 5 (right). Topotype: CU-MWC 186.2, from same surface LOCKLEY, 1998:431, fig. 3. at adjacent site G (Fig. 5). LOCKLEY, 1999:139, fig. 5.4 (top right). Paratypes: Upper part of Summerville Forma­ tion, Del Monte mines locality, north of Bullfrog Utah. CU-MWC 188.10 and 188.13 natural casts of three tracks. CU-MWC 188. 14 replica of natural cast (see TherangospoduS oncalensis Fig. 6) (MORATALLA, 1993) Amended Distribution: Lower part of Upper Jurassic of Fig . 1,7 North America, Europe and Central Asia. Systematic note: This ichnospecies was first North America: Moab Member-Summerville (up­ proposed by MORATALLA (1993), in an unpublished per tongue) near Moab, Utah; Summerville-Bluff thesis, based on a distinctive track type found at the transition zone near Blanding, Utah; Upper Sum­ Fuenteosalvo locality (FTS in TABLE I). Becausethis merville, near Bullfrog Utah, and at Cactus Park is an unpublished source, in Spanish, we herein Colorado; ?Bell Ranch Formation, northwestern translate and amend the original description and dis­ Oklahoma. cussion, so as to establish a formal description. Europe: ? Upper Jurassic-Lower Cretaceous, Description. Fuentesalvo trackway 1 consists of Oncala Group, Fuentesalvo, Spain. six, relatively well preserved, tridactyl tracks (mean foot length 21.7 cm; mean foot width 19.6 cm: see Central Asia: Upper Jurassic carbonates TABLE I). Digits robust without sharp termination . (Oxfordian-Kimmeridgian) Kugitang Suite, Khodja­ Pads not differentiated beyond slight marginal con­ Pil Ata, Turkmenistan. stri ctions. Digit III well-developed and V shaped. Therangospodus pandemicus ichnosp. nov. Heel relatively symmetrical. Step averaging 53.4 cm and stride 107.5 cm. Fig. 3-6 Etymology: "Oncalensis" meaning from the On­ Description: Medium sized, elongate asymmet­ cala Group, Spain. ric theropod track averaging 28 cm long by 20.5 cm Type locality: Fuentesalvo, north of Soria, La wide (n = 33 trackways). Digital pad impressions and Rioja Province, Spain. casts always elongate, cigar-shaped and without creases separating discrete phalangeal pads. Faint Type horizon: Huerteles "Aloformation" of indentations at the margins of pads, sometimes re­ GOMEZ FERNANDEZ (1992), Oncala Group, Came­ veal location of phalangeal pads and suggest a 2-3- ros Basin. 4 phalangeal formula corresponding to digits 11-111- Age: Possibly? Berriasian, or within the undiffer­ IV. Claw impressions preserved in some impres­ entiated Upper Jurassic (? Kimmeridgian or Titho­ sionsand casts. Trackway narrow with pace angula­ nian) to ? Lower Cretaceous (Berriasian), see tion averaging about 170°, (n=6) and trackway width TISCHER (1966), GOMEZ FERNANDEZ (1992). averaging 35 cm (n=29). Step variable, averaging 94 cm (n=36). Type specimen: MORATALLA, 1993: fig. 8.5.1-2.

347 M.G. LOCKLEY; C.A. MEYER & J.J. MORATALLA

Referred material: specimen CU-MWC 186,2 (Fig. 5-6). Such differ­ LOCKLEY & MEYER, 1999:207, fig. B.B. ences are insufficient to suggest that the pads be­ LOCKLEY, HUNT & LUCAS, 1996: 252, fig . 5 (center and left). came more clearly defined through time, though this Distribution: Currently known from type locality possibility can not be ruled out. As shown in Figure 6, at Fuentesalvo, and from localities at Bretun and in comparison with T oncalensis most tracks of T Santa Cruz de Yanguas (see MORATALLA, 1993). pandemicus appear to be elongate, without diver­ gent digits (compare with Fig. 1, and MORATALLA, DISCUSSION OF THERANGOSPODUS ONCALENS/S 1993: fig. 8.5.1). In orderto preserve historical conti­ nuity with the original unpublished work of It is difficult to compare Therangospodus on­ MORATALLA (1993), and Simultaneously do justice to calensis with T pandemicus, because the two sam­ a description of the large sample oftracks from North ples originate from different continents and are America and Asia, we have interpreted the afore­ samples of markedly different sizes. It is also mentioned morphological differences as sufficient thought that T oncalensis is a younger ichnite, to justifying different taxonomic labels at the ichnO'­ though the exact age is unknown. The type material species level. Future work may establish that the of T oncalensis consists of a trackway with short Spanish sample is more appropriately considered step and in this regard differs notably from T pande­ conspecific with the Asian and North American ma­ micus, which has an average step that is almost terial. twice as long (TABLE I). Differences in the morphol­ ogy of the actual tracks between the two ichnospe­ GENERAL DISCUSSION cies are subtle. MORATALLA (1993) reports that two digital pads can be recognized on digit II of track It is evident that Therangospodus pandemicus is number FTS-2. Such features are not observed in not only a widespread ichnospecies, but also one the large T pandemicus sample except faintly in that appears to have a definable stratigraphic and paleoecological range. The ichnospecies is cur-

70

N=l08

60 Mega/osauripus .. .. E u .. c 50 "- .. .. .c ~~ ...... 'C .. "i- 40 ...... c .. titt .. ";:- I.... Co ...... t .. -0 30 .. -:1,0 0 ..... u.. -. - :'1- I- 20 .. -II-- I-- cD - . Therangospodus -. I I 10 20 30 40 50 60 70 80 Footprint length in cm

Fig. 8 - Bivariate plot of length-width of Therangospodus tracks from North America and Asia with Mega/osauripus tracks from same localities in North America and Asia for comparison. Note non overlapping distribution. (MegaJo­ sauripus data includes 10 trackways from Portugal and Spain see text for details).

348 THERANGOSPODUS: TRACK EVIDENCE OF A JURASSIC THEROPOD WITH WELL-PADDED FEET

rently known only with certainty from the Moab couple of isolated footprins, at the type section Megatrackiste in Utah and one or two other nearby (Fuentesalvo), two trackways at Santa Cruz de Yan­ localities that represent approximately the same guas, and one trackway and other footprints from stratigraphic level in the Entrada-Summerville For­ Bretun (MORATALLA, 1993), comprising only four mations (LOCKLEY & HUNT, 1995; LOCKLEY & MICK­ complete trackways, as compared with five of the ELSON , 1997). These deposits are thought to be world's longest trackways in Asia (LOCKLEY, MEYER Oxfordian to Kimmeridgian in age. The ichnospe­ & SANTOS, 1996) and several dozen measured cies is not known from deposits that are either trackways from North America (Fig. 8, TABLE I). The younger or older in this region. The ichnospecies is age of Therangospodus onca/ensis is also some­ also only known from deposits in which Mega/o­ what uncertain as are many of the tracks and track­ sauripus occurs. Given that these two theropod bearing levels of the thick terrestrial sequences of track types co-occur at dozens of sites, it is reason­ the Cameros Basin of the la Rioja region. According able to ask if there is the possibility that the smaller to prevailing opinion (MORATALLA, 1993), the age of ichnotaxon Therangospodus might not be a juvenile the Therangospodus-bearing beds is possibly Ber­ of the larger ichnotaxon Megalosauripus. The an­ riasian, within the Upper Jurassic (Tithonian) - swer appears to be no on the basis of a number of Lower Cretaceous (Berriasian) Huerteles "Alofor­ consistent morphological differences as follows: the mation" of the Oncala Group; see TISCHER (1966), two samples are discrete and non-overlapping in GOMEZ FERNANDEZ (1992). their size ranges (Fig. 8); Therangospodus, lacks If, as the lack of age diagnostic evidence sug­ distinct pad impressions whereas Megalosauripus gests, this sequence could be anywhere from Upper does not; Therangospodus has a narrow trackway Jurassic to Lower Cretaceous in age we must ask if and relatively long step, whereas Megalosauripus further suggestions about the age of this group can has a wide and irregular trackway and a short step be made based on the ichnofauna. First we should relative to its large size (TABLE I and tables I-III in expect that if Therangospudus is not a very long LOCKLEY, MEYER & SANTOS, 1998). This latter char­ ranging ichnogenus, spanning the entire Oxfordian­ acteristic of a narrow trackway and moderately long Berriassian interval, perhaps it is confined to just a step is very consistently seen throughout available part of the Upper Jurassic, suggesting that the samples (TABLE I). Thus we conclude that in North Spanish Therangospodus might be Jurassic. If how­ America there is a distinct Megalosauripus­ ever, it were the case, that Therangospdus is older Therangospodus track assemblage. than Berriasian, one might ask why Theran­ In central Asia, specifically at the large Khodja Pil gospudus does not co-occur with Megalosauripus Ata site in eastern Turkmenistan, we find tracks in­ as it does in North America and Asia. There are sev- distinguishable from Therangospodus, along side those of Megalosauripus. Again there is a distinct size differentiation, different trackway pattern and lack of differentiated pads. We therefore consider the Asian ichnites to be attributable to Theran­ gospdus cf. pandemicus. Thus we conclude that in Central Asia there is also a distinct Megalosauripus­ Therangospodus track assemblage. We have not yet found any Upper Jurassic tracks in Europe that we consider to be unequivocal examples of the Therangospodus morphology. The only example of track of approximately the same size and age, that is ! preserved in a collection, is the specimen illustrated , .' in Figure6 H, from Silo Martinhodo Porto (Portugal). , -{ Furtherwork is required to look at a larger sample of well-preserved tracks of this type to see if they re­ veal typical Therangospodus morphology and occur , in rocks of the same age, and in conjunction with Megalosauripus. Also while this paper was in re­ view, another track attributable to Therangospodus was found in the Summerville Formation of Colo­ 10 em rado (Fig. 9).

Therangospodus oncalensis is less well-known Fig . 9 - Therangospdus from the Summerville Forma­ than Therangospodus pandemicus, having been re­ tion , Cactus Park Colorado (CU-MWC 188.63). Length ported from only one continuous trackway and a 22.5 em, width 20.2 em (compare with TABLE I).

349 M.G. LOCKLEY; C.A. MEYER & J.J. MORATALLA

eral possible answers. First, perhaps Therangospo­ indication of the Late Jurassic age of some of the dus does co-occur with Megalosauripus in Spain, strata in this region. but the co-occurrence has not been documented. A It is worth noting here the age and composition of large track from Bretun (La Mantecasa sublocality) Late Jurassic and Early Cretaceous terrestrial verte­ is 43 cm long and characterized by well defined digi­ br<3te faunas and ichnofaunas is still not well known . tal pads (MORATALLA, 1993: fig 8.6.1 C). Such pro­ LUCAS (1993) attempted to define three land verte­ nounced padding is not characteristic of many of the brate ages (LVA) for western North America, span­ large theropod tracks found in overlying Cretaceous ning the interval Kimmeridgian to mid Albian, but he sequences. A second possibility is that Theran­ recognized that there were intervals during this time, gospodus had a greater stratigraphic range than when there are gaps in the skeletal record. The Megalosauripus and so can still be found after question therefore arises, as to whether the ichno­ Megalosauripus has disappeared. Third, we might logical record fills in these gaps. Forexample his old­ consider that Megalosauripus did not range into this est LVA -the Comobluffian- is based on the Morriso]l part of Spain. fauna, of western North America, which he regards At Santa Cruz de Yanguas we find pterosaur as Kimmeridgian, Tithonian and Berriasian, possi­ tracks (cf. Pteraichnus) in beds of the same age as bly extending into the Valanginian. If this were the those containing Therangospodus (LOCKLEY et a/., case the skeletal fauna, which in it's initial stage was 1995). Although originally referred to as Berriassian the time equivalent to the Mega/osauripus-Theran­ in age, following TISCHER (1966) and GOMEZ FER­ gospodus assemblage, at least in the Kimme­ NANDEZ (1992), this ichnogenus is widely distributed ridgian, would still be represented by a similar fauna in North America in Oxfordian-Kimmeridgian beds some twenty million years later, in Late Berriasian to which also contain Therangospdus and Megalo­ Early Valanginian time. Again if the Comobluffian sauripus (LOCKLEY, HUNT & LUCAS, 1996). With one fauna remained unchanged, we might expect a cor­ Upper Cretaceous exception, all Cretaceous ptero­ responding continuity in the ichnofauna. Thus it saur tracks are much larger that those found at would be possible to predict that the Megalo­ Santa Cruz de Yanguas. While this combination of sauripus-Therangospodus ichno-assemblage evidence does not prove that Spanish Therango­ would still be found in the Lower Cretaceous. If this spodus tracks are somewhat older than previously line of reasoning is accepted, it provides a simple ex­ thought, we should at least consider this possibility planation forthe presence of Therangospodus in the in further studies of the tracks or in pursuit of other Lower Cretaceous (Berriasian) of Spain, as well as lines of evidence that might provide age dates. in Upper Jurassic deposits elsewhere. However, the lack of large ornithopod tracks in association with A final line of evidence, presented by MORATALLA Therangospodus suggests that this assemblage (1993: fig. 9.2) and MORATALLA & SANZ (1997) con­ probably did not extend far into the Cretaceous (e.g. , cerns the overall paleoecological composition of the WRIGHT et a/., 1998). ichnofauna. They noted that the Oncala Group has an ichnofauna in which Therangospodus is actually Some authors however infer that the Morrison dominant, and in which typical large ornithopod or does not extend into the Cretaceous, and so con­ iguanodontid tracks are rare. By contrast the overly­ clude that the Comobluffian fauna is restricted to the ing Urbion and Enciso Groups reveal an ichnofauna Upper Jurassic. If this is the case, then there is no in which the ichnofauna is dominated by large reason to assume that the Megalosauripus-Theran­ iguanodontid footprints and much larger theropod gospodus assemblage has such an extended stra­ tracks (referred to the ichnogenus Bueckebur­ tigraphic range, and the argument that gichnus KUHN, 1958), and where Therangospdus is Therangospodus from Spain is Jurassic - is virtually absent. The appearance of abundant strengthened. DODSON (1990: 7609-7610) dis­ iguanodontid tracks is often seen close to the cussed the longevity of dinosaur genera and con­ Jurassic-Cretaceous boundary, again indicating the cluded that "the average longevity is 1 .1 stages per significant possibility of a pre-Cretaceous age for genus, or 7.7 million years". In our experience few the Therangospodus dominated assemblages. Re­ Jurassic ichnofaunas extend temporally, or chrono­ cent studies indicate that the large Cretaceous logically, for more than a single stage, so we are in tracks from Spain are not assignable to Bueckebur­ agreement with Dodson. We might predict, there­ gichnus (LOCKLEY, in press). It has also been noted fore, that the longevity of faunas (LVAs) and ichno­ that there are virtually no narrow gauge sauropod faunas might ultimately be resolved to the scale of trackways in any well-dated Cretaceous rocks. about 7.7 million years rather than the gross scale of Thus the report of narrow gauge tracks at the Valde­ about 20 million years (LUCAS, 1993). murillo site in Spain (MORATALLA et ai, 1994: fig. 3; In conclusion, all we can say at present is that the LOCKLEY & MEYER, 1999: fig. 8.9) could be further temporal range of Therangospodus is between a minimum of about one stage and maximum of about

350 THERANGOSPODUS: TRACK EVIDENCE OF A JURASSIC THEROPOD WITH WELL-PADDED FEET

three, though evidence for the latter suggestion is Upper part of Summerville Formation, Del Monte thin at best. This margin of error is due to uncertainty mines locality, north of Bullfrog Utah. about age dating, owing to lack ofdiagnosticskeletal CU-MWC 188.1 0 - Thick slab with two tridactyl di­ remains, and does not in any way reflect inherent nosaur tracks. problems with the ichnofaunas. Indeed the ichno­ faunas may ultimately playa role in helping us date CU-MWC 188.13 - Cast of dinosaur track, Del the Spanish succession more precisely. The impor­ Monte loc. tance Megalosauripus-Therangospodus track as­ CU-MWC 188.14 - Replica of natural cast dino­ semblage has been established in Upper Jurassic saur track. (Oxfordian-Kimmeridgian) track assemblages in North America and Asia, with Megalosauripus also being present in Europe, in Portugal and probably in Upper Part of Summerville Formation, Cactus Spain (LOCKLEY, MEYER & SANTOS, 1998). It is now Park, Colorado. important to try and establish independent evidence CU-MWC 188.24 - Impression of theropod track of the age of the Therangospodus-bearing beds in (possibly Therangospodus, not mentioned in text). the Cameros Basin and determine whether Megalo­ sauripus is present. Ichnologists should also try and CU-MWC 188.63 -Impression of theropod track establish whether Therangospodus is present in the (probably Therangospodus, mentioned in text, Fig. Upper Jurassic of Portugal, where the age of track­ 9). bearing beds is well-known, so that the spatial and temporal extent of the Megalosauripus Therangos­ REFERENCES podus assemblage in Europe can be better estab­ BAIRD, D. (1957) * Triassic reptile footprint faunules from Milford, lished. New Jersey. Bull. Museum Comparative Zool. , 117: 449-520. DEMATHIEU, G. (1990) - Problems in the discrimination of tridactyl ACKNOWLEDGMENTS dinosaur footprints, exemplified by the Hettangian trackways , the Causses, France. Ichnos , 1: 97-110. Research on tracks in Utah and Arizona was sup­ DEMATHIEU , G. (1993) - Empreintes de pas de dinosaures dans Ie ported mainly by National Science Foundation Gausses (France). Zubia, 5: 229-252. grants to University of Colorado at Denver. Re­ DEMATHIEU , G. & SCIAU , J. (1995) - L'ichnofaune Hettangienne search in Central Asia has been supported by the D'archosauriens de Sauclieres, Aveyron, France. Buff. Soc. His!. Nat. Autun, 151: 5-46. National Geographic Society. We thank Iberduero, DODSON, P. (1990) - Counting dinosaurs: how many kinds were the government of la Rioja and the Friends of Arnedo there? Proc. Nat!. Acad. Sci. , 87: 7608-7612. for support offield work and research in the La Rioja ELLENBERGER, P. (1972) - Contribution a la classification des piste region of Spain, and we thank Vanda Faria dos San­ de vertebres du Trias: les types du Stormberg d'Afrique du tos and the Geological Survey of Portugal for assis­ Sud (1). Palaeovertebrata, Mem. Extr. tance with field work and access to specimens. She ELLENBERGER, P. (1974) - Contribution a la classification des pis­ and Carlos Marques da Silva are also acknowl­ tes de vertebres du Trias; les types du Stormberg d'Afrique du edged for their efforts to produce this volume. Sud, (2). Palaevertebrata, Mem. Extr., 170 pp. FARLOW, J.O. (1987) - A guide to Lower Cretaceous dinosaurfo­ otprints and tracksites of the Paluxy River Valley, Somervell APPENDIX County, Texas. Geol. Soc. Am., Field Trip Guidebook, 21 st An­ nual Meeting, Waco, Texas, 50 pp. MATERIAL IN CU-MWC TRACKS COLLECTION FARLOW, J.O. & LOCKLEY , M.G. (1993) - An Osteometric Appro­ ach to the Identification of the Makers of Early Mesozoic Tri­ Selected examples of Therangospodus tracks in dactyl Dinosaur Footprints. New Mexico Museum Nat. Hist. the joint University of Colorado at Denver -Museum Sci. Bull., 3: 123-131. of Western Colorado (CU-MWC collection). List FARLOW, J.O. & CHAPMAN, R. (1997)- The Scientificstudyofdino­ based on citations in text. Bold print indicates an saur footprints, in FARLOW, J.O. & BRETT-SURNAM, M.K. (Eds.), The complete dinosaur. Indiana University Press, original specimen. Bloomington, Indiana, pp. 519-553. Moab megatracksite area: tracks from transition GOODKNIGHT, C.G. & ERTEL, O.B. (1991) - Second day road log from Moab to Price via dinosaur trackways along salt valley zone between the Entrada Formation (Moab anticline , the San Rafael Swell , and the Cleveland-Lloyd Tongue) and Summerville Formation (upper tongue ) Quarry, in AVERETT, W.R. (Ed.), Guidebook for dinosaur quar­ near Arches National Monument, Utah. ries and tracksites tour, western Colorado and eastern Utah. Grand Junction Geol. Soc., Grand Junction, pp. 83-92. CU-MWC 186.2 - Natural cast from site G. GIERUNSK1, G. (1991) - New dinosaur ichnotaxa from the early Ju­ rassic of the Holy Cross Mountains, Poland. Paleogeog. , Pa­ CU-MWC 186.5 - Replica of five tracks ( = four leoclimato!., Paleoeco!. , 85: 137-148. steps) from section of long trackway, Site E. (Holo­ GIERUNSKI, G. & AHLBERG, A. (1994) - Late Triassic and Early Ju­ type of Therangospodus pandemicus). rassic dinosaur footprints in the Hoganas Formation of southern Sweden.lchnos, 3: 99-105.

351 M.G. LOCKLEY; C.A. MEYER & J.J. MORATALLA

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