GAIA N°15. LlSBONLISBON. DEZEMBROIDECEMBER 199B. pp. 63-74 (ISSN: OB71-5424)
GLOBAL CORRELATION OF THE TRIASSIC THEROPOD RECORD
Andrew B. HECKERT Department of Earth and Planetary Sciences, University of New Mexico. ALBUQUERQUE, NM 87131. USA
Spencer G. LUCAS New Mexico Museum of Natural History and Science. 1801 Mountain Road NW, ALBUQUERQUE, NM 87104. USA
ABSTRACT: Theropod dinosaur body fossils are known from Upper Triassic (Carnian Rhaetian) strata in North and South America, Greenland, Europe, and India. Theropod foot prints, usually assigned to the ichnogenus Grallator HITCHCOCK and related ichnotaxa, have been described from the Upper Triassic of North America, Greenland, Europe, and Africa. These theropod occurrences are readily correlated as latest Carnian, Norian, and Rhaetian records. The earliest theropods are among the first dinosaurs and appear essentially syn chronously in the Upper Triassic of the United States, the United Kingdom, Brazil, Argen tina, and India. These theropods include herrerasaurs and ceratosaurs from the lower Chinle Group in the southwestern United States, the problematic theropod Saltopus e/ ginensis HUENE from the Lossiemouth Sandstone in Scotland, the herrerasaurid Stauriko saurus pricei COLBERT from the Santa Maria Formation in Brazil, the herrerasaurid Herrerasaurus ischigualastensis REIG and the basal theropod Eoraptor lunensis SERENO, FORSTER, ROGERS & MONETTA from the Ischigualasto Formation in Argentina, and the prob able basal theropod A/walkeria maleriensis (CHATTERJEE) from the Maleri Formation in India. All of these occurrences are of late Carnian age, approximately 228 Ma . Thus, the oldest di nosaurs are not from South America, as commonly claimed, but instead appear synchro nously across Pangea in the upper Carnian fossil record of four modern continents. There are several ceratosaurs and herrerasaurs of Norian age, including the herrerasaur Chindesaurus bryansmalli LONG & MURRY, several unnamed herrerasaurs, and at least two ceratosaurs from the middle Chinle Group, USA, as well as the ceratosaurs Procompsog nathus triassicus HUENE and Liliensternus liliensterni (HUENE), and problematic theropods such as Halticosaurus HUENE from the Middle Stubensandstein, Germany. The Rhaetian fos sil record of theropods is characterized by abundant tracks, particularly ofthe ichnogenus GrallatorHITcHcOCK, in the USA, Europe, and Africa, and numerous theropods, including the Whitaker quarry theropods Coelophysis bauri COPE and Syntarsus? RAATH from the United States, the ceratosaur Liliensternus aire/ensis CUNY & GALTON and other, indeterminate forms from France, and a poorly known theropod fauna from the Los Colorados Formation of Argentina. With the notable exception of the Whitaker Quarry at Ghost Ranch, which pre serves dozens of theropod skeletons, Late Triassic theropods never dominated the tetra pod predator guild.
INTRODUCTION gle locality, we cannot discuss their actual stra tigraphic ranges in a quantifiable fashion, butwe feel In this paper, we document the stratigraphic suc that detailed documentation of the lithostratigraphic cession of Triassic theropod dinosaurs worldwide. and biostratigraphic superposition of these thero At this time, all Triassic theropods appear to be en pods provides important information regarding their demic, with the possible exception of the Rhaetian early evolution and diversification. Whitaker quarry theropod(s). However, correlations based on other tetrapods, principally aetosaurs, al Theropods are among the first dinosaurs, and low us to order chronologically the appearance of their body fossils occur in Upper Triassic sediments theropod taxa. Because most of these taxa are in North and South America, Greenland, Europe, known from relatively few specimens, often at a sin- and India, with footprints known from North America,
63 artigos/papers A.B. HECKERT & S.G. LUCAS GLOBAL CORRELA TlON OF THE TRIASSIC THEROPOD RECORD
Greenland, Europe, and Africa (Fig. 1). Theropod vertebrate faunachron (Ivf) of LUCAS & HUNT LUCAS (1991 a, 1993) and SULLIVAN et a/. (1996). A body fossils are known from rocks as old as Upper (1993a, 1993b) and is of early late Carnian age. new ceratosaur, to which at least one element of the Carnian, and occur throughout rocks of Norian and HUNT et a/. (1998) reported a single theropod podial original (COPE, 1887) type material of Coelophysis Rhaetian age. The ichnogenus Grallator HITCH from the Popo Agie Formation of Wyoming, which is can be assigned, was recentlly described as Eucoe COCK and related ichnotaxa are widely accepted as als'O considered Otischalkian based on the pres 'Iophysis baldwini (SULLIVAN & LUCAS, 1999). the footprints of theropod dinosaurs. These trace ence of the primitive phytosaur Paleorhinus (LUCAS, PADIAN (1986), LONG & MURRY (1995) and HUNT et fossils are rare in pre-Rhaetian rocks, but are locally 1993, 1994). a/. (1996) have also reported ceratosaurs from the abundant in uppermost Triassic rocks in the Chinle Numerous theropods from the lower Chinle Painted Desert Member of the Petrified Forest For Group and Newark Supergroup in North America Group co-occur with the aetosaur Stagonolepis rob mation in the PFNP. and in the lower Elliot Group in South Africa. ertsoni AGASSIZ, the phytosaur Rutiodon EMMONS The type material of the putative bird Protoavis For the purposes of this paper, we consider (sensu BALLEW, 1989), or both. Stagonolepis and texensis CHATTERJEE (CHATTERJEE, 1991, 1995) in Eoraptor lunensis SERENO, FORSTER, ROGERS & Rutiodon are index taxa of the Adamanian (latest Fig. 1 - Upper Triassic theropod localities. A - Chinle cludes bones of an aberrant, non-avian theropod MONETTA and herrerasaurids to be theropods. We Carnian) Ivf, so numerous latest Carnian dinosaurs Group, southwestern U.S.A. B - Newark Supergroup, east (LONG & MURRY, 1995; HUNT et a/., 1998). The puta are well aware of recent debate surrounding the ori are known from the lower Chinle. These include a ern U.S .A. and Canada. C - Fleming Fjord Formation, tive ornithomimosaur Shuvosaurus inexpectatus gins and definition of dinosaurs. Specifically, we herrerasaurid and a ceratosaur from the Placerias Greenland. D - Lossiemouth Sandstone, Elgin, Scotland . CHATTERJEE exhibits no synapomorphies of the Di recognize that some consider Eoraptor and herre quarry in the Bluewater Creek Formation of Arizona E - Airel layer, Carentan Basin, Normandy, and Gres a A vi nosauria (sensu GAUTHIER, 1986; BENTON, 1990; rasaurs, including Herrerasaurus ischigua/astensis (LUCAS, HUNT & LONG, 1992; LUCAS, HECKERT & cula contorta , Nancy, France. F - Keuper, Germany. SERENO, 1991,1995; SERENO&NOVAS, 1992, 1993; G - Santa Maria Formation, Brazil. H - Ischigualasto Forma REIG and Staurikosaurus pricei COLBERT, to lie out HUNT, 1997; LONG & MURRY, 1995 - HUNT et a/. NOVAS, 1993, 1996; SERENO eta/., 1993). Further, side of the Dinosauria (e.g., Ornithischia + [1998] named the ceratosaur Camposaurus ari tion, Argentina. I - lower Elliot Formation, Stormberg Gro up, southern Africa. J - Maleri Formation, India. we are not convinced that this specimen preserves Saurischia) (e.g., PADIAN & MAY, 1993; HOLTZ, zonensis), two relatively derived (non any material diagnostic of the Theropoda, and the 1994,1995; FRASER & PADIAN , 1995; HOLTZ & PA herrerasaurid) theropods from the Bluewater Creek supposed synapomorphies ofthe Ornithomimosau DIAN, 1995). However, we agree with the ongoing Formation near Fort Wingate, New Mexico (HECK ria (CHATTERJEE, 1993a) are unconvincing because work of Sereno and Novas (e.g., NOVAS , 1992, ERT , 1997), a theropod of unknown affinities (CASE , lected from the Painted Desert Member of the Petri of poor preservation. Hence, we follow LONG & 1993, 1996; SERENO, 1993, 1995; SERENO & No 1922,1927; HUENE, 1930; HUNTeta/., 1998)named fied Forest Formation in the PFNP. This moderately MURRY (1995) and consider Shuvosaurus to repre VAS, 1992, 1993; SERENO et a/., 1993) and consider Spinosuchus caseanus by HUENE (1930), an herre large (3 to 4 m long) herrerasaur is diagnosed on the sent a problematic archosaur, perhaps the aberrant Eoraptor and herrerasaurids to represent dino rasaurid (CASE, 1922, 1927, 1932a; MURRY, 1989; basis of several features of the astragalus (LONG & rauisuchian Chatterjeea elegans LONG & MURRY. saurs, specifically basal theropods, and thus include LONG & MURRY, 1995) from the Tecovas Formation MURRY, 1995: 173). Consequently, we disagree with them in our review of Triassic theropods. in Texas named Caseosaurus crosbyensis by HUNT LONG & MURRY (1995) and refer no specimens to These theropods co-occur with the aetosaur Ty et a/. (1998), and a fragmentary theropod from the this genus outside of the holotype, as the astragalus pothorax coccinarum COPE and the phytosaur WESTERN NORTH AMERICA Garita Formation in New Mexico (HUNT & LUCAS, is not preserved in any of the specimens they refer to Pseudopa/atus MEHL, both of which are index taxa 1995). Recently, HUNT et a/. (1996) reported two ad Chindesaurus, and many of these specimens are of the Revueltian IvfofLuCAS & HUNT (1993a, bland All Triassic theropod body fossils in North Amer ditional theropods from the Blue Mesa Member of not even convincingly dinosaurian. Other herre thus of early-mid-Norian age (Fig. 2). Thus, prelimi ica are derived from the Upper Triassic Chinle the Petrified Forest Formation in the Petrified Forest rasaurs from the middle Chinle Group include three nary analyses indicate the presence of at least three Group, which encompasses nonmarine deposits National Park (PFNP) in eastern Arizona. There forms from the Bull Canyon Formation described, herrerasaurs, as many as three ceratosaurs, and from Texas to Idaho and Oklahoma to Nevada (LU fore, we document numerous theropods from the but not named, by HUNT (1994) and HUNT et a/. several problematic theropods in the middle Chinle CAS , 1993). To date, numerous theropod fossils lower Chinle Group, including two herrerasaurs, a (1998). Group of early-mid-Norian age. have been reported from Triassic sediments in ceratosaur, and as many as four other theropods of Texas, New Mexico, and Arizona, with isolated CARPENTER (1997) described a large cerato Unlike the underlying rocks, strata of Apachean unknown affinities. specimens known from Wyoming. saur, Gojirasaurus quayi CARPENTER, from the Bull (Rhaetian) age in the Chinle Group contain abun Theropods from the middle Chinle Group in Ari Canyon Formation in eastern New Mexico. This dant tetrapod footprints and relatively few body fos The first theropod reported from the Chinle zona, New Mexico and Texas are generally more theropod was previously described, but not named, sils. Outside of the extremely prolific Whitaker Group was Coe/ophysis (COPE, 1887), with other fragmentary than the late Carnian dinosaurs. Herre by PARRISH & CARPENTER (1986), and also men quarry, which produces abundant ceratosaur skele notable theropod remains reported later by HUENE rasaurs include Chindesaurus bryansmalli LONG & tioned by CARPENTER & PARRISH (1985), HUNT & tons, almost all theropod fossils from the upper (1915), CASE (1922,1927, 1932a), and CAMP MURRY from the Painted Desert Member of the Petri LUCAS, (1989), LUCAS & HUNT (1989), PARRISH Chinle Group are trace fossils assigned to the ichno (1930). Even after discovery of the Whitaker quarry fied Forest Formation in the PFNP (LONG & MURRY, (1989), and LONG & MURRY (1995). This theropod is genus Grallator HITCHCOCK. in 1947 (COLBERT, 1947, 1989), other theropod sites 1995) and a number of fragmentary herrerasaurids known from fragmentary ribs, centra , a scapula, a were rare. Recently, intensive collection and study The Whitaker quarry produces abundant re from the Bull Canyon Formation in eastern New pubis, a tibia, and a metatarsal (CARPENTER, 1997). of the Chinle has greatly increased the number of mains of the ceratosaur Coelophysis (COLBERT, Mexico and West Texas (HUNT, 1994; HUNT et a/., Based on the tibia, CARPENTER (1997) estimated a known theropods from the Chinle Group. To date, 1989) and possible individuals of the genus Syntar 1998). Ceratosaurs include Gojirasaurus quayi body length of 5.5 m, which would make this one of only the Whitaker quarry theropod is named and sus (PAUL, 1993; SULLIVAN, 1994; HUNT et a/., CARPENTER from eastern New Mexico, a new taxon the largest Upper Triassic theropods. well-known. Here, we briefly review recent Chinle 1998). These are the most derived ceratosaurs and from northern New Mexico, and at least one cerato Group theropod discoveries. Ceratosaurs from the Painted Desert Member of are among the most derived Triassic theropods saur from the PFNP (PADIAN, 1986). Problematic the Petrified Forest Formation in northern New Mex known (e.g., ROWE & GAUTHIER, 1990; HOLTZ, A fragmentary centrum from the Salitral Forma theropod taxa from the middle Chinle Group include ico almost certainly represent the type material of 1994). tion of northern New Mexico reported by HUNT & the putative bird Protoavis texensis CHATTERJEE Coelophysis COPE (SULLIVAN et a/., 1996; SULLIVAN LUCAS (1990a) may represent a theropod dinosaur. and the putative ornithomimosaur Shuvosaurus in Various workers have assigned tridactyl tetrapod & LUCAS 1999). However, in 1996 the ICZN ruled in tracks in the Chinle to the ichnogenera Grallator This is the only putative theropod that co-occurs with expectatus CHATTERJEE. favor of COLBERT et a/.'s (1992) petition and estab the aetosaur Longosuchus meadei (SAWIN). Longo HITCHCOCK, Agia/opous BRANSON & MEHL, Coelu LONG & MURRY (1995) named Chindesaurus lished a neotype for Coelophysis bauri COPE from suchus is an index taxon of the Otischalkian land- rosaurichnus KUHN, and Atreipus OLSEN & BAIRD. based on a partial skeleton of an herrerasaur col- the Whitaker quarry specimens, contra HUNT & With the possible exception of Atreipus, these tracks
64 65 A.B. HECKERT & S.G. LUCAS GLOBAL CORRELA TlON OF THE TRIASSIC THEROPOD RECORD
North Europe South CUSHMAN, and Stenonyx (LULL), many of which tol Channel area and in South Wales also produce America (UK) I(C(mti rl ent,.I~ America have multiple ichnospecies. Recently WEEMS fragmentaryTriassiccoelurosaurs (FRASER, 1994). (1992) argued that of these, only Eubrontes, Gralla In Germany, the holotypes of the ceratosaur Pro tor, and Kayentapus are valid. In general, Triassic compsognathus triassicus HUENE and the problem theropod footprint assemblages are dominated by atic taxa Halticosaurus longotarsus HUENE and H. Grallator, and Jurassic assemblages by Eubrontes, orbitoangulatus HUENE were found in a marly inter with a few Kayentapus known also from the Jurassic val just above the Middle Stubensandstein (HUENE, section. 1908,1921 ,1932; SERENO & WILD, 1992), strata of Noteworthy recent additions to the ichnofossil undisputed Norian age (AIGNER & BACHMAN, 1992). record of the Newark Supergroup are the new dino Recently, SERENO & WILD (1992) dismembered the sauromorph ichnogenus and ichnospecies Banis holotype of Procompsognathus triassicus, assign §:~~ terobates boisseaui FRASER & OLSEN from the Dry ing the skUll to the sphenosuchian Sa/toposuchus ~~-;.~.~ Fork Formation in the Danville River Basin. This ich connectens and retaining the postcrania in Pro c <0 ~ ~m ~~ ci. nogenus strongly resembles Grallator and has a compsognathus. They then redescribed Procomp 'c c. ~.oS!2~ w ~ ~ w ~ , probable theropod trackmaker (FRASER & OLSEN, sognathus as a moderately derived, "Segisaurus ~ Q) 2 (l) iij.~ .'1 1996). SULLIVAN, RANDALL & HENDRICKS (1994) re like" ceratosaur (SERENO & WILD, 1992: 455). CHAT ,c:luCIl('(j !ll ~ ~i.§.9 ~ ported a new ichnofauna from the Lockatong For TERJEE (1993b) argued that the skull is actually E~E'O'e (0 c <0 ~.s ~(!)Q mation of Pennsylvania that includes Atreipus and ceratosaurian. Thus, there is at least one ceratosaur 'u c"" C <0 ~<.:> ~ Grallator(=Coelurosaurichnus). Thus, the entire Tri in the Middle Stubensandstein. --' assic theropod fossil record of the Newark Super Halticosaurus HUENE is a problematic Late Trias group consists of a variety of Grallatorand Grallator sic taxon represented by several fragmentary speci multiple like footprints from strata ranging from late Carnian c herrerasaurs, mens that Huene identified as three different Z .~ to Rhaetian in age (Fig. 2) . ..> species, H. longotarsus, H. orbitoangulatus, and H. cr: ~ f- lilienstemi HUENE (HUENE, 1908, 1932,1934) Of « GREENLAND a these, H. longotarsus and H. orbitoangulatus are JENKINS et al. (1994) first documented the Upper from the Middle Stubensandstein and H. lilienstemi Triassictetrapod fauna of Greenland in detail. So far, was found in the Knolienmergel of Germany the only theropod body fossil this assemblage con (HUENE, 1908, 1932, 1934; SERENO & WILD, 1992). Fig. 2 - Biochronology of Late Triassic theropods. tains is an indeterminate theropod consisting of "dis WELLES (1984) removed the type of H. lilienstemi associated vertebrae and ribs, a partial pelvis and from the genus and designated it the type of his new hindlimb, including a femur(length, 33 cm) and pha genus Lilienstemus lilienstemi (HUENE). ROWE & langes" (JENKINS et al., 1994: 14) from the upper GAUTHIER (1990) assigned Lilienstemus to the Portland Formation and thus are of Jurassic age Bjergkronerne beds in the 0rsted Dal Member olthe Ceratosauria, and NORMAN (1990) considered the represent theropods of varying sizes, and we agree Fleming Fjord Formation. They also reported nu remaining species nomina dubia. Both H. longotar (OLSEN, SCHLISCHE & GORE, 1989). with LEONARDI & LOCKLEY (1995) that Agialopous merous trackways assignable to Grallator sp. from sus and H. orbitoangulatus are represented by frag and Coelurosaurichnus are probably junior subjec With the relatively recent recognition that much the 0rsted Dal Member. The occurrence of Aetosau mentary material (a partial jaw and fragmentary tive synonyms of Grallator. OLSEN & BAIRD (1986) of the uppermost Newark Supergroup is of Jurassic, rus with the theropod fossils olthe 0rsted Dal Mem postcrania for H. longotarsus and a crushed skull for believe that the nature and association of manus not Triassic age, most of the famous Newark Super ber indicates an early-mid Norian age for this fauna H. orbitoangulatus) that have thus far prevented prints with tridactyl pes prints of Atreipus indicate group theropod footprint localities - are now part of (Fig. 2). workers from incorporating these taxa into a modern that the probable Atreipus trackmaker was an or the Early Jurassic ichnofauna (OLSEN, 1980; phylogenetic hypothesis of early dinosaur evolution nithischian(s). However, WEEMS (1992) believes OLSEN, MCCUNE & THOMSON, 1982; OLSEN, EUROPE (e.g., NORMAN, 1990; HOLTZ, 1994). Pending such that the manus prints are inconclusive and could SCHLISCHE & GORE, 1989; SILVESTRI & SZAJNA, studies, we retain the genus Halticosaurus as a The Triassic theropod fossil record of Europe in represent occasional manus tracks by a theropod . 1993). However, unlike the Chinle Group, theropod problematic Norian theropod. The co-occurrence of cludes body fossils from the United Kingdom, The vast majority of Chinle Group theropod foot footprints are known from nearly the entire stra these forms with numerous taxa, particularly the ae France, and Germany, with numerous footprints prints occur in rocks of, or correlative to, the Rock tigraphic section of the Newark Supergroup. Foot tosaur Aetosaurus, indicates an early-mid Norian from several localities. The Lossiemouth Sandstone Point Formation (LUCAS & HUNT, 1993a,1993b; prints of Grallator and related ichnotaxa occur in ageforthe Triassictheropods of the Keuper, with the of Elgin, Scotland, has produced theropod fossils LOCKLEY & HUNT, 1995). Therefore, the upper rocks of late Carnian to Rhaetian age throughout ceratosaur Lilienstemus liIienstemi representing assigned to Saltopus elginensis HUENE. The Lossie Chinle Group is depauperate in theropod body fos much of the Newark Supergroup and have been re the youngest Keuper Norian theropod . sils outside of the Whitaker quarry, but has a rich ported by numerous workers (e.g., BAIRD, 1957; mouth Sandstone is the type stratum of the aetosaur Stagonolepis roberisoni AGASSIZ, which is of Ada The holotype of the ceratosaur Lilienstemus ichnofossil record of theropod footprints. OLSEN & BAIRD, 1986; WEEMS, 1987, 1992; OLSEN, airelensis (LARSONNEUR & LAPPARENT, 1966; CUNY 1980; OLSEN & FLYNN , 1989; OLSEN, SCHLISCHE & man ian (latest Carnian) age. OSTROM (1981) and NORMAN (1990) reviewed the problematic thero & GALTON, 1993) was derived from the Airel layer of EASTERN NORTH AMERICA GORE, 1989) (Fig. 2). pods, and NORMAN (1990) considered Saltopus el the Carentan Basin in Normandy, France. This local The Newark Supergroup in eastern North Amer A variety of names have been assigned to tridac ginensis to be a nomen dubium. Because of its ity and the Saint-Nicolas-de-Port locality near ica yields no body fossils of Triassic theropods. The tyl tetrapod footprints in the Newark Supergroup. strong theropod affinities and latest Carnian age, we Nancy, in the "Gres a Avicula contoria", which has holotype of Podokesaurus holyokensis TALBOT and These include Grallator HITCHCOCK, Eubrontes include it in our review here as one of the oldest also produced fragmentary theropod material, are of casts of theropod bones attributed to Coelophysis HITCHCOCK, Kayentapus WELLES, Anchisauripus known theropods. The fissure-fill assemblages de Rhaetian age (LUCAS & HUBER, 2000). sp. (COLBERT & BAIRD, 1958), originally considered HITCHCOCK, Atreipus OLSEN & BAIRD, Otouphepus veloped in Carboniferous limestone around the Bris- to be Triassic in age, were probably derived from the
66 67 A.B. HECKERT & S.G. LUCAS GLOBAL CORRELA TlON OF THE TRIASSIC THEROPOD RECORD
SOUTH AMERICA nary lines of evidence indicate that it is probably of South India Africa late Norian age (COOPER, 1982; LUCAS & HUBER, land IICcmtirlental\1 America The Triassic theropods of South America are 2000), not late Carnian age, as argued by GAUFFRE very well-studied, and consist of the herrerasaurids (1993a), who mistakenly thought that traversodon Staurikosaurus pricei COLBERT from the Santa tids, present in the upper Elliot, do not occur in post lower Maria Formation of Brazil and Herrerasaurus is APACHEAN CLiFTONIAN Carnian strata. Elliot chigualastensis REIG from the Ischigualasto Forma Group tion of Argentina, which co-occurs with the basal INDIA theropod Eoraptor lunensis SERENO, FORSTER, c ROGERS & MONETTA, 1993. Ischisaurus cattoi REIG CHATTERJEE (1987, 1994) described the primi .~ is a junior subjective synonym of Herrerasaurus tive theropod Alwalkeria maleriensis (CHATTERJEE) ;;; m> (NOVAS, 1992, 1993; SERENO & NOVAS, 1992, from the lower Maleri Formation in India. NORMAN (f) 1993). Some fragmentary theropods are known (1990) considered Alwalkeria to be a problematic z z from the Los Colorados Formation in Argentina (A. possible dinosaur. For purposes ofthis analysis, we Z c « « :;;
68 69 A.B. HECKERT & S.G. LUCAS GLOBAL CORRELA TlON OF THE TRIASSIC THEROPOD RECORD
ACKNOWLEDGMENTS CASE, E.C. (1932b) - A perfectly preserved segment of the armor groups. Regardless of this inconsistency, we note saurs are the best-represented theropod group, al of a phytosaur, with associated vertebrae. Contrib. Museum here that in the Chinle Group the aetosaur Stagono though some of the remaining herrerasaurs are the A. Arcucci provided useful discussion regarding, Paleontol. Univ. Michigan, 4: 57-80. lepis occurs above strata bearing the phytosaur Pa largest known Norian theropods, approximately 5-6 and access to specimens of, theropods from the Los CHATIERJEE, S. (1987) - A new theropod dinosaur from India with leorhinus, known from marine strata of undisputed m in length (HUNT, 1994; HUNT et al., 1998). Colorados Formation. Reviews by T. R. Holtz Jr. and remarks on the Gondwana-Laurasia connection in the Late Triassic, in MCKENZIE, G.D. (Ed.), Gondwana Six: Strati late Carnian age in Germany (HUNT & LUCAS, By the ·end of the Triassic (Rhaetian), the herre B. Perez-Moreno improved the manuscript. J. Pow graphy, Sedimentology and Paleontology, Am. Geophys. 1991 b). Therefore, by the correlation we propose Union , Washington, pp. 183-189. rasaurs had apparently become extinct, and the vast ell and W. Sill provided access to aetosaur here, the co-occurrence of early theropods with Sta majority of the theropod fauna consists of moder specimens from the Ischigualasto Formation and in CHATTERJEE, S. (1991) - Cranial anatomy and relationships of a gono/epis indicates that those theropods are of late new Triassic bird from Texas. Phil. Trans. Roy. Soc. London, ately to highly derived ceratosaurs, such as Lilien formation on South American Triassic reptiles. The Carnian age. Ser. B, 332: 277-342. stemus airelensis and Coelophysis. Abundant New Mexico Museum of Natural History and Sci ence, the Petrified Forest National Park, and Na CHATTERJEE, S. (1993a) - Shuvosaurus, a new theropod. Nail. We note here that the oldest well-known thero footprint evidence indicates that small- to medium Geograph. Res. Explor., 9(3): 274-285. tional Geographic Society supported this research. pods, Staurikosaurus, Eoraptor, and Herrerasau sized theropods (1-3 m long) had become common, CHATTERJEE, S. (1 993b) - Procompsognathus from the Triassic of rus, appear at the same time as more fragmentary, although, with the exception of the Whitaker quarry, Germany is not a crocodylomorph. J. Vertebr. Paleontol. yet more derived forms in the Chinle Group and Sal theropods never dominated the Late Triassic body REFERENCES 13(3): 29A. topus from the Lossiemouth Sandstone. Accord fossil record to the extentthat prosauropods did, and AGASSIZ, L. (1844) - Monographe des poissons fossiles du vieux CHATTERJEE, S. (1994) - A/walkeria (Theropoda) and Motumeria Gres Rouge au systeme Devonien (Old Red Sandstone) des (Plesiosauria), new names for preoccupied Walkeria CHAT. ingly, the oldest known probable theropods are they remained a conspicuously rare element in Isles britanniques et de Russie. Extf. Edinburgh New Phi/os. TERJEE, 1987 and Tumeria CHATTERJEE & SMALL, 1989. J. actually those of early late Carnian (Otischalkian) prosauropod-dominated faunas. J. , 41: 1-171 . Vertebr. Paleontol. , 14: 142. age, including fragmentary remains such as those AIGNER, T. & BACHMAN, G.H. (1992) · Sequence stratigraphic fra CHATTERJEE , S. (1995) - The Triassic bird Protoavis. Archaeop published by HUNT & LUCAS (1990a) and LUCAS CONCLUSIONS mework of the German Triassic. Sediment. Geol., 80: 115- teryx, 13: 15-31. (1994) from the Chinle and, possibly, A/walkeria 135. COLBERT, E.H. (1947) - The little dinosaurs of Ghost Ranch. Nat. The oldest theropods include several fragmen from the Maleri Formation in India. These theropods BAIRD , D. (1957) - Triassic reptile footprint faunas from Milford. Hist. , 56: 392-399, 427-428. tary theropods, including the ceratosaur Campo New Jersey. Mus. Compo Zool. Bull. , 117: 447-520. co-occur with the phytosaur Paleorhinus, and are COLBERT, E.H. & BAIRD, D. (1958) - Coelurosaur bone casts from saurus arizonensis, from the lower Chinle Group, thus of late Carnian age. BALLEW. K.L. (1989) - A phylogenetic analysis of Phytosauria from the Connecticut Valley Triassic. Am. Museum Nov., 1901 : 1- Eoraptor lunensis and Herrerasaurus ischigualas the Late Triassic ofthe western United States, in LUCAS, S.G. 11 . Dawn of the age of dinosaurs in the Ame & HUNT, A.P. (Eds.), COLBERT, E.H. (1964) - The Triassicdinosaurgenera Podokesau Refining the biostratigraphy of Triassic thero tensis from the Ischigualasto Formation in Argen rican southwest, New Mexico Museum Nat. Hist., Albuquer rusand Coelophysis. Am. Museum Nov., 2168: 1-12. pods enables us to make several observations tina, Staurikosaurus pricei from the Santa Maria que, pp. 309-339. about the nature of the original dinosaur diversifica Formation in Brazil , the problematic, fragmentary, COLBERT, E.H. (1970) - A saurischian dinosaur from the Triassic BARBARENA, M.C.; ARAUJO, D.C. & LAVINA, E.L. (1985) - Late Per of Brazil. Am. Museum Nov., 2405: 1-39. tion. In addition to the oldest theropods reviewed Saltopus elginensis from the Lossiemouth Sand mian and Triassic tetrapods of southern Brazil. Natl. Ge here, several ornithischians and prosauropods are stone in Scotland, and possibly, Alwalkeria maleri ograph. Res., 1: 5-20. COLBERT, E.H. (1989) - The Triassic dinosaur Coelophysis. Mu seum Northern Arizona Bull., 57: 1-160. known from the same or correlative strata. These in ensis from the lower Maleri Formation of India. BENTON, M.J. (1990) - Origin and interrelationships of dinosaurs, in WEISHAMPEL, D.B.; DODSON, P. & OSM6LSKA, H. (Eds.), COLBERT, E.H. ; CHARIG, A.J. ; DODSON, P.; GILLETTE, D.O .; os· clude the ornithischians Pisanosaurus from the Is These dinosaurs have a nearly synchronous firstap The Dinosauria, Univ. California Press, Berkeley, pp. 11 -30. TROM, J.H. & WEISHAMPEL, D. (1992) - Coelurus bauri COPE chigualasto Formation in Argentina (CASAMIQUELA, pearance during the late Carnian and are among the 1887 (currently Coe/ophysis bauri; Reptilia , Saurischia): pro BONAPARTE, J.F. (1978) - EI Mesozoico de America del Sur y sus posed replacement of the lectotype by a neotype. Bull. Zoo/. 1967), Pekinosaurus from the Pekin Formation in oldest known dinosaurs. By Norian time, the thero tetrapodos . Opera Ulloana, 26: 1-596. North Carolina (HUNT & LUCAS , 1994) and the pro pod dinosaur fauna was dominated by ceratosaurs Nomenclature, 49: 276-279. BRINKMAN , D.B. & SUES, H.D. (1987) - A staurikosaurid dinosaur COOPER, M.R (1982) - A mid-Permian to earliest Jurassic tetra sauropod Azendohsaurusfrom the Argana series in at the expense of herre- rasaurs. The Norian thero from the Upper Triassic Ischigualasto Formation of Argentina pod biostratigraphy and its significance . Arnoldia Zimbabwe, Morocco (DUTUIT, 1972; GAUFFRE, 1993b). Conse pod fossil record includes the herrerasaur Chinde and the relationships of the Staurikosauridae. Palaeontology, 30: 493-503. 9: 77-104. quently, we recognize that dinosaurs comprise a saurus bryansmalli and other herrerasaurs and COPE, E.D . (1887) - The dinosaurian genus Coelurus. Am. Natu CAMP, C.L. (1930) - A study of the phytosaurs with description of very small, yet diverse, component of late Carnian ceratosaurs from the middle Chinle Group, the cera ralist, 21 : 367-369. new material from western North America. Mem. Univ. Cali tetrapod faunas. These first dinosaurs had already tosaurs Procompsognathus triassicus and Lilien fornia, 10: 1-170. CUNY, G. & GALTON , P.M. (1993) - Revision of the Airel theropod dinosaur from the Triassic-Jurassic boundary (Normandy , diversified and included representatives of the or stemus lilienstemi and the problemat,c theropod CARPENTER, K. (1997) - A giant coelophysoid (Ceratosauria) the nithischian, prosauropod, and theropod clades. Fur France). Neues Jahrb. Geol. Palaontol., Abhandf., 187(3): Halticosaurus from units of Norian age in Germany. ropod from the Upper Triassic of New Mexico, U.S.A. Neues 261-288. thermore, by the latest Carnian the theropods had Of these, Lilienstemus appears to be the youngest, Jahrb. Geol. PaIBont., Abhandl., 205: 189-206. DUTUIT, J.M . (1972) - Decouverte d'un dinosaure ornithischien already diverged from other dinosaurs and included making its first appearance in the Knollenmergel, CARPENTER, K. & PARRISH , J.M. (1985)- Late Triassicvertebrates dans Ie Trias Superieur de l'Atlas occidental marocain. numerous basal forms (Eoraptor and many of the whereas the other Norian theropods in Germany are from Revuelto Creek, Quay County, New Mexico. New Mexi Compl. Rend. Acad. Sci. Paris, D, 275: 2841-2844. co Geol. Sci. Guidebook, 36: 197-198. problematic taxa), herrerasaurids, and, rarely, cera found in the Middle Stubensandstein. Rhaetian and ELLENBERGER, P. (1970) - Les niveaux paleontologiques de pre CASAMIQUELA, RM. (1960) - Noticia preliminar sobre dos nuevos miere apparition des mammiferes primordiaux en Afrique du tosaurs, as well as several problematic, but appar probable Rhaetian theropods include Coe/ophysis estagonolepoideos Argen tinos . Ameghiniana, 2: 3-9. ently derived, forms. This divergence suggests that bauri from the upper Chinle Group, Lilienstemus Sud et leur ichnologies: Esablissement de zones stratigraphi CASAMIQUELA, R.M. (196 1) - Dos nuevos estagonolepoideos Ar ques detaillees dans Ie Stormberg du Lesotho, (Afrique du tetanurine theropods should also be present at this airelensis from the Triassic of Normandy, a poorly gentinos. Rev. Asoc. Geol. Arg., 16: 143-203. Sud) (Trias Superieur a Jurassique), in HAUGHTON , S.H. time, although none have been identified from Trias known theropod from the Gres a Avicula contorta in (Ed.), I.U.G.S., 2nd Symposium on Gondwana Stratigraphy CASAMIQUELA, RM. (1967) - Un nuevo dinosaurio ornitisquio and Paleontology, Council for Scientific and Industrial Re sic strata at this time. Almost all of these taxa were France, and poorly known theropods from the Los Triasico (Pisanosaurus mertii; Ornithopoda) de la Formaci6n search, Pretoria, pp. 343-370. small, usually considerably less than 2 m long, with Colorados Formation in Argentina. All known Rhae Ischigualasto, Argentina. Ameghiniana, 4: 47-64. ELLENBERGER, P. (1972) - Contribution a la classification des pis CASE , E.C. (1922) - New reptiles and stegocephalians from the the exception of Azendohsaurus and some herre tian theropods are at least of ceratosaur-grade, and tes de vertebres du Trias: Les types du Stormberg d' Afrique Upper Triassic of western Texas. Carnegie Inst. Wash. Pub!., rasaurs, wh ich may have reached lengths of 3 - 4 m. the herrerasaurids had apparently gone extinct by du Sud (I). Palaeovertebrata, Mem. Extraord., 152 pp. 321 : 1-84. the Rhaetian. The most significant implications of By Norian time, prosauropod dinosaurs domi ELLENBERGER, P. (1974) - Contributions a la classification des pis the ichnofossil record of early theropods is the docu CASE, E.C. (1927) - The vertebral column of Coelophysis COPE. tes de vertebres du Trias : Les types du Stormberg d' Afrique nated the more terrestrial, dry ecosystems, as evi Contrib. Museum Pa/eontof Univ. Michigan, 2: 209-222. mentation of their increasing abundance, particu du Sud (II eme Patrie Le Stormberg Superieur. Le biome de la denced by the abundant specimens of pro CASE, E.C . (1932a) - On the caudal region of Coelophysis sp. and zone B/1 ou niveau de Moyeni: ses biocenoses). Palaeover larly at the end of the Triassic and into the Early sauropods known from the Norian portion of the on some new or little known forms from the Upper Triassic of tebrata, Mem. Extraord., 141 pp. Jurassic. western Texas. Contrib. Museum Paleontol. Univ. Michigan, Keuper (HUNT, 1991). The theropods, however, re FRASER , N.C. (1994) - Assemblages of small tetrapods from Bri 4:81-92. tish Late Triassic fissure deposits , in FRASER, N.C. & SUES, mained a minor component of all faunas. Cerato-
70 71 A.B. HECKERT & S.G. LUCAS GLOBAL CORRELA TION OF THE TRIASSIC THEROPOD RECORD
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