Reu. Mus. Argentino Cienc. Nut., n.s. 611): 67-103, 2004 Buenos Acres ISSN 1514.5158
Cretaceous theropods from India: A review of specimens described by Huene and Matley (1933)
Fernando E. NOVAS1," Federico L. AGNOLIN 1 and Saswati BANDYOPADHYAYZ
'Museo Argentino dr? Ciencias Naturales -B. Rivadavia-, Av. A. Gdlardo 470, (1405) Huenos Aires, Argentina. [email protected] 2Geological Studios Unit, Indian Statistical Institute, 203 Barrackporc Trunk Road, Kolkata 700 035, India. [email protected] "esearcher of CONICET.
Abstraot: The Lato Cretaceous (Maas~richtian)Lameta Ihrmation of central India has yielded dissociated elements ofavariety ofpredatory dinosaurs, most ofthem comingfrom aquarry named the e~Carnosaurbed. The materials were described hv Huene and Matlev nearly. 70 "years ago.- They recomized- nine iheronod. species, . which they sorted out into the theropod subgroups 4!arnosaiirian and eaCoelurasauria~~,I-iuene and Matley also described a considerable amount of theropod hindlimb bones (e.g.,femora, tibiae, metat,arsals, and pedal phalanges) that they oilld not refer to any of these species, hut vaguely interpreted as corresponding to ~~allosaurid~~or ,~coelurosaurid,~ theropods. We reviewed the available collection of Cretaceous therapods from Bara Simla housed at the Geological Survey of India. Kolkata. avrivlne- to the foliowine- conclusions: 1) Indosuchus and Indosaurus are abelisaurids, as recognized by previous authors, hut avsilable information is not enough to judge whether they axire synonyms; 2) Lmt~oisuchusindicus is a small abclisauroid, related to Noasaurus and Masiahasaurus on the basis or their pecu- liar cervical vortcbrae: 3: the coiitroversial taxa *Clorn~sosuc~ius-.~iDruolosauroidesn, ,~Ornithornirnoides*. and also exhihit ahelisauroid characters. and bones of iaree size are tentatively referrod to as corresnondine- to h~dosi~chus or Indosai~~zs,whereas some pedal bones of smaller size may belong to i,aeoisuckus; 5) two kinds of abelisaurid feet arcapparent:. . ansin which the nhaianpes- ofdieit- 111 andIVarerobust, and another typein~. which the iiliaianze-es of digit IV are transversely narrow and dorsoventrally deep. This reviow demonstrates that all of the theropod eloments discovered at the ,,Carnosaur bed,) belong to a single theropod clade, tho Abelisauroidev.
Key words: Theropods, Abelisaurids, Lameta Formation, Cretaceous, India.
The only available comprehensive contributiou corresponding to different positions in the verte- on Cretaceous theropod dinosaurs from India is bral column. that of Frederich von Huene and Charles Matley These authors sorted out the Indian theropods published in 1933. Their descriptions were based into Carnosauria and Coelurosauria, the main two on a large, but mixed assemblage of isolated bones lineages Huene had previously recognized within collected from a sixiglc fossiiiferous spot (the predatory dinosaurs (e.g., Huene 1914, 1920). ~Carnosaurbed.) within the Late Cretaceous Indosuchus and Indosaurus were considered to Lameta Group, extcnsivelji exposed in the be members of the carnosaurian family Provinces of Madhya Pradesh, Grujat, and Allosauridae mainly on the basis of their large size Maharashtra in NW Iridia (Fig. 1; Matley, 1921; and primitive features resembling the Jurassic Chatterjee, 1978; Chatterjee & Rudra, 1996). Allosaurus, while the remaining seven species Within this multispecies bone association, IIuene were gathered within the Coelurosauria on the and Matley (1933) recognized the following nine basis of their smaller size and more slender species of Cretaceous predatory dinosaurs: proportioxis. Compsosuchus was related to the Indosuchus raptorius and Indosaurus matleyi, both Jurassic Compsoguathus, whereas Laeuisuchus, coined on the basis of incomplete basicrania, and Jubbulpuria, Coeluroides, and Dryptosauroides Compsosuchus solus, Laeuisuchus indicus, were assembled within the c.Coeluridaem, a group Jubbulpuriu tenuis, Coeluroides largus, that in Huene's concept also included, among Dryptosauroides grandis, Ornitholnimoides others, the Jurassic Ornitholestes hermanni and mobilis, and Ornitho~nimoides(?) barasimlensis, Coelurus agilis (see for example, Huene, 1956). coined on the basis of isolated vertebrae In addition, Ornithomimoides mobilis and 68 Reuista del Museo Argentino de Ciencias Naturales, n. s. 6 flj, 2004
5a0 km
Fig. 1. Map of India showing the Late Cretaceous (Maastrichtian) fossil site at Jabalpur.
Ornithomimoides (?) barasimlensis were referred Xenotarsosaurus bonapartei, Ilokelesia by Huene and Matley (1933) and later by Huene aguadagrandensis, Majungatholus atopus, (1956) to the Ornithomimidae. But, most of the Aucasausus garridoi; Martinez et a1.1987, bones collected from the < Fig. 2. Basicrania of different abelisaurid theropods, in lateral (A-CJ, and dorsal (D-H) views. A, D, Majungatholus atopus (from Sampson et al., 1998); B, E, Zndosaurus matleyi E271565) (Dam Huene & Matley, 1933); C, F, Indosuchus raptorius (K 271685) (from Huene & Matley, 1933); G, Indosuchus raptorius (11201350) (from Iiuene & Matley, 1933); and H, Abelisaurus comahuensis (modified, from Bonaparte & Novas, 1985). Not to scale. Abbreviations: hoc hasioccipital; fr, frontal; frh, frontal horn; lac, lacrimal; prf, prefrontal; sc, sagital crest; soc, supraoccipital. K201350 the suture between frontal and lacrimal comahuensis and Majungatholus atopus (mislabeled as eprefrontalx in the original (Sampson et al., 19981, except for Carnolaurus description) closely matches that of Abelisaurus (MACN-CH 894) in which the lacrimals are comahuensis, especially in the subquadrangular transversely narrow. The narrow parietal crest of outline of the suture and the sharp < 1964).Abelisaurid traits are present in the sagital constitute autapomorphic features of indo.~uclzus parietal crest of specimen GSI K271685: in dorsal raptorius. "iew, the rostral half of this crest has a cup-shaped Indosaurus matleyi. This species was contour, beirig transversely narrow towards the founded by Huenc on the basis of a single piece of rear (Fig. 2F). This condition is only documented skull, catalogued with the number GSI K2'7/565. in abelisaurids among Theropoda (e.g., Chatterjee (1978) later declared this specimen to Carnotaurus sastrei, Abelisaurus comahiiensis, be the holotypf of indosaurus matleyi. The Majungatholus atopus). Notable for specimen GSI specimen consists oftlle posterior part of the skull, m71685 is the presence of an inter-orbital wall the dorsal surface of which is partially damaged (presumably made up by the parasphenoid bone), and includes tho right frontal bone, the temporal which is vertically hanging below the mid-frontal region, and the area for articulatiori with the suture (Fig. 3C). Such inter-orbital wall, also seen postorbital (Fig. 213, El. In dorsal view the frontal in specimen GSI KZ71565 of Indosaurus matleyi is suhtrianplar, with an ar~terolateralnotch for (Figs. 2B, 3B) is almost identical to that of articulation with the lacrimal. Tile dorsal surface ahelisaurids Aklisaurzis comaizuens~s(Bonaparte is slightly rugose, although not to the degree seen & Novas, 1985), Carnolaurus sastrei (MACN-CII in Abelisuurus (Bonaparte & Novas, 1985). In Pa- 894), and Majz~ngatholusalopus (Sampson et al., tcral view (Fig. 2B) the articular surface for the 1998). Moreover, in GSi K271685 the cranial half postorbital and lacrimal bones is rugose, being of the parasphenoid ends in a diamond-shaped dorsoventrally deep in the postorbital portion struclure (eventuaily the orbitosphonoid; Currie (roaching 5 cm thick), and becoming shallower & Zhao, 19Y3), which bears a double-foramen for rostrally (nearly 3 cm thick). The posterior surface the exit of thc olEactory nerve. Ossified of the frontal is high, except for the surface parasphenoids tightly fused to tho skull roof and hounding the suprate~nporalfbssa, w-hich is with a double exit for nerve I, are features not excavated. Both dorsai and posterior surfjces of exclusi\,e for Abelisauridae, since they are present the frontal are separated by a sharp border AIL also in Ceratosaurus (Madsen & !Are1les, 20001, interorbital wall, vertically hanging below tho Acrocanthosaurus (Stovall & bangston, 1950),and mid-frontal suture and presumably made up by some tymnnosaurids (Russell, 1970). Albeit such tho parasphenoid, is a character that Inilosuurus conditions for the parasphenoid and orbitosphenoid matle.yi (GSI K271565) shares with Indosuchus may be not syriapomorphic ibr Abelisauridae, at raptorius (GSI K271685) and other ahelisaurids least their presence in the Indian basicrania is (see above; Fig. 3). congruent with other ahelisaurid features. Bonaparte and Novas (1985) found similarities Sonie differences among the Indian basicrania between slbelisauriis comah.uensis and Indosaurus and other ahelisaurid taxa are discernable on the matlqi based on the broad interorbital region, and basis of the figures given by Huene and Matley Molnar (1990)noted that thisIndian taxon resembles (1933).For example, in specimen GSI K201350 the Carnotaurus sastrei in the massive frontals and fronto-nasal suture appears to be rostrally placed supraoccipital and markedly elevated sagittal crests with respect to thelacrimais (Fig. 2G), in contrast oftheparietals. Later, Bonaparte (1991b)pointedout to t,he remaining abelisaurids in which such suture that the supratemporal openings of Indosau,ras are is more caudally placed, approximately at level of anteropos&riorly short, resembling~belisaurusand the rostrolateral notch of the frontals for C~rmtaurm.Theabovementioncdauthorsobviouslv articulation with the lacrimals. GSI K201350 also concluded that Indosaurus is a member of exhibits on its caudal half a niedian suture Abelisauridae, an interpretation also followed by between both frontals, as well as a clear fronto- Cllatterjee and Rudra (1996). parietal suture. The presence of visible dorsal Indosuchus and Indosaurus shows some sutures in GSI K201350 is in agreement with the distinctions with respect to other abelisaurids. Tho lack of fusion wit11 the parasphenoidal bone, thus Indian taxa lack, at least, the prominent central exposing the ventral furrow for the olfactory ca- dome on frontal bones autapomorphic of nal (Huene & Matley, 1933). Both frontals and Majungatholus atopus, or the paired frontal horns parasphenoid are completely fused in Abelisaurus that characterize the Patagonian Carnotaurus. In co~izahuensisand Carnotaurus sastrei, as well as this regard, the morphology of the skull roof of in specimens GSI K27168.5 ofIndosuchus raptorius I~zdosuchusand Indosaurus is more conservative, and GSI K271565 of Indosaurz~smatleyi. and looks similar to Abelisaurus in being Presumably the lack of ossification among the dorsoventrally thick but without prominences above skull roof and braincase bones may be duo to the skuli roof. 72 Reuista del Museo Argentzno de Ciencins Naturales, n. s. G (I), 2004 A K201350 B K271565 C K271685 Fig. 3. Ventral view of basicrania of A, indosuchus raptorius (K201350), B, indosaz~r-usmatleyi (K27/ 5651, and C, Indosuchus raptorius (K 271685). E'igmes taken from Huene and Matley 11933). Not to scale. Abbreviations: fr, frontai; iow, interorbital wall. askwhether they arevalidspecies. Since Huene's specimen, and thus the validity of this fcature is description, many authors have accepted the here dismissed; preservation of the braincases anatomical distinctions between indosuchus and does not prove the presence of horn-like hdosaurus, supporting them as valid taxa. tuberosities in indosaurus, nor a dorsally smooth Moreover, they were interpreted as belonging to postorbital in indosuchus. Other possible quite different theropod clades: while Indosuchus distinctions recognized by previous authors was considered as a tyrannosaurid, Indosanrus between li~dosuchusand ir~dosaur-usconcerning was interpreted as representative of a lineage that the thickness of the skull roof, the anteroposterior inherited primitive features from Jurassic forms extension of supratemporal fbssa, the lusion of such as ~~megalosaurs~~(Chatterjee, 1978; Walker, sutures, and the degree of development of 1964). Huene and Matley (1933), and later rugosities on the skull bones, may reflect indivi- Chatterjee (1978) and Chatterjee and Rudra dual variations. In thoseregards, the frontal dome (19961, offered a list of anatomical distinctions ofMajungatholus shows a variety ofshapes, from between the hasicrania of both taxa. including being inflated in some soecimens (Sues & Taauet.. , differences in the transverse width of (he parietal 19797, to slightly dcveioped in others (Sampson sagital crest, the presence or absence of a etal., 1998).Such development of the frontaldome "transverse crest" on the dorsal surface of the also affects the width and shape of the sagital skull, the dorsoventrai thickness of the frontals, parietal crest ofMajungatholus. This possihle case and the contour of the supratemporal fossa. of individual variation in the Malagasy ahelisaur However, it is difficult to evaluate such serves as an alert when distinctions between the distinctions, not only because most of the poorly preserved skulls of indosuchus and basicrania were unavailable tbr t,hepresent study, Ii~dosaurusare evaluated. In sum, anatomical but also because the preservation of the skulls is distinctions between Indosuchus andindosaurus far from optimal. For example, in the available arc doubtful, at least. specimen ofindosaurus (GSI K271565) the dorsal surface of the braincase is eroded, thus no features 11. Skull bones originally described as of the frontal bones or sagital parietal crest are < AMNII 1'753 a shallow but distinct notch is a strong medial premaxillary process, which is identified on the posterior border of the premaxilla, located high on the medial aspect of the bone, corresponding with the subnarial foramen. The constituting another abelisaurid character. The narial fossa is deep and well delimited, in cont,rast lateral surface of the maxilla exhibits strong with Carnotaurus sastrei and Abelisaurus decoration that include foramina and grooves. The comahuensis. The external surface of the right grooves, which are predominantly oriented premaxilla (K201619) is not decorated with the dorsoventrally, split and join in a compiex pattern foramina and tuberosities present in the remaining as occurs in other abelisaurids. The grooves are abeiisaurids, and the ascending ramus looks more moro marked on the ascendingramus than in other robust and complex than in AMNH 1753. regio~lsof the maxilla, specially in larger specimens Maxilla. Chatterjee (1978, following Huene (e.g., GSI K271538). The maxillary ascending and Matley (19331, referred a fairly complete ramus is almost verticaily oriented, with the maxilla (K271548) to Indos~~chus,on the basis of rostra1 margin slightly convex in lateral view. The its considerable thickness. Moreover, Chatterjee caudal margin of the ascending ramus is (1978) took this bone as a -Rosetta stone>>, transversely wide and deeply excavated, and allowing further reference of a left maxilla provided with a presumed promaxillary fenestra (AMNH 1955; Fig. 5B) to that species. Because (hidden in side view). A maxillary fencstra is no means exist to compare with Indosaurus lacking. The dorsal margin of the maxilla is malleyi (for which no maxiiiary bone has been transversely convex and affected by deep, preserved or identified), we follow Lammana et presumably pneumatic, excavations. 011 the al. (2002) in considering specimen AMNH I955 internal side is seen a roui of dental foramina as belonging to Abelisauridae gen. et sp. indet. along the contact between the dental plates and The following description of the abelisaurid the remainder of the medial surface of the maxilla. maxillae from India is based on observations made Two conspicuous abelisaurid synapomorphies on AMNW 1955, AMNH 1753, GSI K271538, and are identified on the available Indian maxillae: a GSI K271544 (it must be noted that at the minute antorbital fossa, and dental plates collections of the GSI, specimen GSI K271538 is dorsoventrally deep, strongly fused, and decorated mistakenly labeled as GSI K271548, while specimen by obliquely oriented striations. GSI K271544, not illustrated by Huene and Matley, MNH 19S5 was referred to the subfamily is incorrectly labeled as GSI K271538; specimen Carnotaurinae by Lamanna et al. (2002),because GSI K271548 is missing). The maxilla GSI K271 it shares a promaxillary fenestra obscured by the 538 (Fig. 5) is characteristically triangular, lamina iateralis of the ascending ramus. anteroposteriorly short, and bas a proportionally Additionally, these authors suggested that an low ascending process, characters also present in anteroposteriorly short lnaxillary body with AEielisaurus, Carnotauras andMajurigatholus. The parallel dorsal and ventral margins is articulation with the pre~naxillais made through synapomorphic of this subfamily. EIowever, the Novas et al Cretaceous theropods from Indza Fig. 6. Leftjugals of Aheiisauridae in lateral (A,B) and medial (C,D) views. A, C, Majungatholus atopus (FMNH PR 2100); B,D, Ahelisauridae indet. (K271577). diagnostic value of such characters is debatable, referred to the presumed postorbital (K271580) because the maxilla is unknown in other as Indosuchus. GSI K271580 is here reidentified ahelisauroids (e.g. Ilokelesia) or it is incompletely as a portion of a right jugal (Fig. 7). preserved in others (e.g. AbeZisaunls). In other Specimen GSI N271580 has a triangular aspect words, such features may exhihit a wider in lateral view, with a slender and rod-like dorsal distribution among abelisaurids. extremity. The ventral half of the bone is Several isolated teeth were recovered from the transversely narrow, and exhibits a concave late- quarry. However, they are lost in the GSI ral surface. Towards the dorsal end the hone collection, and the figures given by Huene (pl. becomes transversely thicker, constituting the XIII, figl-10) are not detailed enough. However, most laterally projected portion of the jugal. Its many of these dental pieces agree in general shape lateral surface is decorated by grooves, being with the teeth of other ahelisaurids (e.g., intensely sculptured on the posterior andventral Abelisaurus, Mqjungatholus) in the great orbital portions. A distinct oblique groove is transverse compression and degree of backward present on its lateral surface. The medial surface curvature. of the lacrimal is smooth, with a caudal depression Jugal. The following description of this hone surrounding the infratemporal opening. The is hased on specimens GSI K271577 and GSI K271 medial surface of the ascending process ofjugal 680. We could not access another two specimens forms a longitudinal prominence (Fig. 7). (K271635 and GSI K271681) that were descrihed Jugals GSI K271580 differs from Carnotaurus, (but not illustrated) by Huene and Matiey (1933) Abelisaurus and A4ajungatholus mainly in the as portions of right and left jugals. Specimen GSI presence of a deep, rounded notch on the caudal K271577 was originally descrihed as a right lacri- margin of the ascending ramus. Specimen GSI mal, but it matches well with theascendingramus K271577 exhibits rugosities with a different of the left jugal of Carnotaurus and pattern than these seen in Carnotaurus and Majungatholus (Fig. 6). Besides, specimen GSI Majungatholus. K271580 (Fig. 71, originally was interpreted by .~~Lacrimaln.Specimen GSI K271708 was Huene and Matley as a right postorbital, interpreted hy Huene and Matley (1933 pl. XI, interpretation accepted hy Chatterjee (1978) who fig, 5) and later by Chatterjee (1978) as 76 Revzsta del Museo Argent~node Czenczas Naturales, n, s. 6 flj, 2004 Fig. 7. Right jugal of Abelisauridae in lateral (A,B) and medial (C) views. A,C, Abelisauridae indet. (K271580); B, Carnotaurus sastrei (from Bonaparte et al., 1990). Not to scale. Abbreviations: asp, ascending process; ito, infratemporal opening. corresponding to the upper portion of a right la- ornamentations made up by loramina, grooves crimal. However, the bone lacks the pattern of and prominences, resembling those of rugosities and the wide contact for the postorbital Carnotaurus and LMajungatholus.As in the latter as seen in the lacrimal of Carnotaurus and two taxa, a clear separation exists between the Majungatholus. We are unable to identify strongly decorated ventral half relative to the specimen GSI K271708. smooth dorsal (or -labial,,) halfof the dentary The Quadrate.This bone was originally described line defined by these two surfaces describes a as a right astragalus (Huene & Matley, 1933, dorsally concave curvature, which in Carnotaurus pl.XIX, fig.1; GSI K27/684), but it corresponds in and Majungatholus is lined by a number of large fact to a left quadrate. The specimen preserves foramina. The abovementioned resemblances the distal articular condyles, the base of the clearly support the hypothesis that all of the pterigoid ramus, and a rugose lateral surface for tberopod deutaries recovered in the "Carnosaur the attachment of quadratojugal. The anterior bed" belong to Abelisauridae. facet of the distal condyles is nearly flat, as it Huene and Matley (1933:50) and later occurs in the abelisaurids Ilokelesiu, Chatterjee (1978) cited some distinctions among Majungatholus and Carnotaurus (see Wilson et these dentaries, but because of the fragmentary al. 2003, character 53). nature of the material, plus the impossibility of Dentary bones. The following specimens comparing them directly, we prefer do not address were studied: GSI K271550, GSI K271709, GSI this aspect. K271529 (incorrectly catalogued as GSI K271527 Surangular. Specimen GSI K271693 was in the GSI collections, a number corresponding originally described as a left articular (Huene & to a left articular), and AMNH 1960 (a number Matley, 1933, pi. XII, fig. 31, but it is identified that also applies to a caudal vertebrae). Huene here as a left surangular The dorsal surhce of and Matley (1933) listed, althou~bdid not des- this bone is almost flat and transversely wide, ,z . 550 (Figs. 8, 9), GSI K271709 (Fig. 91, and GSI portion of the suranylar, there are two large K271529 exhibit on the ventral half of their foramina (nearly 5 mm in diameter), separated external surface a distinct pattern of each other by nearly 20 mm. Both foramina Novas et al.: Cretaceous theropods from Indza Fig. 8. Right denlary of Ahelisauridae indet. (K271550) in A, lateral, B, medial, and C, dorsal views Abbreviations: idp, interdental plates; s, symphysis. Fig. 9. Dentaries of ahelisaurid theropods in left lateral view. A, Abelisauridae indet. (K271709) (from Huene & Matley, 1933); B, Abelisauridae indet. (N271550); C, Majungatholus atopus (from Sampson et al. 1998); and D, Carnotaurw sastrei (from Bonaparte et al., 1990). Not to scale. 78 Reuista del Museo Argentzno de C7.q ?nczus Naturales, n. s. 6 [li,2004 continue forward and inward, perforating the anterior surface of the articular bone: The surangular resembles that of Carnotauras sastrei (Bonaparte et al., 1990) in the presence and position of the pair of fbramina near the glenoid cpvity. 111. Theropod tsxa based on vertebral elements Seven theropod species were coined by Huene and Matley on the sole basis of vertebrae: Compsos~~chussolus, Laeuisz~chusindieus, Jubbalpu~ia tenuis, Coeluroides largus, Dryplosauroides grandis, Ornithomimoides mobilis, and Ornithornimoides (?) barasimlensis. Also, some isolated vertebrae were described as corresponding to "ailosauroids" or "coelurosaurs". These specimens are reviewed as follows: Compsosuchus solus. This taxon was described on the basis of a single axis with fused atlantal intercentrurn (GSI K271578; Fig. 10). Most of this vertebra is preserved, except for the the upper portion of its neural arch, which is broken. The axial centrum bears one large pleurocoel, and a pneumatic opening postero- ventrally to the diapophysis. The anterior articu- lar surface of the intercentrum is slightly convex and kidney-shaped, while the posterior one is slightly concave. The diapophyses are small and blunt. A sharp lamina extends obliquely from the diapophysis to the postzygapophysis. The neural arch is wide and low. Molnnr et al. (1990) found that the axis of Contpsosuchus resemhies ttiat ofAllosaurus in the similar position of the upper pleurocoel, the cylindrical aspect of she axial intercentrum in ventral view, the axial pieurocentrnm loss than twice the length of the axial intercentrum, find the broad condition of the neural canal. This lead Molnar et al. (1990) to include Compsosuchus within Allosauridae. However, GSI K271578 exhibits the following resemblances with Carnotaurus: presence of a pneumatic pore posteroventrally to the diapophysis, and at least one large pleurocoel on the axial centrum, proportionally small and rod-like diapophyses, presence of a sharp lamina extending obliquely Fig. 10. Compsosuchus solus (K271578), axis in A, from the diapophysis to the postzygapophysis, and left lateral, B, dorsal, and C, anterior views. a neural arch laterally expanded and triangular- Abbreviations: dp, diapophysis; ic, intercentrum; shapedin dorsal view. s he axis that served as basis nc, neural canal; op, odontoid process; poz, to create Comososuchus closelv resembles that of postzigapophysis. IS1 R9111, referred to Inclosaurus by Chatterjee and Rudra (1996). In sum, the generalmorphology of this cervical vertebra indicates that it pertains (IS1 R 9111) and no evident autapomorphies are to an ahelisaurid theropod. Since there are no recognized, we conclude that Compsosuchus is a substantial differences with the axis of indosuchus nornen dubium. Novas e2 al.: Cretaceous theropods from Ind~a Fig. 11.1,aeuisuchus indicus (K20/613), cervical vertebra in A, posterior, B, right lateral, C, anterior, and D, dorsal views. Abbreviations: dp, diapophysis; ep, epipophysis; nc, neural canal; ns, neural spine; pf, pneumatic fossa; pl, pleurncoei; poz, postaygapophysis; pp, parapophysis; prz, preaygapophysis. La,evisuchus indicus. This taxon was connects the preaygapophyses with the described (Huerre 1932; Huene & Matley 1933:60- epipophyses, thus bounding laterally the dorsal 61, pl.XX, figs. 2-5) on the basis of three cervicals surface of the neural arch. The dorsal surface of (GSI K201613, GSI K201614, and GSI K271696) the neural arch is concave between the lateral and one dorsal ~rertebra(K271588). Unfortunately, rnargin and the neural spine. The latter is from these elements only a mid-cervical vertebra pyramidal, low (7 mm height) and craniocaudally was located at the GSI collections (GSI K271696; short (9 mm). Its cranial surface is damaged, but Fig. 11). The vertebra presumably corresporlds on the caudal surface exist ligament scars that do with cervical 5. The centrum is dorsoventrally low not reach to the top of the spine. On the cranial (the cranial surface is 19 mm high), and long surface of the neural arch, and ventroniedial to (nearly 42 mm), with an almost flat and the prezygapophyses, exist a pair of deep and transversely wide ventral surface. The cranial elliptical pneumatic fossae. The articular surface articular surface olthe centrum is kidney-shaped, of the prezygapophysis is smooth and slightly slightly concave, and. with rised borders. The cau- convex (transversely and craniocaudaily). The dal articular surface is also concave. A pair of caudal surface of tho neural arch is deeply pleurocoels are present on the sides of the excavated between the postzygapophyses and centrunl, and a pneumatic depression exists more diapophyses. The postzygapophyses are broken dorsally on the right side (this may correspond to dorsally, and corisequently their respective the "third" pleurocoel cited by Hnene & Matley, epipophyses are incomplete. However, some 1933). The parapophyses are prominent. The inlormation about their morphology is still neural arch is low and transversely wide (the available: the epipaphyses are projected dorsally distance between external margins of the andlaterally (as seen from behind; Fig. 11A).They prezygapophyses is 43 mm). Asharp dorsal margin are craniocaudally extended, roughly representing 80 Reuista del Museo Argentina de CienciasNaturnles, n. s. 6 il),2004 Fig. 12. Cervical vertehrae of abelisauroid theropods in right lateral (A-D) and dorsal (E-H) views. A, E, Masiahasaurus knopfleri (from Carrano et al. 2002); B, F,Laeuisuchus indicas; C, G, Noasaurus leali (from Bonaparte and Powel!, 1980); D, EI, Carrzotaurus sastrei (from Bonaparte et al., 1990),Not to scale. 75% of the maximum diameter of the (junior synonym of Cala?nospondylw Fox, 1866). postzygapophyseal articular surface. The caudal I-Iowever,the vertebra of Calamospondylus differs portions of the epipophyses have not been from Laeuisuchus in that only one pieurocoel is preserved. However, the epipophyses lack the present, the cranial articular surface is convex, and slender and conical cranial projections present in the dorsal surface of the neural arch is not Noasaurus (Bonaparte & Powell, 1981), for transversally wide and well defined as in example. 1,aeuisuchus. In sum, there are no coelurosaurian Huene and Matley (1933, pl.XX, figs. 2 a.nd 4) i'eatures in Laeuisuchus. On the contrary illustrated another two cervicals of Laeuisuchus Laeuisuchus shows the following abelisauroid (GSI K201613 and GSI K201614). Cervical GSI features: elongate epipophysis, pair of foramina on K201613 is remarkable fbr the extensive, iahle- centrum, pyramid-shaped, low and transversely shaped dorsal surface of the neural arch, ciosoly thick neural spines (Fig.121. resembling that of Noasaurus, Majungatholus his~~chushas cervicals that are proportiondly and Carr~otaurus.Cervical GSI K201613 is here longer than in Carnotaurus and Majungatholus. interpreted as more cranial in position than the Also, in Lueuisuchus the anterior articular surface proviously described cervical GSI K271696. is slightly concave, instead of being convex as in Reassons supporting this include a centrum with Carnotaurus. The articular surfaces of the diiYerently inclined cranial and caudal articular prezygapophyses are anteroposteriorly \-vide in surfaces, a proportionally smaller centrum Lueuisuchus, opposite to the transversaily expanded diameter with respect to the neural arch ones of Carnotaurus and Majiingatholus. (proportions that are also noticed in cranial cer- Laeui.suchus has large pneumatic cavities below the vical vertebrae of Carnotaurus, for example), the prezygapophysis, whereas in Carnotaurus and dorsal surface of the neural arch is wide, the late- Majungutholus the cavities are smaller in diameter. ral margin of the dorsal surface is straight in side Within ahelisauroids, Laeuisuchus more closely view, the neural spine seems to he absent, and the resemblesNoasaurus and Masiahasaurus. Carrano epipophysis seems to be well developed and et al. (2002) suggested that these ihree taxa could dorsally projected. be included within Noasauridae because they Norman (1990: 3021, following Huene and shares cervical vertehrae with anteriorly placed Matley (1933: 60-61),pointedout that thevertebra neural spines and cervical epipophysis that are of Laeui.suchus resemhles that of "Aristosuchus" reduced posteriorly Laeuisuchus and Noasaurus Novas et al.: Cretaceous theropods from India 81 Fig. 13. Omiihomirrtoides mobilis (K201614B1, dorsal vertebra in A, dorsal, and B, left lateral views. are similar in the development pneumatic cavities, crest, the neural spine is anteroposteriorly exten- the presurried abserice of neural spines on cranial ded, the base of the transverse processes is ventrally cervicals, and in the position of the both pre- and buttressed and excavated, and the apneumatic postzygapopliyses. centrum is longer than deep. We did not recognize Laevisuchm mersfromNoasaurus in havingthe autapomorphic features diagnostic of a~tediapophysial,postdiapphyi3ialanddiapophysial Ornithomimoides. On the contrary, the caudal cavities shallower, the diapophysis are wider and less vertebrae referred to this taxon look closely simi- ventraUy directed, and the neural spine is less exten- lar to caudals of other abeiisauroids. In sum, we dedanteroposteriorly Indorsalview Laeuisuchus has follow previous authors (Norman, 1990) in shorter prezygapophyses and the postzygapophyses considering Ornithomimoides as a nomen dubium, are caudally rounded (not acute as in Noasaurus). the set of vertebrae representing proximal caudals Imuisuchus differs from Masiaizasauras in having of an Abelisauroidea gen. et sp. indet. the space between the postzygapophyses less Dryptosauroides grandis. Dryptosauroides excavated, the prezygapophyses are thinner, and the was recognized by Huene and Matley, (1933; pl. infrapostzygapophysial and infraprezygapophysial XXII, figs. 1, 2, 3 and 4) on the basis of six dorsal cavities shallowec vertebrae (K201334, GSI K201609, GSI K271549, Omitl~omimoides.Iluene & Mattey (1933, pi. GSI K271601, GSI K271626, and GSI K27!602), but YX, fig. 8-10) created Lhis genus (with a pair of they also referred to this taxon a cervical vertebra species, 0. mobilis and O.? barasimlensis! on the (K271555) and several dorsal ribs (GSI K201615, basis of several vertebrae that this author GSI K271547, GSI K271623, GSI K271624, and GSI interpreted as dorsals similar to those of the K271625). This set ~Evertebraedoes not belong, ornithomimids Orr~ithomimusandStruthiomimw. in fact, to the dorsal, but to the caudal region. One of the species (0.mobilis! is represented hy Among them we have only accessed specimen GSI iive large and elongate vertebrae (GSI K201610, GSI K201609 (Fig.14) which exhibits almost the same K20/614B, GSl K271586, GSI K271597, arid GSI morphology as caudal vertebra GSI K201610 of K271600), and the second species (0.1 Ornithomimoides mobilis (Fig. 14). Caudal barasimlensis) by a set of smaller vertebrae (GSI vertebrae of Dryptosauroides match well with the K271531, GSI K271541, GSI II271604, and GSI K271 proximal caudals ofMajnngatholus @ers.obs.).As 682). Review of these specimens indicate that they is the case for Ornithomimoides the vertebrae of are not dorsals but caudalvertebrae (Fie.13)..",their Drvotosauroides., . corresaond to the nroximal morphology corresponding to those of caudals of an indeterminate abelisauroid. Majungatholus (pers. obs.).Asit occursin theiatter Consequently, Dryptosauroides grandis is abelisaurid, the prezygapophyses are close each considered as anomen dubium, followingprevious other, they lack of the ventral projections present authors (Molnar, 1990). The size of the caudals in dorsal vertebrae of neoceratosaurs (see Fig. 18 indicate the presence of a very large animal, foraniUustrationofsuchprojections),theprespinal surpassing the size of Carnotaurus (MACN-CH depression is deep and divided by a tiny sagital 8941, for example. 82 Reuista del Museo Argentina de Ciencias i?iaturales, n. s. 6 (11, 2004 Fig. 14. Left lateral views of dorsal vertebrae of Ornithomimoides mobilis (A) and Dryptosauroides grandis BE).From Huene and Matley (1933). Jubbulpuria tennis. This taxon was crected Matley (1933) erroneously interpreted as on the basis of two small vertebrae (K201612 and corresponding to the dorsal region. They all belong GSI K271614), identified by Huene and Matley to the caudal region, as already recognized by (1933, pl. XX, figs. ti and 7) as corresponding to Welles (1984) and Molnar (1990). Specimen GSI the dorsal region. Review of' available specimen K271595 (Huene & Matley, 1933, pl.XXI fig.5) is a GSI K201612 indicates that it is not a dorsal but a neural arch that closely resembles proximal distal caudal vertebra (Fig. 15). The centrum is caudals of Majungatholus (pers.obs.) and low and elongate, the transverse processes are "0rnithomimoide.s" (Fig. 13). However, specimens expanded and dorsally cxcavated (as in GSI K 271562 and GSI K27i574 of Coeluroides Coeluroides largus GSI K271562), and the neural largus show distinctive fcatures that merit more spine is anteroposteriorly extended. The spine is detailcd consideration. Caudal GSI K 271562 (Fig. represented by a tiny axial crest between the 16) is distinguished by its wide, almost prezygapophyses, but towards the rear it becomes horizontally oriented and wcll separated pre- and transversely stouter and was probably postzygapophyses; also, the transverse processes dorsoventrally higher. The postzygapophyses are notably expanded and triangular-shaped in (Huene & Matley, 1933, pl.XX, fig, tia,bj were dorsal view, with their dorsal surface deeply laterally facing, as it occurs in mid to distalcaudals excavated, thus resulting tho anterior margins of of other theropods (e.g., Majungatholus, the transverse proccsses beingraised. The ncural Allosaurus, Tyrannosaurus). spine is broken, but its basc is axially extended The caudal vertebra described as Jubbulpuria and transversely robust Zygapophyseal has wing-shaped transverse processes, which look rnorphology of GSI K271562 suggests that it is a well developed for such a distal caudal. Distal mid-caudal vertebra. The peculiar morphology caudals of abelisaurids (e.g., Majungatholus, described above is also seen in AMNH 1957 (Fig. pers.obs.j lack well developed transverse 171, a caudal vertebra catalogued as Indosuci&us processes. Also, its dorsal surkce is excavated, mploriils, which also has an elongate and low, different fro111 the dorsally flattened of apneumatic centrum. Moreover, specimens GSI abelisaurids. However, the Neocomian basal K271562 and AMNH 1957 are similar to the abelisauroid Ligahueino andesi (Bonaparte, 1996) fragmentary caudal vertebra GSI K201612 shows similarly developed transverse processes on referred to Jubbulpuria (Fig. 15) in the distal caudals, supporting referal of vertebrae of morphology of the transverse processes (e.g., Jubbulpuria as to Abelisauroidea. extensive, triangular shaped, and dorsally Romer (1956) agreed with Huene and Prlatley excavated). Interestingly, GSI K 271562 (1933) in that Jubbulpuria is a member of (Coeluroides largus), AMNI-I 1957 (catalogued as different from Abelisauroidea. Similarities noted above between distal caudals of Jubbulpuria tenuis and Ligabuieno andesi (Bonaparte, 1996) argues in favor that other candais with delta- shaped transverse processes (e.g., Coeluroides largus, AMNH 1957) also belong to Abelisauroidea. IV Axial skeleton remains originally referred to as "allosaunid" and "coelurosawid" "Allosaurid cervical vertebra" (K 271590) This was described as a cervical vertebra (Huene & Matiey, 1933, pi. XIV, fig. I), hut it is here reinterpreted as a dorsal vertebra because the prezygapophyses are close each other, and the parapophyses are projected outwards occupying a high position on the neural arch (Fig. 18). GSI K271590 is similar to dorsal 9 of Sinraptor (Currie & Zhao, 1993) in the morphology of the prezygapophyses, with pendant ventral processes, and the pattern of laminae connecting the diapophysis with the parapophysis. Also, the prezygapophysis and diapophysis are connected by a ridge that is dorsally convex in lateral view, a Fig. 15. Jubbulpuria tenuis (K20/612), distal cau- deep pneumatic cavity is located between dal vertebra, in A, right lateral, and B, dorsal prezygapophysis and parapophysis, and an "Y- views. Abbreviations: ns, neural spine; pre, shaped crest connects the parapophysis with the prezygapophysis; tp, transverse process. diapophysis. The same description also applies to dorsal 5 of Carnotaurus and dorsai 7 of Ceratosaurus (Welies & Madsen, 2000), but in these twoiaxa and GSI K271590 the parapophyses are more laterally prominent than in Sinraptor. 595, GSI K201610. GSI K20/614B, GSI K271614, Also, the prespinal cavity is large and deep, a GSI K271586, GSI K271597, GSI K271600). It is synapomorphic trait shared by all neoceratosaurs iniportant to note that mid-caudal AVNH 1957 (Holtz, 2000). In sum, dorsal GSI K271590 exhibits is not only different in morphology from proximal neoceratosaurian features, and because its caudal AMNH 1960 (also catalogued as morphology is congruent with that of abelisaurids Indosuchus raptorius), but it is iarger than the and it was found in association with abelisaurid latter, thus indicating that they do not belong to bones, we refer this specimen to Abelisauroidea a same individual (and presumably pertain to indet. different species). "Allosaurid cervicalvertebra"(K271572). Possibly Coeluroides largus may represent a This is a large vertebra (16 cm height), wish an valid taxon of an indeterminate ahelisauroid opisthocoelous centrum, albeit the cranial articu- theropod. Mid and distal caudals of Coeluroides lar surface is almost flat (Huene & Matley, 1933, retained notably developed neural spines. lam. XIV fig. 2). Huene cites that a single Comparing AMNH 1957 with the similarly pleurocoel is present below the diapophysis. This elongated and low caudal centra of caudals of cervical does not resemble that of Aheiisauridae Majungatholus, it becomes evident that in AMNH in the shape of the neural spine (which is axially 1957 the transverse processes are well developed, extented and transversely narrow, instead of aliform structures, whereas in Majungatholus craniocaudally short and transversely wide as in they are absent or represented by a faint abelisaurids), and the apparently poor longitudiilal ridge. The evidence is not enough to development of the epipophyses (in contrast with evaluate whether Coeluroides largus and the high and craniocaudally extended epipophyses Jubbulpuria tenuis are synonyms, but their cau- of abelisauroids). Unfortunately, specimen GSI dal vertebrae may represent theropod lineages K271572 is lost at the GSI collections, and first 84 Reuista del Museo Agentino de Ciencias Naturules, n. s. 6 ilj, 2004 Fig:. 16. Coeluroides largus (K.271562),mid caudal vertebra, in A, left lateral, B, dorsal, and C, poste- rior views. Abbreviations: ns, neural spine; poz, postaygapophysis; prz, prezygapophysis; tp, transverse process. Fig. 17. Mid-caudal vertebra of an indeterminated abelisauroid (AMNH 1957) in A, dorsal, and B, left lateral views. Abbreviations: ns, neural spine; poz, postaygapophysis; prz, prezygapophysis; tp, transverse process. hand observations are needed to test whether this gle, rod-like structure similar to that of represents clade of theropod other than Carnotaurus. Although we do not dismiss that Abeiisauroidea in the "Carnosaur bed". specimens GSI K271533 and GSI K271554 belong Sacral vertebrae. Huene and Matley to Abelisauridae, their morpholog~more closely illustrated some portions of fused sacral.vertebrae resembles that of tainetasaurus (Matley, 1923), under the numbers GSI K271.554 (twopieces), GSI Rajasaurus (Wilson et al., 2003) and K27/533 (two pieces) and GSI K271571 (Fig. 19). iWasiakasaurus (Csrraro etal., 2002) in that each More recently, Bonaparte (1991b) referred sacral element is transversely broad and the specimens GSI K271533 and GSI K271554 to contact between succesive vertebra is well Abelisauridae because they are fused into a sin- marked. In Carnotaurus, instead, the sacral cen- Nouas et al.: Cretaceou s theropods from. lndla 85 tra are strongly reduced in transverse diameter and the contacts among succesive sacrais are slightly marked. Notably, specimen GSI K271571 (Eirstly interpreted by Huene and Matley as coelurosaurian) responds to the Carnotaurus morphotype, thus suggesting the presence of two different large abelisauroids in the fossil assembiange. There is a large isolated centrum (GSI Pi271 598; 19C,E) with a strong constriction at mid- lenght, but with highly expanded articular facets. This morphology remember that of sacral 1 of Rajasaurus Wilsonet al., 2003), thus we interpret specimcn GSI K27i598 as a probable sacral 1. Proximal caudal vertebra (AMNH 19601.This proximal caudal is represented by a neural arch with elongate transverse processes, which are not entirely preserved at their extremities (Fig. 20). General morphology of AMNH 1960 is congruent with that of the proximal caudals of Majungatholus, Carnotauras and Abelisauridae indet. MPM 99. However, derived traits of Abelisauridae (e.g., distally fan- shaped transverse processes, and presence of a slender anterior projection on the transverse processes that contacts with the transverse process of the contiguous anterior caudal; Martinez et al., in press) are not identified in AMNH 1960. AMNIl 1960 lacks well developed hyposphene-hypantrum articulations, thus differing from Aucasaurus and Carnotaurus in which hyposphene-hypantrum are present in the proximal and middle sections of the caudal series (Curia et al., 2002). Also, in AMNH 1960 the transverse processes are laterally oriented, instead of dorsoiaterally as in the abovementioned Patagonian abelisaurids. Although this orientation may depend on the position of the vertebra in the caudal series, Majur~gatholusand a new abelisaurid specimen from Patagonia (Abelisauridae indet. MPM 99; Martineia et al., in press) also exhibit laterally projected transverse processes. Medium and distal caudals. Some distal caudals (Fig. 22) are characterized by a po1yg.onal centrum in cross-section, transverse processes represented by thick ridges overiapping both sides of the centrum, flat ventral surface of the centrum, rounded and short prezygapophyses, and sY>>- Fig. 18. Dorsal vertebra of an indeterminated shaped neural arch in dorsal view (being cranially abelisaurid (JC.271590) in A, lateral, B, anterior, and bifurcated towards the prezygapophyses). Distal C, dorsal views. Abbreviations: nc, neural canal; caudals with these features are: AMNH 1958, GSI pp, parapophysis; prz, prezygapophysis; psf, K271596 (Fig. 221, K271532, and GSI K271594 prespinal fossa; pvp, pendant ventral process. ,(Huene & Matley, 1933, pl. XXIII, fig. 2, and pl. 86 Reuista del Museo Argentino de Ciencias Naturales, n. s. 6 fli,2004 Fig. 19. Ahelisauroid sacral vertebrae in ventral (A, B, F, H, J) and lateral (D, E, G, I, K) views. A, specimerl GSI K271533 (from Huene & Matley, 1933); B, D, specimen GSI K271554 (fi.om Huene & Matley, 1933); C, E, specimen GSI K271698 (from Huene & MaLley, 1933); F, G, Rajasaurusnarmden,sis (from Wilson et al., 2003); H, I, Carnotaurus sastrei (from Bonaparte et al., 1990); J,K, specimen GSI K271571 (from Huene & Matley, 1933). Not to scale. Fig. 20. Proximal caudal vertebra of an indeterminate ahelisaurid (AMNH 1960) in A, dorsal, B, ante- rior, and 6,posterior views. Novas et al.: Cretuceous theropods from lndza Fig. 21. Distal caudal vertehra of abelisauroids in lateral (A,U) and dorsal (C,D) views. A, C, indeterminated abelisauroid (K271599), andB,D, Masiakasauri~sknopfleri (from Carrano et al. 2002). Abbreviations: ns, neural spine; poz, postzygapophysis; prz, prezygapophysis; tp, transverse process. XY fig. 6, respectively). Caudals GSI K271589 transverse processes. This suggests the presence (t~coelurosaurid~~,Huene & Matley, 1933, pl. XXIII, of abelisauroids with different kinds of caudal fig. 3) and GSI K271705 exhibit a similar pattern, processes although their prezygapophyses and centra are Obviously, more work needs to be done on the longer and thus they correspond to the distal end caudal anatomy of abelisauroids in order to resolve of the tail. the allocation of isolated vertebrae, recognize the Tlie kind of caudais described previously morphological variations alongthe tail series, and contrasts with another group in which the evaluate the taxonomic significance. transverse processes are well developed, delta- Haemal arches. Several isolated haemal arches shaped (i.e., caudoiateraily expanded), and have elongate centra and prezygapophyses. This group of vertebrae includes caudals of Jubbulpuria. An Except for GSI K271680, the haemal arches are isolated caudal vertebra GSI K271599 (Fig. 21), elongate and rod-like, and lack a distal expansion, assigned by Huene and Matley (1933) to a resembling the condition seen in other ahelisanrids and Ceratosaurus Wilson et al. 2003). In contrast coelurosaurid, resembles Masiakasaurus with Carnotaurus, at least, the Indian chewrons (Carrano et al., 2002) in the general shape and posses a haemal canal that is proximally open. principally in the shape of the transverse processes and elongate prezygapophyses. The latter suggests V Pelvie and hind limb bones originally this is amid- to distal caudal which retained well referred to as "allosaurid" and "coeluro- developed transverse processes. This combination saurid" of features sharply differs from the condition seen in other caudals with equally longcentra (Fig. 221, With the exception of two fragmentary ischia, but with short prezygapophyses and nearly absent no other pelvic bone was describcd from the 88 Revista del Museo Argentino de Cieneias Naturales, n. s. 6 (11, 2004 Fig. 22. Distal caudal vertebra of indeterminaced abelisauroids in left lateral (A-Cj and dorsal (D-F), views. A, D, specimen GSI K271705; B,E, specimen GSI K271596; C,E: specimen AMNH 1958 (reversed). Cranial is to the left. "Carnosaur bed". However, several hind limb Carnotaurw, and that both bones would not have elements (femora, tibia, metatarsals, and belonged to the larger forms found in the phalanges) have been recovered and they offer a "Carnosaur bed". good source of anatomical information. Huorie Femur. Most of the femora described by and Matley (1933) distinguished two types oi' Huene and Matley (1933) are between 60 cm and femora, one of a robust animal and another more 74 cm in length (Fig. 23), thus belonging to slender one. IInene associated the robust kind of animals of large size. We did not locate the great femora with a single, equally stout tibia, and majority oi'the femora at the GSI collections, and referred them to the eallosanrids.>.Walker (1964) our comments will mostly rely on Huene and considered the slender kind of femora as Matley 's i:lustrations. The exception is a proximal belonging to Inilosaurus and the stouter type to portion of a left femoral shaft newly cataloged Indosuchus. with number GSI 296 which lost the correspon- Ischiurn. Two fragmentary proximal ischia ding numbers of the GSI K series (Fig. 24A). We were described by IIuene (K271686 and GSI K271 presume that it may belong to any of the left 546; pl. XVI, figs. 7, 8). They are poorly femora (either GSI K271564 or GSI K271563) cited, informative, and their morphology matches with but not figured, by Iluene and Matley (1933:55). most basal theropods. The distance between the As mentioned before, Hnene and Matley distal extremity of the obturator process and the sorted out the available 9 theropod femora from than 4 cm. This is in contrast to the holotype of femora (GSI K271560, GSI K271563, GSI K271564, Carnotaurus, for example, in which the distal tip GSI K271569, GSI K271621, GSI K271627). The of the obturator process is 30 cm from the iliac robust femoraare characterized by their relatively pedicle, and the craniocaudal diameter of the shaft short and robust shafts, thus looking sharply (immediately distal to the obturator process) is 7 diff'erent from the remaining non-avian cm. This indicates that ischia GSI K271686 and theropods. They may belong to a single taxon (e.g., GSI K271546 belonged to animals smaller than species) characterized by stout hindlimbs, Nooas et al Cretaceous theropods from Tndio C K271570 D K271558 Fig. 23. Abelisaurid femora in caudal view. A, specimen GST K271569; B, specimen GSI K271560; C, specimen GSI K271570; D, specimen GSI K27155S. Abbreviations: at, anterior trochanter; 4t, fourth trochanter. although some distinctions are observed among femora described by Huene and Matley belong to them, for exaniple GSI K271570 appears to exhibit Abelisauroidea, a conclusion that is in agreement a well-developed (i.e., proximally projected) an- with the whole bone assemblage, mostly (if not terior trochanter, whereas in GSI Ii271558 this entirely) made up by abelisauroid hones. trochanter is smaller. This last specimen, at least, Tibia.Huene and Matley (19333 described three exhibits features resembling Xenotarsosaurus theropod tibiae corresponding to large theropods, (UNPSJB-PV 184-6121and Indosuchus (IS1 R911 none of which was available for tho present study I), including: rounded femoral head, anterior at the GSI collection. Only GSI K271568 was trochanter low with respect to the femoral head, illustrated by Hueno and Matley (Fig. 25A). As 4'" trochanter convex in side view, and presence earlier suggested by Bonaparte (1991b1, this of a prominent mediodistal crest. The slender specimen resembles Abelisarlridac in havinga well- specimen GSI 296 (Fig. 24A) exhibits the developed cnemial crest, and apoorly differentiated following resembidnces withXenotarsosaurus and outer condyle on the proximal end, which is located Carnotaurus: the anterior trochanter is cranially at almost the same level as the inner condyle. The convex in lateral view, the trochanteric shelf is tibia bears an elonbwte cnemiai crest as usual in prominent and located at level of the distal end neoceratosaurs, but the shaft is remarkably short of the anterior trochanter, the 4"' trochanter is and stout, being clearly different from other also convex in side view, and a conspicuous pit theropods, including most ahelisauroids (e.g., fbr attachment of the M. caudofemoralis is present Aucasaurus, Xenotar.sosaurus, Majungatholus, on the medial surface of femur, cranial to the 4l" Masiakuswrus). Theonly exceptionsare thelndian trochanter. Larnetasuurus (Matley, 1923; Fig. 25B) and the All of the 9 femora discovered in the Braziiian Pycnonenzosaurus (Kellner & Campos, "Carnosaur bed" are morphologically congruent 2002), in which the tibiais proportionally short. This with the femora of other abelisaurids (e.g. peculiar condition of the tibia is not due to Xenotarsosaurus, Carnotaurus, and IS1 specimens deformation, loss of its distal portion, or a of Indosuchus). Our iriterpretation is that the pathological case, and therefore it constitutes a ,:% Reiiisia del Museo Argmtino de Ciencias Natumles, n. s. 6 ili, 2004 .a>sz >% ,; Allosaurus (Madsen, 1976). The fibula GST K271 620 and that of Rajasaurus differ from that of the abelisauroid Deltadromeus (Wilson ei al., 20031, because in the latter taxon the fossa, albeit deep, is not subtrianylar but proximodistally elongate. In sum, GSI K271620 is recognized as an abelisauroid fibula. Astragalus and caleaneum. As mentioned in previous pages, the purported astragalus (K271 684; Huene & Matley, 1933, pl. XIX, fig.l) is in fact a left quadrate. Besides, the purported calcaneum (K201396; Huene & Matley, 1933, pl. XIX, fig. 2) is considered here to be an indeterminate bone. Metatarsals. Soveral isolated metat,arsals were recovered in the "Carnosaur-bed". Review of these elements (either in the GSI collection or on the basis ol Huene & Matley's illustrations) indicate that: 1) the elen~entsoriginally thought as helouging to the manus correspond in fact to the pes; 21 available metatarsals correspond to metatarsal 11, 111 or IV; and 3) all these uieces exhibit abelisauroid leatures. Fig. 24. Abelisaurid femora in lateral view. A, Metatarsal I1 (Fig. 27) is represented by specirnen GSI 296; B, Xenotarsosaurus bonapartei specimen GSI K271671 (Huene & Matley'e. (UNPSJB-PV 184 and 612). Abbreviations: at, "allosauroid, distal extremity of mLt 11';) and anterior trochaiiter; fh, femoral head; ts, presumably GSI K271667 (Huene & Matley's trochanteric shelf; 4t, fourth trochanter, "coelumsaurid, distal end ol metacarpal"), both of which exhibit resemblances to metatarsal I1 of Masiakasaurus (Carrano et al.. 2002). They bear a double-flanged distal condyle of which the late- derived trait only documented in GSI K271568 and ral flange is substantially larger than the medial Lametasaurus. one, as in Masiakasaurus. Huene and Matley (1933) also described some Bones recognized here as metatarsal 111 (Fig. other tibiae interpreted as belonging to 28) include specimen GSI K271665 (Huene & aCoelurosaurian (GSI K271526, GSI K271670, GSI Matley 's "coelurosaurid, probably rnt,t III"), GSI K271552, GSI Ii271556, GSI K271662, and GSI K271658 (Huene & Matley 's "allosaurid, left mtt K27/669), but none of them could be located at III"), and GSI K271697+ GSI K271681 (Hueue & the GSI collections, with the exception of GSI K271 Matley's "coelurosaurid, probably mtt It"). In 669. We doubt that specimens GSI K271526 and them, the ginglymus is dorsoventrally low, as in GSI K271669 were correctly identified as tibiae, the metatarsal I11 of basal ceratosaurians and and we prefer identify them as indeterminate limb abelisaurids (Valais et al., 2002). It is interesting bones. to note the differences in size, proportions and Fihula. An incomplete left fibula (K271620; shape of metatarsal I11 of specimens GSI K271658 Fig. 26Aj resembles that of abelisaurids (e.g. (a large abelisauroid with a metatarsal 25.4 em Xenotarsosaurus, 12ajasaurus) in having a long and with a distal ginglymus 5.2cm thick; Fig. prominent iliolibularis tubercle and a well 28A-Dl, and specimens GSI K271665 and GSI K271 excavated fossa on the medial surface of the 697+681 representingsleiider forms withadistal proximal end. Reserriblances between GSI K271 ginglymus 2.8cm wide. 620 and the fibula of Rajasar~rus(Wilson et al., Finally, ~netatarsalsinterpreted here as 2003) include the subtriangular contour of the metatarsal IV (Fig. 291 include the lollowing proximal fossa, which is bounded by strong cranial specimens: GSI K271539 (Huene & Matley's and caudal ridges. Distal to the fossa, both ridges "allosaurid, right metatarsal N";25cm long), GSI join to form asingle, prominent longitudinal ridge K271659 (IJuene & Matley's "allosaurid, right extending along the fibular shaft. GSI K271620 metatarsal IV"), GSI K271666 (Ifuene & Matiey's and Rajasaurus appear to lack the longitudinal "coelurosaurid, distal hall' of metacarpal"), and groove present in tile allosauroids Sinraptor and GSI K201337C (Hueile & Matley 's "coelurosaurid, Nouar : et al.: Cretaceous theropods from Indm E3 G D E Fig. 26. Tibiae of several ahelisauroid in right lateral view. A, specimen GSI K271568 (from Huene & Matley, 1933); B, Lametasaurus (from Matley, 1924); C, Indosuchus (IS1 R9111); D, Pycnonemosaurus (fom Kellnor and Campos, 2001); E,Aucasaarus (from Coria et al., 2001);Xenotarsosauras (Martinez et al., 1986). probably loft metatarsal I"). All these hones exhibit deep and transversely compressed distal ends, with asymmetrically developed articular condyles ii.e., the inner condyle is more developed than the outer one), features that also apply to metatarsal IV of the ahelisauroids Masiaka.sauras (Carrano et ul. 20021, Deltadromeus (Sereno et al. 1996), Aucasaurus (Coria et al., 20021, and Abelisauridae indet (MCA 56). As far as Huene & Matley's figures suggest (pl. XIX, figs. 5 and 6, and pl. XXIX fig. 41, two kinds of metatarsal IV may be recognized: one in which the distal ginglyrnus is relatively robust (K271539) and others with a transversely narrower girrglymus (K271659 and GSI K271666). Also, the shaft exhibits adifferent contour in transverse section: in GSI K271539 it is trapezoidal-shaped wiih the longest side dorsal, instead in GSI K271659 the transverse section is subtriangular, with the longest side ventral (Fig. 29). Such differences may correspond to two kinds of pes within Abelisauroidea, one in which metatarsals and their respective phalanges are robust, and another kind in which side metatarsals (11 and especially IV) are more slender, as well as their respective phalanges, as it occurs in Velocisaurus (Bonaparte, 1991a). Fig. 26. Fibulae of abelisaurids in medial view. A, Pedal phalanges. Huene and Matiey specimen GSI K271620 (from Huenc & Matley, (1933:57) noted that "there are more than 40 1933); B, Rajasaarus narmadensis (right fibula, phalangeal bones", hut they described three reversed; from \Nilson et al. 2003). phalanges (e.g, GSI K2716.51, GSI K271652, GSI 92 Revzsta del Museo Argenttno de Ctenctas Naturales, n. s. 6 (I), 2004 phalanges, we have only accessed GSI K20/626B, B GSI K271648, GSI K271524 (this last number has ~27m been also applied to a pedal ungual; Huene & Matley, 1933, pl.XIX, fig. 13). With the aim of determining their tentative positions in the pes, we have sorted out the phalanges illustated by Huene and Matley (1933, pls. XIX and XXIV) on the basis of their morphology, size, and relative proportions, identifying themas belonging to digits 11,111and IV This task was also supported by comparing this set of pedal elements with phalanges of other theropods !e.g., Allosaurus, Sinraptor, Velocisaurus. Aucasaurus). It is clear that the phalanges correspond to an~malsofd~fferent slze and robustness The phalanges of digit I1 are represented by specimens GSI K271654 (Huene & Matley's "allosaurid pedal digit I") and GSI K271524 (Huene & Matley's "coelurosaurid manual digit I"). The morphology of these phalanges matches well with that of pedal phalanx 1 of digit 11 of Velocisaurus (Bonaparte, 1991aj in being elongate, strongly asymmetrical, with a dorsoventrally deep proximal end, and a pair of well developed proximoventral longitudinal ridges (Fig. 30, A,B). However, specimens GSI K271654 (8 cm long) and GSI K271524 (6 cm long) correspond to an animal considerably larger than Velocisaurus. in which nhalanx 111-1 reaches 2.3 cm long. The phalanges of digit I11 are represented by the following GSI specimens (Fig. 30, C-F): GSI K271653 (Huene & Matley's "allosaurid foot phalanx"), GSI K271646 (Huene & Matley's "coelurosaurid pedal phalanx of digit IV"), GSI K271525 (Huene & Matley's "allosaurid pedal - phalanx"), and GSI K271644 (Huene & Matlev's "coelurosaurid pedal phalanx"). Thev are Fig. 27. Metatarsal I1 ofabelisaurids in dorsal and symmetrical and dorsbventra~l~depressed distal views, A, Rajasaurus narmadensis (from phalanges, which are more robust than the Wilson et al. 2003); B, specimen GSI K271667 remaining phalanges. Their proximal ends are (*om Huene & Matley, 1933); C, specimen GSI laterally and medially flared for articulation with K271671 (*om Huene & Matley, 1933). the corresponding metatarsal 111 or the preceeding phalanx. In proximal aspect they are crescent-shaped. Available phalanges of digit I11 are characterized by the presence of a low and wide proximal articular surfice, in congruence K2716541, of which only GSI K271654 has been with the subrectangular distal condyle of illustrated (Huene & Matley, 1933, pl.XIX, fig. 7). metatarsal III..In lateral view, the dorsal margin They are relatively large, measuring between 8 of these phalanges is more or less straight, and and 7 cm long. Specimen GSI K271654 was the ginglymus lacks a dorsally expanded articu- originally interpreted as belonging to digit I, hut lar facet, being slightly more depressed than thc we interpret it as corresponding to phalanx 11-1. rest of tho dorsal margin of the bone, a condition In addition, 18 non-unguals phalanges were also contrasting with most other theropods (e.g., listed by the same authors as belonging to smaller Allosaurus and Sinraptor). Specimen GSI K271 theropods ("coelurosaurids"). From this set of 653 (Fig. 30, D) is interpreted here as a pedal F Nouas et a1 Cretaceous theropods from Indza 93 F Fig- 28. Metatarsal 111 of abelisauroids. A-D, specimen GSI K271658 (left metat.arsalII1) in medial (A), dorsal (B), ventral (C), and lateral (D) views; E,F, specimen CXiI KC271665 in dorsal (E), and side (F) views (from Huene & Matley, 1933); G, specimens GSI K271697+681 in dorsal view (from Huene & Matley, 1933). phalanx 111-2 and GSI K271646 (Fig. 30, E) GSI K271642 (Huene & Matley's "coelurosanrid probably corresponds to pedal phalanx 111.1. The manual phalanx"), and GSI K27164R (Huene & latter one resembles Aucasaurus in its robust Matley's "coelurosaurid manual phalanx"). In proportions, albeit its size (3.6cm long and 2.6cm agreement with the distal condyle of metatarsal wide proximally) indicates that it probably IV described above (K2716591, phalanges of this corresponds to a juvenile individual of a robust digit are transvei.se1y narrow and dorsoventrally abelisauroid. In addition, GSI K271525 is deep, in sharp contrast with those of non- considered to be pedal phalanx 111-1 (it measures abelisauroid theropods such as Sinraptor, 7.6cm long and 5cnl wide proximally), and GSI AIZosauru~sand Deinonychus, in which the K271644 as pedal phalanx 111-2 (4.6cm long and phalanges of'digit IV are proportionally lower and approximately 2.3cm wide proximally). In sum, wider This is a notable character not described two subsets of digit 111 phalanges seem to be before for abelisauroids, except for distinguished by their relative proportions: GSI Masiakasaurus (Carrano et al., 2002). In contrast K271653 and GSI 11271646 are proportionally with the above described phalanges of digit 111, robust, whereas GSI K271525 and GSI K271644 those from digit IV have deep dorsoventral are of more slender proportions, in particular the grooves on their distal ginglymoids. latter specimen which resembles Velocisaurus Specimen GSI K201626B was interpreted by (Bonaparte, 1991a). Huone and Matley (1933, pl. XXIV, fig. 7) as a The phalanges of digit IV are represented by "coelurosaurid manual phalanx". However, this the following specimens (Fig. 30, G-M): GSI K201 is a pedal element that closely resembles pedal 337B (Huene & Matley's "coelurosanrid foot phalanx IV-1 of Velocisaurus (Bonaparte, 1991a). phalanx"), GSI K271637 (Huene & Matley's As in the latter taxon, GSI K201626B is "coelurosaurid foot phalanx"), GSI K271638 proximodistally short hut transversely (Huene & Matley's "coelurosaurid foot phalanx"), compressed (Fig. 30, G-I). GSI K201626B is pcdal GSI K271647 (Huene & Matlev's "coelurosaurid phalanx IV-1, and GSI K271648 (Fig. 30, I) is 94 Reuista del Musro Argeutino de Cz encias Naturales, n. s. 6 fli, 2004 identification of such pedal elements, but the phalanges of digit IV lack of the proximovcntral ridges characteristic of the proximal phalanges of digit 11. Nso, the medial surface of digit IV phalanges is high and almost flat, and exists on its proximoventral corner a deep excavation. Although specimens GSI K20162fiB and GSI K271 648 exhibit a similar morphology, they appear to belong to different individuals: GSI K201626B is 5cm long and 2.2cm wide, but GSI K271648 is 2.6~~1long and 1.2cm wide, suggesting that individuals in different. growth stages, and presumably belonging to a same species (e.g., a gracile ahelisauroidj, are represented in the "Carnosaur bed". Huene and Matley say (1933:67) that phalanx GSI K201626B ~crfectlvarticulateswith the distal halfof a bone that they thought as a "metacarpal of a coelurosaur" (K271666, here reindentitied as distal end of metatarsal IV). There is another group of phalanges of digit IV (Fig. 30, J-M) which are very short, deep and transversely wide, showing a more conservative morphology similar to that present in other theropods (e.g., Sinraptor, Allosaurus, ornithomimids). Digit IV phalanges of the robust kind are similar to those of the abelisaurid Aucasaurus (Coria et al., 2002), and they can he sorted out on the basis of their size: GST K201337B (probably a pedal phalanx IV-2; it is 2.4cm long and 1.2cm wide proximally) and GSI K271647 (in- terpreted as a pedal phalanx IV-4) may correspond to a single specimen of small size. Instead, GSI K271638 (identified here as phalanx IV-3, being 3.8cm long and 2.6cm wide proximally), and GSI K271!337(interpreted as phalanx IV-4; it is 2.8cm Fig. 29. Metatarsal IV of abelisauroids in dorsal long and 2.4cm wide proximally), are short, wide and distal views. A, specimen GSI K271539 (mid- and deep pedal digit IV phalanges, responding to shaft cross-section indicated on its right); B, an Aucasaurus kind of hot but belonging to a specimen GSI K271659 (mid-shaft cross-section larger specimen. indicated on its left); C, specimen GSI K271666; Summing up, available pedal digits indicate D, specimen GSI K201337 (all figures taken from that: 1) they are congruent with abclisauroid Iluene & Matley, 1933). anatomy; 2) differences with the set of trans- versely narrow digit TI1 and IV phalanges de- scribed before may reveal the presence of more than one type of abelisauroid species in the quarv, i.e., an Aucasaurus-like foot with more robust in being transversely compressed, with a distal phalanges on digits 111 and especially IV (Figs. ginglymus asymmetrically developed (i.e., the 29, 321, and a Velocisaurus-like foot with slender inner condyle is wider and deeper than the outer phalanges (Fig. 31). one), and with a deep dorsoventral groove on There were many more phalanges (Haene & distal ginglymus. They also exhibit well excavated Matley, 1933:67), originally described as belong- collateral and extensor ligament pits. The ing to a single foot, but such pedal elements were proximal end ofboth phalanges is triangular, with not illustrated with the exception of GSI K271646, the long axis oriented dorsoventrally This GSI K271647 and GSI K291337B (see Figs. 30 and condition resembles the proximal phalanx of digit 321, so there are no possibilities to evaluate such TI, and may lead to confusion regarding the an association. Fig. 30. Pedal phalanges of ahelisauroids. A, phalanx 1.11 (K2715241, in proximal, lateral, dorsal and ventral views; B, phalanx 1.11 (K271654), in proximal, lateral, dorsal and ventral views; C, phalanx 1.111 (K271525) in lateral and dorsal views; D, phalanx 2.111 (K271653) in dorsal view; E, phalanx l?.III (K271646) in proximal, lateral and dorsal views; F, phalanx 2.111 (K271644) in lateral and dorsal views; G, phaianx 1.1V? (K271642) in proximal, lateral and dorsal views; H, phalanx 1.IV (K201626B) in proxirnai, lateral and dorsal views; I, phalanx 3.1V (K271648) in proximal, lateral and dorsal views; J, phalanx 3.IV (K271638) in lateral and dorsal views; K,~phalanx4.IV (K271637) in lateral and dorsal views; L, phalanx 2?.IV (K201337B) in proximal, lateral and dorsal views; M, phalanx 4.IV (K271647) in proximal and lateral views. All figures taken from Huene and Matlej: 1933. Reuista del Museo Argentina de Ciencias Naturales, n. s. 6 ili, 2004 Fig. 31. Abelisauroid foot. A, composite reconstruction based on different pedal elements (indicated on the figure). B, articulated metatarsals and phalanges of the left foot of Velocisaurus (from Bonnparte, 1991a), in dorsal view. C-E, matatarsal IV (K271666) articulated with phalanx 1.IV (K20/626B), in dorsal (C), distal, proximal, and side views; F, composite reconstruction of pedal digit IV in side view, based on different pedal elemcnts (indicated on the figure), and compared with same digit of' Velocisaurus (from Bonaparte, 1991a). Unguals. As analyzed elsewhere (Novas & CONCLUSIONS Bandyopadhyay, 2001) the set of ungual phalanges figured by Huene and Matley (1933) correspond The review of the theropod hones collected at to the pes, thus dissmising interpretations of these the "Carnosaur bed" demonstratcs that authors that at least some unguals belong to the differences in shape, size and proportions of hand. Also, these pedal unguals exhibit the same postcranial and cranial bones support the morphological pattern (e.g., presence of proxi- mally bifurcated grooves, rounded bump on the presence of individuals at different growth stages. lateral side of pedal unguals, and ventral surface Cranial and postcranial elements exhibit excavated or witb a narrow deep furrow), de- ceratosaurian, abelisauroid or abelisaurid traits, scribed for other abelisauroids (Abelisauridae or they are n~orphologicallycongruent witb indet. MCA 56, Masiahasaurus; Novas & abelisauroids. In other words, most, if not all, of Bandyopadhyay, 2001; Carrano et al., 2002). For the theropod specimens ducumented in the quarry a more exhaustive review of the pedal unguals, correspond to Abelisauroidea. The only possible see Novas and Bandyopadhyay (2001). exception may be a large cervical vertebra (K271 Novas et al.: Cretaceous theropods from Indza Fig. 32. Abelisaurid Coot. Different pedal elc~nentscorresponding to individuals of different sizes, eventually referable to as Ahelisauridae. A, metatarsal IV (K271659); B, metatarsal IV (K271539); C, metatarsd 111 (K271658); D, metatarsal I1 (K271671);E, phalanx 3 IV (K271638), F, phalanx 4 IV (K271 637); G phalanx 1111 (K271525);H, phalanx 1111 (K271646); I, phalanx 2 I1 (K271654);J, phalanx 4 IV (K201337); K, phalanx 2 I11 (K271653). Ail figures taken from Huene and Matley, 1933. 672) which does not exhibit features consistent grandis, Of-nithomimoides barasimlensis, and with this clade. Jubbulpuria tenuis, Coeluroides Eargus) are here Most of the taxaoriginally described by Huene considered nomina duhia, in agreement with and Matley (1933) on the basis of vertebral previous authors (Molnar, 1990; Norman, 1990; remains (Con~psosuchussolus, Dryptosauroides Welles, 1984). The axis of Compsosuchw solus 98 Reuista del Museo Argentina de Ciencias Naturales, n. s. 6 /I), 2004 may belong to an abelisauroid the sizeof Carnotaurw among abelisauroids, but the more Carnotaurus, and it is not illogical to refer this conservative type (in which each sacral eleinent vertebra to Indosuchus raplorius because of its is transversely broad and the contact between similarities with this taxon. Besides, the proximal succesive vertebrae is well marked) are caudals of llryptosauroides grandis and documented among abelisauroids hot11 in Orfiithomimoides barasimler~sis are Ahelisauridae (e.g., Rajasaurus) and Noasauridae morphologically congruent with the proximal (e.g., Masiakasaurusj. For this reason, it is pro- caudals of abelisaurids (e.g., Majungatholus). The bable that the "Carnosaur bed" includes remains caudals of Jubbulpuria tenuis and Coeluroides of different forms ofAbelisauridae. In this regard, largus may also helongto Abelisauroidea (because we concur with Huene and Matley about the of their resemblance with the Patagonian distinctions in femoral proportions ("slender" vs Ligabueilao), albeit they seem different from the "robust") see11 iii the femora of large abelisaurids. caudals of abelisaurids, thus suggesting that the However, referral of this set of hindlimb bones fossil assemblange at the "Carnosaur bed" either to Indosuchus or Indosaurus is not includes the remains of individuals corresponding warranted at the moment. to different abelisauroid species. As already mentioned in the Introduction, Sorting out the available cranial and the ahelisaurid Lametasaurus indicus was postcranial materials into discrete individuals and found in the "Carnosaur bed.Since this taxon taxa is currently not possible, but some postcranial characterizes by a stout and transversely wide bones (sacral and caudal vertebrae and pedal tibia, we interpret the short and stout femora bones) show contrasting morphological patterns. (GSI K271558, GSI K271570, GSI K271618) and Among equally elongate distal caudals, some bear tibiae (GSI K2715681 discovered in the same short prezygapophyses and almost absent quarry as probably belonging to Larnetasaurus transverse processes, instead other caudals bear indicus. Moreover, it would not be dismissed elongate prezygapophyses and well developed, that Lametasaurus indicus may constitutes a triangular-shaped transverse processes. We also senior synomym of the also robust Indosaurus recognize two different pedal types (i.e., rnatleyi and Rajusaurus narmudensis. transversely wide vs. transversely narrow pedal The fragmentary nature of the Indian digit IV) and two types of sacrum (i.e., a abelisaurids, including Lametasaurus, Rajusaums, conservative one in which sacral centra -although Indosaurus and Indosuchus (except specimen IS1 fused- remain distinct from each other, vs. a RBI11 referred to as Indosuchi~sraptor-ius, still Carnotaurus-like sacrum in which the centra are awaiting a detailed description), obstructs easy rod-like, with smooth i~~tervertebralcontacts). recognition of the taxonomic validity of each of Such distinctions in sacral, caudal, and pedal these taua. Eventual solution of their respective morphologies probably reflect the presence of two taxonomic validity will need direct comparisons main abelisauroid clades, informally large among all specimens as well as new, more comple- abelisaurids (represented in the quarry by te discoveries. We keep the names Indosuchus, Indosuchus and Indosaurusj aiid smaller Indosaurus, Rajasaurus and Larnetasaurus noasaurids (represented in the quarry at least by pending new studies or more discoveries that might Laeuisuclzusj. Because Laeuisuchu.~is referred to clarify whether these taxa can be diagnosed on the as the Noasauridae, and since some metatarsals basis of autapomorphies. and pedal phalanges resemble these of the Although our knowledge on the anatomy and noasaurids Velocisaurus and Masiakasuurus, we phylogeny of Indian abelisauroids is far from tentatively associate the slender foot bones with being settled, it seems clear that these theropods Noasauridae or Laeuisuchus. Caudals with well were numerically dominant and taxonomically developed transverse processes are also diverse in the Late Cretaceous of India, as tentatively referred to Noasauridae. Instead, documented in different fossil sites of the Lameta distal caudals devoid of transverse processes and Formation (e. g., Bara Simia, Rahiolij, a view that is in concert with the information available from having short prezygapophyses, and short and other Gondwanan localities of Late Cretaceous robust pedal digits, more probably belong to age (Madagascar, Patagonia). abelisaurids. To this list of robust abelisaurid bones, we add all the hindlimb bones (femora, ACILWOWLEDGEMENTS tibiae, metatarsals, phaianges) of large size as well as those of small size but with robust proportions. The senior author received support from Agen- With respect to the sacral vertebrae, the slender cia Nacional de Promoci6n Cientifica y Thcnica, and rod-like kind is documented so far in The National Geographic Society, CONICEX, Nouas et al.: Cretaceous theropods from Indza 99 FundaciCin Antorchas, and The Jurassic Earth Sci. 30: 2037-2081. Foundation. Thanks to Ashok Sahni for the Holtz, TR. 2000. A new phylogeny of the carnivorous support received during the trip of FEN to India. dinosaurs. GNA 15: 5-61. Our gratitude lo the personnel of the Geological Hoene, I?V 1914. Dasnaturiiche System der Saurischia. Survey of India (Kolkata) for allowingus to study Zer~trfur Min., Geol. Palaront., B 1914: 154.158, the tberopod collection, and I? Ghosh for his - 1920. Stammesgeschichtliche Ergebnisse einiger information on Indian geology. M. Carrano arid S, Untersuehungen an Trias-Repcilien. Zeits, fur Chatterjee offered vainable observations on the Induk. Absl. 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Science 280: 1048-1051. Trans. Royal Soe. London, 248 (744):53-134. Stovall, JW & UTL. Langston. 1950.Acroear~thosaurus Weiles, S. P 1984.Dilophosaurus welherilli (Dinosauria, atohensis, a new genus and species of Lower Theropodai. Osteology and comparisons. Cretaceous Theropoda from Oklahoma, Am. iMidl. Palaeontographica A 185 (4-6):85.180. Nat. 43:696-728 Wilson, J.A., PC. Sereno, S. Srivastava, D.K. Bhatt, A. Sues, H.D. & I? l'aquet. 1979. A pachycephalosaurid Khosla & A. Sahni. 2003. A new sihelisaurid dinosaur from Madagascar and a Laurasia- (Dinosauria, Theropoda) from the Lameta Gondwanaland connection in the Cretaceous. Formaiton (Cretaceous, Maastrichtian) of India. Nati~rt.279: 633-635. Contrib. Mus. Paleont., Uniu. Michigan 31(1):1-42. Recihido: 1.0-XI-2003 Aceptado: 19-11-2004 Appendix. List of theropod specimens described by Huene and Matley (19331, indicating current determination of bones and their respective taxonomic interpretation. Taxa Collection Original Material Taxonomic Current (original number illustration (original interpretation determination description) interpretation) (this paper) of bones Indosudzus K201350 PI. IX, fig. 1 Skull-roof lndosuc1,us raplorius Skull-roof raptoriris Indosuchus K271685 Pi. IX, fig. 2 Skull-roof Indosuchus raptorius Skull-roof ~t~nus I~~dosaurus K271565 PI. IX, fig. 3 Cast of braincavity Indosaurus rnatlfyi Cast of braincavity rnatleyi Indosai~rus K271565 PI, IX, fig. 4 Middle part of skull Indosaurus matleyi Middle part of skull nratleyi Indosuchus K271690 None Skull Indosuchus raptorius Sku11 rnptorius Indosuchus AMNH None Premaxilla Abelisauridae indot. Premaxilla rnplorius 1753 Irrdosuchus NANH None Left maxilla Abelisauridae indet. Left maxilla raptorrus 1955 Indosucl~us AMNIl None Right dentary Abelisauridae indet. Right deiitary raptorius 1960 Ir~dvsuchus AMNH None Caudal vertebra Abelisauridae indet Caudal vertebra raptorius 1960 Indosuchus AMNH None Caudal vertebra Abeiisauroidea indot Caudal vertebra raptorilia 1957 Indosuclius kMNH None Caudal vertebra Ahelisauroidea indet Caudal vertebra raptorius 1958 Indosuchus AVNH None Proximal caudal Abelisauroidea indet Proximal caudal raptorius no number vertebrae Ailosaurid K271548 PI. X, fig. 2 Right maxilla Abelisauridao indot. Right maxilla Allosaurid K271628 PI. X, fig. 3 Basioccipital Abolisauridae indet. Basioccipital Allosaurid K271577 PI. X, fig. 4 "Lacrimal" Abelisauridae indet. Left jugal Ailosaurid K201619 PI. XI, fig. 1 Right premaxilla Ahelisauridae indet. Premaxilla Aliosaurid K271710 PI. XI, fig. 2 Left premaxilia Abelisauridae indet. Yremaxilla Novas et aE.. Cretaceous therop,gds from India Allosaurid Right maxilla Right maxilla .Ulosaurid PI. XI, fig. 3 Left lnaxilia Left Maxilla Allo~aurid P1. XI. fie. 4 "Transverse bone" Indet, bane Nlosaiirid Pi. xi; fig. 5 "Right Lacrimal" Indet. bane Allosaurid 1'1. XI, fig. 6 Basioccipital Basioccipital Nosayid Pi. XI, fig. 7 Right Postorbital Abelisauridae indct. Right Jugal Allosaurid Pi. XII. fie 1 Right Dental Abelisauridae indet. Dentary Allasaurid PI. x11; fig. 2 Right Dental Abbelisauridae indet. Dentary Allosaurid Pi. XII, fig. 3 Left Articular Abelisauridao indet. Left surangular Allosaurid PI. XII, fig. 4 Left Articular Abelisauridae indet Articular Allosaurid PI. XIil, fig. 1 Tooth Indet. Tooth Allosaiirid P1. XIII, fig. 2 Tooth Indet. Tooth Allosaurid P1. XIII. fir. 3 Tooth Indot. Tooth Allosaurid PI. x111; fig. 4 Tooth Indet. Tooth Allosaurid Pi. XIII. fir. 5 Tooth Indet. Toorh Allosaurid Pi. XIII., fi;.- 6 Tooth Indet. Tooth Allosaurid Pi. XIII, fig. 7 Toath Indet. Tooth Allasaurid PI. XIII, fig. 8 Tooth Indet. Toath Nlosaurid P1. XIII. fie. 9 Tooth Indet. Toath Coelurosaiirian Pi. x111: fig.10 Tooth Indet. Toath Carnosnuria Pi. XIII, Fig.11 Dorsal vertebra Abelisauridae indet. Sacral 1 Allosaurid Pi. XIV fie. 1 Cervical vertebra Abelisauridae indet. Dorsal vertebra Allosaurid PI. XI\! fig. 2 Cervical vertebra Theropoda indet. Cervical vertebra Allosaurid PI. XIv, fip. 3 Sacral vertebra Abelisauridae indet. Sacral Aliosaurid PI. XI? fig. 4 Sacral vertebra Abelisauridae indet. Sacral Caelurosaurid P!..XXIII, fig. 1 Sacral vertebra Abelisauroidea Sacral vertebra Allosaurid Pi. Xv, fig. 1 Caudal vertebra Abeiisauridae indot. Caudal vertebra Allosaurid Pi. XV, fig. 2 Caudal vertebra Abelisauridae indet. Caudal vertebra Allosaurid Pi. XV, Fig. 3 Caudal vertebra Abelisauridae indot. Caudal vertebra Allosaurid Pi. Xv, fis. 4 Caudal vertebra Abelisauridae indet. Caudal vertebra Allosaurid PI xV, fi.5 Caudal veptebra Abelisauridae indet. Caudal vertebra Allosaurid Pi. XC: fig. 6 Caudal vertebra Caudal vertebra Allosaurid PI. XVI, fig. 1 Hemal arches Hemal arches Allasaurid PI. XVI, fig. 2 Hemal arches I-Iexnal arches Allosaurid PI. XVI, fig. 3 Hemal arches Hemal arches Allosaurid Pi. XVI, fig. 5 Hemal arches Hemal arches Allosaurid Pi. XVI, fi~.4 Hemal arches Hemal arches Allosaurid P1. XVI, fig. 6 Hemal arches Ilemal arches Allosaurid Pi. XVI. fir. 7 Left ischium Ischium Allosaurid Pi. XVI; fig. 6 Left ischium Abelisauridae indet. Ischium Allosaurid PI. XVI, fig. 9 Right femur Abeiisauridae indet. Right femur Aliosaurid Pi. XVI, fig. 10 Left femur Abeiisauridae indet. Left femur Allosaurid PI. XVII, fig.1 Right femur Abelisauridae indet. Right femur Aliusaurid PI. XVII, fig.2 Right femur Abelisauridae indet. Right fomur Allosaurid PI. XVIII, fig.2 Right tibia Abelisauridae indet. Right tibia Allosaurid PI. XVIII, fig.3 Left fibula Abelisauridae indot. Fibula Allosaurid Pl. XIX, fig. 1 Right astragalus Abelisauridae indet. Left quadrate Allosaurid PI. XIX, fig. 2 Caicaneum Indet. Indet, bone Allosaurid Pl. XiX, fig. 3 Right Mtt 2 Abelisauroidea indet. Right Mtt 2 Allosaurid Pi. XIX, fig. 4 Left Mtt 3 Abelisauroidea indet. ~e?tMtt 3 Allosaurid Pi. XIX. fie. 5 Rirht Mtt 4 Abelisauroidea indet. Right Mtt 4 Allosaurid P1 XIX; fig. 6 Right Mtt 4 Abelisauroidea indet. Right- Mtt 4 Allosaurid P1. XIX, fig. 7 Phalanx digit l Abelisauridae indet. Pedal phalanx 1 of digit I1 Allasaurid PI. XIX, fig. 8 Foot phalanx Abeiisauridae indet Pedal phalanx 1 of digit 111. Allosaurid PI. XIX, fig. 9 Foot phalanx Abelisauridae indet Pedal phalanx 2 of digit 111. Aliosaurid PI. XIX, fig. 10 Claw of foot Abelisauridae indet. Ungual phaianx Allosaurid PI. XIX, fie. 11 Claw of foot Abelisauridae indet. Ungual phalanx Allosaurid Claw of foot Abelisauridae indet. Ungual phalanx Allosaurid PI XIX; fig. 13 Claw of foot Abelisauridae indet. Ungual phalanx Compsosuchus PI. XX, fig. 1 1, 2 cerv~cal Abelisauroidea nomen Axis solus vertebrae dubiurn 102 Reuista del Museo Argentino de Ciencias Naturales, n. s. 6 (I), 2004 Laeuisucl~us K201613 Pi. XX, fig. 2 Cervical vertebra Noasauridae Cervical vertebra indieus Laeuisuchus K271696 PI. XX, fig. 3 Cervical vertebra Noasauridae Cervical vertebra indieus Laeuisuchus K201614 Pi. XX, fig. 4 Cervical vertebra Noasauridae Cervical vertebra indicus Laeuisuei~us K271588 PI. XX, fig. 5 Dorsal vortebra Noasauridae Dorsal vertebra indicus Jubbulpnria K201612 PI. XX, fig. 6 Dorsal vertebra Abelisauroidea Caudal vertebra tenuis Jubbulpuria K271614 PI. XX, fig. 7 Dorsal vertebra Abelisauroidea Caudal vertebra tcnuis Omithonzirnoi - K271600 1'1. XX, fig. 8 Dorsal vertebra Abelisauroidea Caudal vertebra des mobilis Omitl~onzin~oi- KZO1GlO PI. XX, fig. 9 Dorsal vertebra Abelisauroidea Caudal vertebra des rrtobilis Orizillzorr~imoi- K201614 P1. XX, fig. 10 Dorsal vortebra Abelisauroidea Caudal vertebra des mobilia I3 Coelurosaurian K271703 Pi. XX, fig. 11 'Pooth lndet. Tooth Omithumimoida? K271541 PI. XXI, fig. 1 Dorsal vertebra Abelisauroidea Caudal vertebra barasirrilensis OrnithonzimorrEesPK271531 Pi. XYI, fig. 2 Dorsal vertebra Abelisauroidea Caudal vertebra barasimlensis Ornithomirnoides?K271604 P1. XXI, fig. 3 Dorsal vertebra Abelisauroidea Caudal vertebra barasirnlensis On~ithornimoide~?K271682None Dorsal vertebra .4belisauroidea Caudal vertebra barasiriil~nsis Coeluroides K271574 PI. XXI, fig. 4 Dorsal vertebra At>elisauroidea Caudal vertebra largus Coeluroides K271595 PI. XXI, fig. 5 Dorsal vertebra ilbelisauroidea Caudal vertebra largus Coeluroides K271562 PI. XXI, fig. 6 Dorsal vertebra Abelisauroidea Caudal vertebra largus D~,ptoptosauroi 11201334 Pi. XXII, fig. 1 Dorsal vertebra Abclisauroidea Caudal vertebra des grandis Dryptosauroi K201609 PI. XYII, fig. 2 Dorsal vertebra Abelisauroidea Caudal vertebra des grandis Dryptusauroi K271549 Pi. XXII, fig. 3 Dorsal vcrtebra Abelisvuroidea Caudal vertebra des grandis Dry~tomaiLrdes K27160l PI. XXII, fig. 4 Dorsal vertebra Abelisauroidea Caudal vertebra grandis. Coelurosaurid K271571 PI. XXIII, fig.1 Sacral vcrtebra Abelisauridae Sacral vertebra Coeluroaaurid 11271532 R.XXIII, fig.2 Caudal vertebra Abelisauridae Caudal vertcbra Coelurosaurid K271589 Pi. XXIII, fig.3 Caudal vertebra Abeiisauroidea Caudal vertebra Coelurosaurid X271599 PI. XXIIl, fig.4 Caudal vertebra Abelisauroidea Caudal vertebra Coeiurosaurid K271587 1'1. XXIII, fig.5 Caudal vertebra Abelisauroidea Caudal vertebra Coelurosaurid K27155Y PI. XXIII, fig.6 Right iiiurn lndet. Indet. Cadurosaurid K271526 PI. XXIII, fig.7 Right tibia Indet. Indet. Coelurosaurid K271669 PI. XXIY fig.l Distal tibia Indet. Indet. Caelurosnurid 11271665 PI. XXIY fig.2 Metatarsal I11 Abelisauroidea indet. Metatarsal III Coelurosaurid K271697 +Pi. .MY fig.3 Metatarsal 11 Abbelisaudaidoa indet. Metatarsal 111 K271681 Coelurosaurid K271666 Pi. XXIY fig.4 Distal metacarpal Abelisauroidea indet. Distal metatarsd iV Coelurosaurid K271667 Pi. XXIY fig.5 Distal metacarpal Abelisauroidea indet. Distal metatarsal I1 Coelurosaurid K201337C Pi. XXIY fig6 Metatarsal I Abelisauroidea indet. Distal metatarsal IV Coelurosauvid K201626B PI. XXIV fig.7 Firat manual Abelisauroidea indet. Pedal phalanx 1 phalanx digit 1V Coelurosaurid K271657 PI. XXIY fig8 Phalanx Indet. Phalanx Coelurosaurid K271637 PI. XXIY fig.9 Phalanx foot IV toe Abelisauraidea indet. Pedal phalanx 4 of digit IV Coelurosaurid K271638 PI. XXIY fig.10 Phalanx A1,elisauroidea indet. Pcdal phalanx 3 of digit IV Caelurosaurid K271645 PI. XXIY fig.l.1 Phalanx Indet. Indet. Nooas et a1 Cretaceous theropods from iizd~a 103 Coelurasaurid K271642 PI. XXI'Y: fig.12 Manual phalanx Abelisauroidea indet. Pedal phalanx 1of dint IV Coelurosaurid K271648 PI. XXIV; fig.13 Manual phalanx Abelisauroidea indet. Coelurosaurid K271646 PI. XXIY fig14 Phalanx of digit IV Abelisauridoidea indet. u Coelurdsaurid K271647 Pi. XXIY fig.15 Phalanx of' digit IV Abolisauroidea indet. I'edal phalanx 4 of digit IV Coelurosaurid K201337B PI. XXIY fig.16 Phalanx Abelisauroidea indet. Pedal phalanx 2? of digit I1 Coelurosaurid 11271524 PI. XXIY fig.17 Phalanx Abelisauroidea indot. Pedal phalanx 1of dieit 11 Coelurosaurid K271644 PI. XX[Y fig.18 Phalanx Abelisauroidea indet. I'ednl phalanx 2 of dieit. 111. Caelumsaurid 11271632 Pi. XWI: fig19 Pedal unguai Abeiisanroidea indet. Pedal ungual phalanx phalanx Coelurosaurid K27162Y PI. XXIY fig.20 Pedal ungual Abelisvuroidea indet. Pedal ungiial phalanx phalanx Coelwosaurid K201626 PI. XXIY fig.?. Coelurosaurian Abelisauroidea indet. Pedal phalanx 1of manual phalanx digit IV