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Abelisaurid Pedal Unguals from the Late Cretaceous of India

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Abelisaurid Pedal Unguals from the Late Cretaceous of India AsociaciónPaleontológicaArgentina.PublicaciónEspecial7 ISSN0328-347X VIIInternationalSymposiumon MesozoicTerrestrialEcosystems:145-149Buenos. Aires,30-6-2001 AbeliSaurid Pedal unguals from the Late Cretaceous of India Fernando E. NOVAS!and Saswati BANDYOPADHYAY2 Abstraet. The ungual phalanges of theropod dinosaurs discovered in the Lameta Formation (Maastrichtian),centralIndia,exhibitdistinctivecharactersunknownin othertheropods.Hence,their tax- onomic identificationremained obscure since their original description in 1933.Recent discoveriesof abelisauridtheropods in Patagoniaindicate that the phalangesfromIndia belong to the foot of the~e~i- nosaurs. New informationon the foot skeletonof this group of CretaceousGondwanan predators lS m- cluded herein. Key words. Theropoda. Abelisauridae. Unguals. Cretaceous. Gondwana. IntroducTion Xenotarsosaurus; Bonaparte and Novas, 1985; Martínez et al., 1986; Bonaparte et al., 1990), India The Late Cretaceous (Maastrichtian) Lameta ilndosuchus; Chatterjee and Rudra, 1996), and Formation of central India has yielded dissociated el- Madagascar (Majungatholus; Sampson et al., 1998). ements of a variety of predatory dinosaurs. The ma- This considerable amount of information about terials were described by Frederich von Huene (in abelisaurid anatomy has served as a valuable guide Huene and Matley, 1933), who recognized nine to resolve confusing taxonomic aspects of the Indian theropod species from a fossil quarry at Bara Simla theropods. Hill (Madhya Pradesh, India). They are: lndosuchus Currently, lndosuchus, lndosaurus, and also rapiorius Huene, Indosaurus matleyi Huene, Compso- Laeoisuchue, are interpreted as members of suchus solus Huene, Laeoisuchus indicus Huene, Abelisauridae (Bonaparte and Novas, 1985; Molnar, [ubbulpuria tenuis Huene, Coeluroides largus Huene, 1990; Chatterjee and Rudra, 1996; Sampson et al., Dryptosauroides grandis Huene, Ornithomimoides mo- 1998; Novas and Bandyopadhyay, 1999). Moreover, bilis Huene, and Ornithomimoides (?) barasimlensis the controversial taxa "Compsosuchus", "Drypto- Huene. Huene also described a considerable amount sauroides", "Ornithomimoides", and "[ubbulpuria" are of theropod hindlimb bones (e.g., femora, tibiae, represented by vertebrae corresponding to different metatarsals, and pedal phalanges) that he could not regions of the neck and tail that also exhibit abeli- refer to any of these species. On the contrary, such saurid features (Novas and Bandyopadhyay, 1999). bones were vaguely interpreted by him as corre- Now, abelisaurid material from the Upper sponding TO "allosaurid" OR "coelurosaurid" thero- Cretaceous rocks from NW Patagonia settles the long pods (Huene and Matley, 1933). standing confusion about the theropod unguals from Phylogenetic relationships of the Indian India, and establishes its abelisaurid affinity. theropods has been unknown since then, and virtu- ally no progre ss has been attained about the system- atic allocation of the abundant post-cranial remains Abbreviations (e.g., Romer, 1956; Walker, 1964; Chatterjee, 1978; AMNH, American Museum of Natural History, Molnar, 1990;Norman, 1990).Fortunately, the last 15 New York;GSI, Geological Survey of India, Calcutta; years has been highly prolific in discoveries of MCA, Museo "Carlos Ameghino", Cipolletti; SUNY, abelisaurid theropods in Upper Cretaceous beds State University of New York, Stony Brook; YPM, from Patagonia (e.g., Abelisaurus, Carnoiaurus, YalePeabody Museum, New Haven. 'CONICET. Laboratory of Comparative Anatomy, Museo Argentino de Ciencias Naturales, Av. Angel Gallardo 470, Morphology of abelisaurid pedal unguals C:14050JR a\.lenQ~Aires, Argentiml, E-mail: [email protected]. from Patagonia and India 'Geological Studies Unit, Indian Statistical Institute, 203 Barraekpore Trunk Road, Calcutta 700 035, India. The new abelisaurid specimen (MCA 56) includes Email [email protected]. two pedal unguals (figure 1) apparently from digit ©AsociaciónPaleontológicaArgentina 0328-347X/01$00.00+50 146 F.E. Navas and S. Bandyopadhyay A E B e o Figure 1. A-E, digit IV ungual, presumably of the left foot, of an indeterminate abelisaurid (MCA 56) from the 'Río Limay Formation, northwestern Patagonia, in A, medial, B, lateral, C, dorsal, D, ventral, and E, proximal views. Scale bar: 2 cm. A.P.A. Publicación Especial 7, 2001 Abelisa urid pedal unguals from India 147 IV of the left and right feet. The unguals were found ,theropods, the ventral surface of the ungual is in association with other hindlimb elements and der- smooth and transversely convex (e.g., Allosaurus, mal skull bones that show a characteristic pattern of Sinraptor; Madsen, 1976; Currie and Zhao, 1993) or ornamentation (e.g., anastomosed grooves delimiting flat (e.g., Deinonychus YPM 5205). Interestingly, a mosaic pattern of raised areas) found in other Noasaurus, the purported sister group of Abeli- abelisaurids (e.g., Carnotaurus, Abelisaurus, Majun- sauridae (Novas, 1992, 1997)also has a ventrally ex- gatholus); distal caudal vertebrae were also collected, cavated ungual, although it is located in a more prox- the morphology of which fits well with that of imal position (Bonaparte and Powell, 1980). Majungatholus (specimen 93161-14 at SUNY) and Pedal unguals of indeterminate theropods from lndosuchus (AMNH 1957,1958). the Lameta Formation (Huene and Matley, 1933) The unguals, nearly 6.5 cm when complete, are were sorted into two subsets: "allosaurid" manual blunt and asymrnetrical. They are arched (presum- (specimen K27/ 633) and pedal unguals (specimens ably medially), with the lateral surface strongly con- K27/524, K27/537, K27/543, K27/551, K27/634, vex and the internal surface almost flat. As a result of K27/635, K27/636, and K20/399), and "coeluro- this asymmetry, the medial surface of the ungual is saurid" pedal unguals (e.g., K27/625, K27/629, also exposed dorsally. Both lateral and medial sur- K27/630, K27/631, and K27/632). We were able to faces have wide canals that show a distinct, proxi- access only K27/634, K27/632, K27/633, and mally bifurcated, "Y"-shaped pattem. The bifurca- K27/524, which are amenable for comparison with tion is at mid-length. The grooves are deeper on the the Patagonian unguals. lateral rather on the medial side of ungual. The "Y"- Ungual phalanx K27/634 (figures 2.A-D) is trans- shaped groove of the lateral surface bounds a con- versely wide and almost symmetrical, and may be spicuous, triangular bump. There is no proximoven- from the central toe (digit I1I).As in abelisaurid un- tral flexor tubercle. The ventral surface of the ungual, guals from Patagonia, it has deep furrows on both instead, is proximally flat, but has a narrow and deep sides that bifurcate at mid-length and bounding a groove centrally. The proximal articular contour is prominent bump. Also, the ventral surface is fur- triangular, and the proximodorsal lip is prominent rowed. Ungual phalanx K27/524 (figures 2.E-F; the and tongue-shaped. same collection number was originally used as well These unguals exhibit the following peculiar fea- for a non-ungual phalanx), was interpreted as be- tures distinguishing them from other theropods (e.g., longing to a "coelurosaurid" by Huene, but may be- Allosaurus, Sinrapior, Tyrannosauridae, Ornithomi- long to pedal digit IVbecause it resembles the ungual midae, Troodontidae, Dromaeosauridae, Alvarez- of this digit of Sinrapior (Currie and Zhao, 1993). sauridae): Ungual phalanx K27/632 (Huene and Matley, 1) Grooves bifurcated proximal1y (figure LB). 1933) has the same characters as in K27/634. Most tetanurines (e.g., Allosaurus AMNH 680, 5750; However, because ungual K27/ 632 is more com- Sinraptor; Currie and Zhao, 1993; Alectrosaurus pressed transversely and asymmetrical than the oth- AMNH 6554)have a simple groove along both later- er unguals described by Huene, it may correspond to al and medial surfaces. Although some coelurosauri- digit 1.It is the smallest of all the available unguals. an theropods have "Y"-shaped grooves (e.g., Also, it has a strongly keeled dorsal margin, in agree- Patagonykus; Novas, 1997;Troodon AMNH 2137),the ment with the first ungual of other theropods (e.g., bifurcation is more proximal than in abelisaurids. Sin raptor; Currie and Zhao, 1993). Moreover, current phylogenetic hypotheses (e.g., Phalanx K27/633, although notably compressed Novas, 1992;Sampson et al., 1998)depict abelisaurids transversely, has a conspicuous ventral furrow and as Cretaceous ceratosaurs, suggesting that bifurcated collateral bifurcated grooves, as the pedal unguals grooves on the pedal unguals were independently described above. Because K27/ 633 is nearly as large acquired in different theropod groups (e.g., as, albeit is transversely narrower, than the presumed Abelisauridae, Patagonykus, Troodon). ungual of digit III (K27/634; figures 2.A-D),we inter- 2) Presence of a rounded bump on the lateral pret K27/ 633 as belonging to digit 11.It is the deepest side of pedal unguals (figure LB). In abelisaurid un- of the available unguals, and resembles the ungual of guals the upper and lower branches of the bifurcated digit 11of Sinraptor (Currie and Zhao, 1993).Ungual lateral groove bound a triangular prominence, not K27/635 (Huene and Matley, 1933,pl. XIX,fig. 10)re- found in other theropods. In coelurosaurs with "Y"- sembles K27/ 633 in its high lateral profile and asym- shaped grooves, this region is less prominent or al- metry, although it is not dorsally keeled. most flat. Another ungual presumably belongs to digit IV. 3) Ventral surface excavated or
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