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From the Late Triassic (Adamanian Placerias Quarry, Arizona, USA) and the Stratigraphic Range of the Drepanosauridae

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From the Late Triassic (Adamanian Placerias Quarry, Arizona, USA) and the Stratigraphic Range of the Drepanosauridae N. Jb. Geol. Paläont. Abh. 2009, vol. 252/3, p. 315–325, Stuttgart, June 2009, published online 2009 The oldest record of drepanosaurids (Reptilia, Diapsida) from the Late Triassic (Adamanian Placerias Quarry, Arizona, USA) and the stratigraphic range of the Drepanosauridae Silvio C. Renesto, Varese, Justin A. Spielmann and Spencer G. Lucas, Albuquerque With 3 figures RENESTO, S. C., SPIELMANN, J. A. & LUCAS, S. G. (2009): The oldest record of drepanosaurids (Reptilia, Diapsida) from the Late Triassic (Adamanian Placerias Quarry, Arizona, USA) and the stratigraphic range of the Drepanosauridae. – N. Jb. Geol. Paläont. Abh., 252: 315–325; Stuttgart. Abstract: Previous detailed descriptions of relatively complete drepanosaurid material make it possible to recognize isolated drepanosaurid elements from other localities. The identification of isolated elements from the USA and Great Britain extended the geographical distribution of the group and encouraged a review of Triassic collections for characteristic elements of this family. In this paper, isolated vertebrae previously described as problematic reptiles from the famous Placerias Quarry, near St. Johns, Arizona, USA are re-identified as drepanosaurids. These specimens represent the oldest occurrence of this family, which is earliest Adamanian. Key words: Drepanosauridae (Reptilia, Diapsida), Chinle Group, isolated vertebrae, biostratigraphy. eschweizerbartxxx author 1. Introduction the only two named drepanosaurid taxa from North America. Recently, Vallesaurus, also from the Italian Originally erected by OLSEN & SUES (1986) and first Calcare di Zorzino, was described (RENESTO & diagnosed by BERMAN & REISZ (1992), the Drepano- BINELLI 2006). Although these forms exhibit dif- sauridae are a family of diapsid reptiles with highly ferent sizes and slightly diverse body form, all share modified postcrania and are interpreted as arboreal characteristic features that unequivocally demonstrate (RENESTO 1994a, b, 2000; RENESTO & BINELLI 2006; their close relationships (RENESTO 1994a, b, 2000; though see COLBERT & OLSEN 2001 and BERMAN RENESTO & FRASER 2003). Nevertheless, there is & REISZ 1992 for differing interpretations). Dre- some uncertainty about the relationships of the panosaurus (PINNA, 1980) and Megalancosaurus Drepanosauridae with other diapsids. BERMAN & (CALZAVARA et al., 1980) from the Calcare di REISZ (1992) considered drepanosaurids to be lepido- Zorzino (Zorzino Limestone, Norian, Late Triassic) of sauromorphs, whereas RENESTO (1994a, b, 2000) Northern Italy were the first taxa of this family to be suggested archosauromorph affinities. Analyses by described. Dolabrosaurus from the Petrified Forest BENTON & ALLEN (1997) and DILKES (1998) lent Formation, Chinle Group, New Mexico (BERMAN & further support to the hypothesis that drepanosaurids REISZ 1992), and Hypuronector, from the Newark are archosauromorphs, specifically closely related to Supergroup (COLBERT & OLSEN 2001) (“deep-tailed Prolacertiformes, a hypothesis also proposed in the swimmer” of OLSEN 1980), were for many years recent revision of Prolacertiformes (Protorosauria) by DOI: 10.1127/0077-7749/2009/0252-0315 0077-7749/09/0252-0315 $ 2.75 © 2009 E. Schweizerbart’sche Verlagsbuchhandlung, D-70176 Stuttgart 316 S. C. Renesto et al. eschweizerbartxxx author Fig. 1. Paleogeographic map of the Late Triassic (~220 Ma) with drepanosaurid localities highlighted. Base map after WING & SUES (1992) RIEPPEL et al. (2003). The phylogeny of SENTER cognized isolated drepanosaurid cervical vertebrae (2004) suggested that the Drepanosauridae are closely among the disassociated vertebrate assemblages of the related to the Permian Coelurosauravidae. However, Upper Triassic fissure deposits at Cromhall Quarry, RENESTO & BINELLI (2006) demonstrated that England. The addition of the isolated elements from SENTER’s analysis reflected much confusion about the USA and Great Britain extended the geographical drepanosaurid anatomy, rather than helping to resolve distribution of the group and encouraged a review of relationships. RENESTO & BINELLI (2006), on the basis Triassic collections for characteristic elements of this of more complete material, proposed a new analysis, distinct family. Here we report some isolated verte- which placed the Drepanosauridae within Archosauro- brae, previously described as “Reptilia Problematica” morpha, close to Protorosauria and basal archosaurs. (KAYE & PADIAN 1994), from the large microverte- Based on detailed descriptions (RENESTO 1994a, b, brate collection of the famous Placerias Quarry, near 2000; COLBERT & OLSEN 2001; RENESTO & BINELLI St. Johns, Arizona, USA, and redescribe and identify 2006) of various drepanosaurs, known from relatively them as drepanosaurid specimen. complete material, it is now possible to recognize These newly recognized drepanosaurid specimens isolated drepanosaurid elements. Indeed, HARRIS & from the Placerias Quarry represent the oldest occur- DOWNS (2002) identified a characteristic drepano- rence of this family of archosauromorphs. Here, we saurid shoulder girdle from the famed Ghost Ranch also review the geographic and biostratigraphic range Coelophysis Quarry, and, more recently, RENESTO & of the Drepanosauridae and discuss the age of the FRASER (2003) and FRASER & RENESTO (2005) re- various drepanosaurid occurrences (Fig. 1). The oldest record of drepanosaurids from the Late Triassic 317 Institutional abbreviations: AUP, Aberdeen Uni- scribed by KAYE & PADIAN (1994); they are a centrum versity Palaeontology Collection, Aberdeen, Scotland; missing its neural arch and a neural arch with a medio- MNA, Museum of Northern Arizona, Flagstaff, Arizona, laterally-compressed, semicircular neural spine. We USA; MPUM, Museo Paleontologico Università di Milano, Dipartimento di Scienze della Terra dell’Università degli interpret these elements to be the centrum of a Studi di Milano, Milan, Italy. drepanosaurid and a cf. drepanosaurid neural arch. 2. Drepanosauridae from the Placerias 2.3. Description and identification Quarry MNA V3637 was originally described as a “posterior 2.1. Geological setting caudal centrum with a thin ridge on the ventral sur- face” by KAYE & PADIAN (1994: 186). It consists of an Near St. Johns in east-central Arizona, USA, the Pla- almost complete small vertebra, about 10 mm long cerias Quarry ranks among the richest fossiliferous (Fig. 2R-V). The centrum is low and elongate, ex- localities for Triassic continental vertebrates (CAMP & tending caudally well beyond the walls of the neural WELLES 1956; JACOBS & MURRY 1980; MURRY & arch, it is wider than high and also slightly constricted LONG, 1989; KAYE & PADIAN 1994; LUCAS et al. in the middle, so that its ventral margin has a concave 1997). Discovered in the 1930s, the quarry takes its outline. The cranial articular surface is concave, name from the abundance of bones of one of the last whereas the posterior surface is convex; thus the Triassic dicynodonts, Placerias, monographed by vertebra is procoelous. Because of the caudal ex- CAMP & WELLES (1956). The stratigraphically equi- tension of the centrum, the zygapophyseal articulation valent Downs Quarry is very close to the Placerias between pre- and postzygapophyses lies well forward Quarry (MURRY & JACOBS 1988). In the 1970s and of the intercentral articulation. The prezygapophyses 1980s, screenwashing yielded a diverse microverte- lie very close to the midline, and their articular facets brate fauna from the Placerias Quarry (MURRY & are oriented close to vertical, allowing good mobility JACOBS 1988; TANNENBAUM 1983; KAYE & PADIAN on the vertical plane, while hindering lateral mobility. 1994; HECKERT 2004), and other elements of its The neural arch is partially broken at the caudal end, macrovertebrate assemblage were described in the revealing a wide neural canal. The neural spine is eschweizerbartxxx author 1990s, including aetosaurs, phytosaurs and dinosaurs almost missing, but judging from the outline of the (LUCAS et al. 1992, 1997; LONG & MURRY 1995; preserved portion, it had to be low. Neither articular NESBITT et al. 2007). areas for ribs, nor can transverse processes be de- The quarry is located in the Bluewater Creek tected. Formation, near the base of the Upper Triassic Chinle MNAV3637 shows a combination of characters Group (LUCAS et al. 1997). It is of earliest Adamanian such as procoelous centra with large neural canal and age, based on the co-occurrence of Stagonolepis, end of the centrum protruding beyond postzigapo- Parasuchus (= Paleorhinus) and Rutiodon-grade physes, praezygapophyses forward protruding and phytosaurs (LUCAS et al. 1997, 2007). with subvertical articular facets, that correspond to that of the caudal vertebrae of the drepanosaurids especially of Vallesaurus in which, the haemal arches 2.2. Material examined are intercentral in position and not fused with the MNA V3637, originally identified by KAYE & PADIAN posteroventral end of the centrum as it occurs in (1994: 184, fig. 9.76) as a “procoelous posterior Drepanosaurus and Megalancosaurus. Thus, we inter- caudal centrum” of a reptile. We interpret it as a pret MNA V3637 as a posterior caudal vertebra of a posterior caudal vertebra of a drepanosaurid (Fig. drepanosaurid. 2R-V). Specimen MNA V3652 consists of three elements – MNAV3652 (Fig. 2A-Q) consists of three ele- a vertebra, a centrum and a neural arch. The vertebra ments, one originally described and figured by KAYE (Fig. 2A-E) was described as “a relatively large, & PADIAN (1994: 185, fig. 9.88)
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