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Guaibasaurus candelariensis (Dinosauria, Saurischia) and the early origin of avian-like resting posture
Federico Agnolin & Agustn G. Martinelli Available online: 02 Mar 2012

To cite this article: Federico Agnolin & Agustn G. Martinelli (2012): Guaibasaurus candelariensis (Dinosauria, Saurischia) and the early origin of avian-like resting posture, Alcheringa: An Australasian Journal of Palaeontology, DOI:10.1080/03115518.2012.634203 To link to this article: http://dx.doi.org/10.1080/03115518.2012.634203

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Guaibasaurus candelariensis (Dinosauria, Saurischia) and the early origin of avian-like resting posture
FEDERICO AGNOLIN and AGUSTIN G. MARTINELLI
AGNOLIN, F. & MARTINELLI, A.G., iFirst article. Guaibasaurus candelariensis (Dinosauria, Saurischia) and the early origin of avian-like resting posture. Alcheringa, 15. ISSN 0311-5518. A specimen of the basal saurisquian Guaibasaurus candelariensis Bonaparte, Ferigolo and Ribeiro (UFRGS-PV-0725-T) from the Faxinal do Soturno locality, Rio Grande do Sul State, Brazil (Caturrita Formation; Late Triassic) lacks any sign of post-mortem transport and burial deformation, and exhibits features (exed forelimbs, folded hindlimbs under the body and curved neck) that indicate a typical avian-like resting position. The presence in Guaibasaurus of an avian-like resting posture and related physiological implications would extend this unique trait, previously considered restricted to derived maniraptoran theropods, to the base of the Theropoda (or even Saurischia) clade. n de Historia Natural Felix de Azara. Departamento de Ciencias Naturales y Federico Agnolin [fedeagnolin@yahoo.com.ar], Fundacio nides. Valentn Virasoro 732 (C1405BDB), Buenos Aires, Argentina, and Seccio n Paleontologa Antropologa, CEBBADUniversidad Maimo ngel Gallardo 470 (C1405BDB), Buenos Aires, Argentina. de Vertebrados, Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Av. A Agustn G. Martinelli, Centro Paleontologico de Pesquisas Llewellyn Ivor Price, Museu dos Dinossauros, CCCP-UFTM, BR-262, km 784, Peiropolis, Uberaba, Minas Gerais, Brazil. Received 1.6.2011; revised 28.9.2011; accepted 11.10.2011. Key words: Guaibasaurus, basal Dinosauria, avian-like resting posture, homeothermy, Triassic, Brazil.

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GUAIBASAURUS CANDELARIENSIS was originally described by Bonaparte et al. (1999) as a very primitive saurischian dinosaur from the Late Triassic of southern Brazil. The holotype, an incomplete but highly informative skeleton, and the paratype speci ria, Rio mens were recovered near the city of Candela Grande do Sul State, southern Brazil (Bonaparte et al. 1999). Later, a partial articulated skeleton clearly referable to Guaibasaurus candelariensis was described by Bonaparte et al. (2007), sourced from Linha Sa o Luiz, near the city of Faxinal do Soturno (Rio Grande do Sul State, southern Brazil; Fig. 1). Finally, a still unprepared specimen of the same species derived from this latter locality was briey mentioned by Langer et al. (2011). All known specimens were collected from outcrops of the Caturrita Formation (CarnianNorian, Late Triassic; Langer et al. 2007b, Bonaparte et al. 2010) attributable to the Riograndia Biozone (Abdala & Ribeiro 2010, Soares et al. 2011). The phylogenetic relationships of Guaibasaurus candelariensis are in a state of ux. The genus was originally described as a very basal saurischian, included in the monotypic family Guaibasauridae, with a combination of features that positioned it
ISSN 0311-5518 (print)/ISSN 1752-0754 (online) 2012 Association of Australasian Palaeontologists http://dx.doi.org/10.1080/03115518.2012.634203

phylogenetically within saurischians, but basal to the Sauropodomorpha-Theropoda split (Bonaparte et al. 1999). Based on the description of the specimen from Faxinal do Soturno, Bonaparte et al. (2007) found resemblances between G. candelariensis and Saturnalia tupiniquim (Carnian, Santa Maria Formation, Rio Grande do Sul State, Brazil; e.g., Langer et al. 1999). The authors placed both species within Guaibasauridae, reinforcing the familys assignment to basal saurischians on the stem lineage to Sauropodomorpha or prior to the divergence of Sauropodomorpha Theropoda (Bonaparte et al. 2010). Cladistic analyses have favoured inclusion of Guaibasaurus either within the basal sauropodomorphans (Ezcurra 2010, Ezcurra & Novas 2009) or as having theropod anities (Langer 2004, Langer et al. 2007a, Yates 2007a, 2007b, Bittencourt & Kellner 2009). Finally, the most recent compelling work on Guaibasaurus candelariensis undertaken by Langer et al. (2011), which included a careful re-description of the available specimens and an extensive character analysis, placed this species within the basal Theropoda, and this assignment is provisionally followed here. This study aims to evaluate the posture of the specimen from Faxinal do Soturno referred to Guaibasaurus by Bonaparte et al. (2007, 2010; see also Langer et al. 2011; Fig. 1) and interpret its palaeophysiological and palaeoecological implications.

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Material
Specimen UFRGS-PV-0725-T (Vertebrate Paleontological Collection, Instituto de Geocie ncias, Universidade Federal do R o Grande do Sul, Porto Alegre, Brazil) referable to Guaibasaurus candelariensis (Bonaparte et al. 2007, 2010; see also Langer et al. 2011) is analyzed (Fig. 1). The specimen was collected in an abandoned quarry 1.5 km northwest of the centre of Faxinal do Soturno city, R o Grande do Sul, Brasil. This quarry is near Santo Antonio village (298340 0800 S, 538300 2300 W), in the Linha Sa o Luis locality (Bonaparte et al. 2010). The specimen was collected from ne-grained red beds of the Caturrita Formation, Mammaliamorpha Biozone (Carnian, Upper Triassic; Bonaparte et al. 2010).

Results
The fossil consists of a nearly complete skeleton lacking information only on some forelimb elements, the skull and most of its neck (Bonaparte et al. 2007). The specimen is relatively intact, although the preservation of each individual bone is aected by weathering (when discovered, most of the skeleton was very close to the weathered surface) and abrasive preparation techniques. The neck and skull are not preserved, very likely owing to recent erosive loss. The specimen lacks any sign of post-mortem transport and burial deformation, most of the bones being preserved in tight articulation (see also Langer et al. 2011). The forelimbs are partially exed, with the right humerus extending posterolaterally and the radius-ulna anterolaterally oriented; the manus is medially oriented (Langer et al. 2011). The hindlimbs are exed under the trunk and pelvic girdle, thus the body sits on its folded legs. The right hindlimb is laterally displaced to a slight extent and the femoral head does not perfectly match the acetabular glenoid. Although incompletely preserved, the base of the neck (last cervical vertebra) and the anteriormost portion of the dorsal vertebral column are slightly curved to the left side, suggesting that the neck was probably entirely curved in the same direction.

Discussion
Xu & Norell (2004) recently proposed that the available specimens of the Cretaceous, derived, bird-like troodontid theropod Mei long exhibited an avian-like resting posture. In this derived dinosaur, this posture is expressed by the body resting on its symmetrically folded hindlimbs, the forelimbs verti-

cally exed and laterally extended with the elbows slightly displaced laterally, and the neck curved posterolaterally to the left side of the body (Xu & Norell 2004). These authors interpreted this posture to be identical to the tuck-in resting posture of living birds. This evidence led Xu & Norell (2004) to argue that this kind of resting was also present in non-avian theropods of the lineage leading to birds (see also Burt et al. 2007). Another troodontid was also found by Russell & Dong (1993) with its forelimbs clearly exed under its body, essentially identical in posture to that of Mei long. This resting posture, once thought to be exclusive to Aves (Wing 1956), has also been reported in some other non-avian derived coelurosaurian theropods, including alvarezsaurids and oviraptorans (Nesbitt et al. 2011). However, such avian-like resting posture has not been reported previously in non-coelurosaurian dinosaurs. The Guaibasaurus specimen UFRGS-PV-0725-T (Fig. 1) bears all the traits typical of an avian-like resting position, including exed forelimbs, folded hindlimbs under the body and curved neck. In fact, Bonaparte et al. (2007, 2010) noted in their description that the fore- and hindlimbs were exed in this specimen, resembling the resting condition of a living chicken. The occurrence of these features in Guaibasaurus, a supposed basal theropod (see Langer et al. 2011), clearly indicates that a typical avian-like resting position was present at least at the base of Theropoda (or even Saurischia if other systematic interpretations are followed). A few pre-dinosaurian taxa also appear to have this resting pattern. The only available and poorly preserved specimen of Saltopus elginensis (Fig. 2) comes from the yellow sandstones of the Lossiemouth Sandstone (Late Triassic) of Morayshire, northeastern Scotland (see Benton & Walker, 2011 for a summary). Although Saltopus was originally considered to be a basal dinosaur (e.g., Huene 1910, Benton & Walker 1985), it was recently re-interpreted as a basal dinosauriform (Benton & Walker 2011). Nonetheless, its poor preservation leaves its phylogenetic placement weakly supported. As in Guaibasaurus, the Saltopus holotype has exed forelimbs and folded hindlimbs near the body, and it is probable that its posture was avian-like (sensu Xu & Norell 2004). Regrettably, the poor preservation of the only available specimen precludes conrmation of the position of the bones and determination of its posture with certainty. Scleromochlus taylori Woodward, 1907 is a basal pre-dinosaurian ornithodiran of uncertain phylogenetic position (see Benton, 1999). All available

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RESTING POSTURE IN ANCIENT DINOSAUR

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Fig. 1. A, Partial skeleton (UFRGS PV0725T) of Guaibasaurus candelariensis Bonaparte, Ferigolo & Ribeiro, 1999 (Late Triassic; southern Brazil) in dorsal view (modied from Langer et al. 2011). B, Detail of left forelimb. C, Detail of left hindlind. Scale bar 10 cm. Abbreviations: a astragalus; c calcaneum; d pedal digit; f femur; bula; h humerus; il ilium; mc metacarpal; r radius; t tibia; tv trunk vertebra. Numbers refer to posititions.

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Fig. 2. Partial skeleton of the holotype of Saltopus elginensis Huene, 1910 (Late Triassic, northeastern Scotland) in dorsal view (modied from Benton & Walker, 2011). Scale bar 10 cm.

specimens consist of bone impressions in ne-grained rocks from the Lossiemouth Sandstone (Late Triassic) of Morayshire, northeastern Scotland (see Benton & Walker, 2011). Although most specimens are dicult to interpret, the holotype of Scleromochlus taylori is relatively well preserved. This specimen shows exed forelimbs, with slight lateral orientation of the wrists, folded hindlimbs and a laterally curved neck, essentially matching an avianlike resting position. On this basis, both Scleromochlus and Saltopus reveal that avian-like posture was probably acquired prior to the origin of dinosaurs. Regrettably, the poor preservation of the available specimens precludes resolution of this matter. Living tetrapods have a broad array of resting postures, but only birds and some mammals rest with their hindlimbs folded (Xu & Norell 2004). In birds, this resting position, together with the exure of the neck to the sides of the body, is considered to be related to heat conservation (Hill et al. 1980, Marsh & Dawson 1989). Consequently, Xu & Norell (2004) considered that this behaviour in dinosaurs was also likely related to physiological/thermal status and that the avian-like resting posture may be a signature of homeothermic tetrapods. The presence of an avian-like resting posture in Guaibasaurus and its related physiological implications would extend this unique behaviour to early saurischians rather than being restricted to derived maniraptoran theropods. The appearance of this behaviour in bipedal dinosaurs by the early Norian (Late Triassic) is coeval with an increase in dinosaur diversity (e.g., Langer et al. 2010). The homeothermic versus ectothermic status of dinosaurs remains hotly debated: see, for example, the contrasting interpreta-

tions summarized by Chinsamy & Hillenius (2004) and Padian & Horner (2004). However, more evidence, such as that provided here, will be necessary to resolve the physiological status of basal dinosaurs.

Acknowledgements
We thank C. L. Schultz and M. B. Soares (UFRGS) for access to collections under their care. We thank three anonymous reviewers and the Editor S. McLoughlin for comments that improved the manuscript.

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