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Simulated Range of Motion and Hind Foot Posture of the Middle Jurassic Sauropod Rhoetosaurus

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Simulated Range of Motion and Hind Foot Posture of the Middle Jurassic Sauropod Rhoetosaurus Nair Jay (Orcid ID: 0000-0001-9218-8124) Jannel Andréas (Orcid ID: 0000-0002-6625-5693) 70 Jannel “Keep your feet on the ground”: Simulated range of motion and hind foot posture of sauropods — a case study of the Middle Jurassic Rhoetosaurus brownei LRH: ANDRÉAS JANNEL ET AL. RRH: BIOMECHANICS OF AN EARLY SAUROPOD PES Andréas Jannel , Jay P. Nair , Olga Panagiotopoulou , Anthony Romilio and Steven W. Salisbury Andréas Jannel. School of Biological Sciences, The University of Queensland, Brisbane, QLD 4072, Australia. E-mail: [email protected]; [email protected] Jay P. Nair. School of Biological Sciences, The University of Queensland, Brisbane, QLD 4072, Australia. E-mail: [email protected]; [email protected] Olga Panagiotopoulou. Monash Biomedicine Discovery Institute, Department of Anatomy and Developmental Biology, Monash University, Clayton, VIC 3800, Australia. E-mail: [email protected] This is the author manuscript accepted for publication and has undergone full peer review but has not been through the copyediting, typesetting, pagination and proofreading process, which may lead to differences between this version and the Version of Record. Please cite this article as doi: 10.1002/jmor.20989 This article is protected by copyright. All rights reserved. Jannel 2 Anthony Romilio. School of Biological Sciences, The University of Queensland, Brisbane, QLD 4072, Australia. E-mail: [email protected] Steven W. Salisbury. School of Biological Sciences, The University of Queensland, Brisbane, QLD 4072, Australia. E-mail: [email protected] This article is protected by copyright. All rights reserved. Jannel 3 ABSTRACT The biomechanics of the sauropod dinosaur pes is poorly understood, particularly among the earliest members of the group. To date, reasonably complete and articulated pedes in Early–Middle Jurassic sauropods are rare, limited to a handful of taxa. Of these, Rhoetosaurus brownei, from eastern Australia, is currently the only one from the Gondwanan Middle Jurassic that preserves an articulated pes. Using Rhoetosaurus brownei as a case exemplar, we assessed its paleobiomechanical capabilities and pedal posture. Physical and virtual manipulations of the pedal elements were undertaken to evaluate the range of motion between the pedal joints, under both bone-to-bone and cartilaginous scenarios. By employing the results as constraints, virtual reconstructions of all possible pedal postures were generated. We show that Rhoetosaurus brownei was capable of significant digital mobility at the osteological metatarsophalangeal and distal interphalangeal joints. We assume these movements would have been restricted by soft tissue in life but that their presence would have helped in the support of the animal. Further insights based on anatomy and theoretical mechanical constraints restricted the skeletal postures to a range encompassing digitigrade to subunguligrade stances. The approach was extended to additional sauropodomorph pedes, and some validation was provided via the bone data of an African elephant pes. Based on the resulting pedal configurations, the in-life plantar surface of the sauropod pes is inferred to extend caudally from the digits, with a soft tissue pad supporting the elevated metatarsus. The plantar pad is inferred to play a role in the reduction of biomechanical stresses, and to aid in support and locomotion. A pedal pad may have been a key biomechanical innovation in early sauropods, ultimately resulting in a functionally plantigrade pes, which may have arisen during the Early to Middle Jurassic. Further This article is protected by copyright. All rights reserved. Jannel 4 mechanical studies are ultimately required to permit validation of this long-standing hypothesis. KEYWORDS: Sauropoda; range of motion; posture; pes; biomechanics; Rhoetosaurus brownei This article is protected by copyright. All rights reserved. Jannel 5 1 INTRODUCTION Sauropods represent the heaviest terrestrial organisms known to have existed, exemplifying one of the most emblematic groups of dinosaurs. With body masses exceeding 40 tonnes (Bonaparte & Coria, 1993; Alexander, 1998; Mazzetta, Christiansen & Fariña, 2004; Benson et al., 2014; Lacovara et al., 2014), one might ask how these gigantic animals could move, let alone support themselves, without their bones failing under such heavy loads. Many studies have reconstructed sauropod locomotion using a variety of approaches, including comparative osteology (e.g., Hay, 1910; Christiansen, 1997; Carrano, 2005; Sander et al., 2011), paleoichnological analysis (e.g., Wilson & Carrano, 1999; Wilson, 2005), biomechanical investigations (e.g., Christian, Heinrich & Golder, 1999; Preuschoft et al., 2011; Klinkhamer et al., 2018), and/or computational techniques (e.g., Sellers et al., 2013). In sum, the majority of these studies proposed that sauropods likely had fairly restricted limb movements with reduced mobility at the main joints. These aforementioned studies have nonetheless focused entirely on analysis of the upper and middle long bone segments (the stylopodium and zeugopodium, respectively). For the vast majority of terrestrial vertebrates, however, the lower limb segments (the autopodium), comprising the manus and pes, represent the only skeletomuscular parts that interact with the substrate. Accordingly, the biomechanical properties of the autopodium not only operate for the support of the animal but also facilitate the transmission of forces into the more proximal parts of the limbs during locomotion (Floyd, 2014). So far, however, most locomotor studies have generalized sauropod autopodia to function as singular and static body segments, with a limited capacity for movement at the wrist and/or ankle joints (e.g., Sellers et al., 2013), if at This article is protected by copyright. All rights reserved. Jannel 6 all. Consequently, little is known about the exact biomechanical abilities of sauropod autopodia, and the extent to which these complex structures could withstand stresses and simultaneously propel these terrestrial giants. As often noted, the manus and pes of sauropods are morphologically divergent structures (Upchurch, Barrett & Dodson, 2004; Bonnan, 2005). While the manus comprises an elevated colonnade of metacarpals, the pes is generally proposed to contain more horizontally positioned metatarsals. However, the attribution of these configurations rests largely on the interpreter’s opinion of how the autopodial elements articulate, lacking rigorous support from biomechanical insights. A detailed understanding of the paleobiomechanical abilities of the autopodia in sauropods has therefore been deficient largely because certain parameters have been difficult to estimate with precision. These comprise ranges of motion (ROM) between ossifications, posture and autopodial-level kinematics. Consequently, the posture and biomechanics of the sauropod autopodia have often been open to subtly variant interpretations (e.g., Marsh, 1878; Hatcher, 1901; Paul, 1988; Gallup, 1989; McIntosh, 1990; Bonnan, 2005; Carrano, 2005; Wilson, 2005). In fact, published observations concerning movements within the sauropod pes remain limited to a handful of studies, mostly detailing motion at a few joints within the parasagittal plane. For instance, Hatcher (1901) observed that the unguals of Diplodocus carnegii (CM 94) were capable of substantial ‘vertical’ movements based on their articular morphology. Gallup (1989: p. 2) noticed that the articulation at the metatarsophalangeal joint of pes specimen FMNH 241-50 (now referred to Cedarosaurus weiskopfae, D'Emic, 2013) allowed for significant motion in the vertical plane. A similar appraisal was made by Bonnan (2005: This article is protected by copyright. All rights reserved. Jannel 7 p. 353), observing that the unique articular surface of the unguals at the most distal interphalangeal joints permit considerable flexion in Plateosaurus engelhardti, an early sauropodomorph. Whilst these observations provide preliminary indication for the flexibility at some joints within the sauropod pes, quantification of the associated pedal ROMs remains lacking (specifically in a three-dimensional space). Given this backdrop, we sought to investigate the paleobiomechanical capabilities and test the likelihood of potential postures in early sauropod pedes, using the Middle Jurassic Rhoetosaurus brownei as an exemplar. The scope of this study is to provide a preliminary constraint-based framework (Gatesy, Baker & Hutchinson, 2009) for the acquisition of more robust and uniform paleobiomechanical data in the reconstruction of sauropod autopodia, an approach fundamentally applicable to other extinct taxa. To evaluate this overarching aim, a set of hierarchical investigations was performed on the pes of Rhoetosaurus brownei. First, we measured the flexibility of each pedal joint using various ROM evaluations in a 3D context (physical or virtual; with cartilage or bone-to- bone), an aspect so far not yet quantified in the literature of sauropods. Second, each ROM analysis of Rhoetosaurus brownei was statistically compared between procedural methods (physical, virtual) and articular assumptions (bone-to-bone, cartilage) with ANOVA in order to determine the most exhaustive technique for paleobiomechanical analysis in extinct taxa. Third, identical ROM evaluations were performed with an African bush elephant (Loxodonta africana) pes to provide corroboration of our methods to modern taxa. As the largest living terrestrial animal,
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