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Comparative Cranial Myology and Biomechanics of Plateosaurus and Camarasaurus and Evolution of the Sauropod Feeding Apparatus

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Comparative Cranial Myology and Biomechanics of Plateosaurus and Camarasaurus and Evolution of the Sauropod Feeding Apparatus Button, D. J. , Barrett, P., & Rayfield, E. (2016). Comparative cranial myology and biomechanics of Plateosaurus and Camarasaurus and evolution of the sauropod feeding apparatus. Palaeontology, 59(6), 887-913. https://doi.org/10.1111/pala.12266 Publisher's PDF, also known as Version of record License (if available): CC BY Link to published version (if available): 10.1111/pala.12266 Link to publication record in Explore Bristol Research PDF-document This is the final published version of the article (version of record). It first appeared online via Wiley at http://onlinelibrary.wiley.com/doi/10.1111/pala.12266/full. Please refer to any applicable terms of use of the publisher. University of Bristol - Explore Bristol Research General rights This document is made available in accordance with publisher policies. Please cite only the published version using the reference above. Full terms of use are available: http://www.bristol.ac.uk/pure/about/ebr-terms [Palaeontology, Vol. 59, Part 6, 2016, pp. 887–913] COMPARATIVE CRANIAL MYOLOGY AND BIOMECHANICS OF PLATEOSAURUS AND CAMARASAURUS AND EVOLUTION OF THE SAUROPOD FEEDING APPARATUS by DAVID J. BUTTON1,2,3,PAULM.BARRETT2 and EMILY J. RAYFIELD1 1School of Earth Sciences, University of Bristol, Life Sciences Building, 24 Tyndall Avenue, Bristol, BS8 1TQ, UK; e.rayfi[email protected] 2Department of Earth Sciences, The Natural History Museum, London, SW7 5DB, UK; [email protected] 3Current address: School of Geography, Earth and Environmental Sciences, University of Birmingham, Edgbaston, Birmingham, B15 2TT, UK; [email protected] Typescript received 24 May 2016; accepted in revised form 13 September 2016 Abstract: Sauropodomorpha represents an important shape changes to the mandible. The skull and mandible of group of Mesozoic megaherbivores, and includes the largest Camarasaurus are also ‘stronger’ under static biting. The Pla- terrestrial animals ever known. It was the first dinosaur teosaurus mandible appears to compromise structural effi- group to become abundant and widespread, and its members ciency and force transmission in order to maintain relatively formed a significant component of terrestrial ecosystems high jaw closure speed. This supports suggestions of faculta- from the Late Triassic until the end of the Cretaceous. Both tive omnivory in basal sauropodomorph taxa. The expanded of these factors have been explained by their adoption of mandibular symphysis and ‘lateral plates’ of sauropods each herbivory, but understanding the evolution of sauropodo- lead to greater overall craniomandibular robustness, and may morph feeding has been hampered by the scarcity of bio- have been especially important in accommodating forces mechanical studies. To address this, the jaw adductor related to asymmetric loading. The functional roles of these musculature of the basal sauropodomorph Plateosaurus and characters, and observed general shape changes in increasing the sauropod Camarasaurus have been reconstructed. These skull robustness, are consistent with hypotheses linking bulk- reconstructions provide boundary conditions for finite ele- herbivory with the origin of Sauropoda and the evolution of ment models to assess differences in structural performance gigantism. between the two taxa. Results demonstrate that Cama- rasaurus was capable of much greater bite forces than Pla- Key words: Sauropodomorpha, finite element analysis, her- teosaurus, due to greater relative adductor muscle mass and bivory, palaeoecology, biomechanics, virtual reconstruction. S AUROPODOMORPHA represents one of the most impor- behaviour influences all parts of an animal’s lifestyle, tant Mesozoic terrestrial herbivore groups, numbering including life history strategy, energetics, habitat prefer- more than 200 genera and occupying all continents (Galton ence and ecological role (e.g. Owen-Smith 1988; Chap- & Upchurch 2004; Upchurch et al. 2004; Weishampel et al. man & Reiss 1999; Schwenk 2000). In turn, information 2004; Mannion et al. 2011; Sander 2013). This clade on feeding in prehistoric animals can inform larger evolu- includes the sauropods, the largest terrestrial animals tionary patterns (Reisz & Sues 2000; Barrett & Rayfield known, with taxa commonly exceeding 20 tonnes in body 2006; Barrett 2014). Although the small size and superfi- mass (Klein et al. 2011; Benson et al. 2014) and the largest cial simplicity of the sauropod skull perplexed earlier reaching in the region of 60–70 tonnes (Mazzetta et al. authors (e.g. Hatcher 1901; Haas 1963; Colbert 1993) it is 2004; Lacovara et al. 2014; but see Bates et al. 2015). The now known that the sauropod skull represents a special- extreme nature of sauropodomorph biology makes explo- ized cropping tool (Christiansen 1999; Upchurch & Bar- ration of their functional anatomy and evolution imperative rett 2000; Sereno et al. 2007; Hummel & Clauss 2011; for understanding the constraints acting upon terrestrial life Sander et al. 2011; Young et al. 2012; Sander 2013). Mul- (Clauss 2011; Sander et al. 2011; Sander 2013). tiple studies of craniodental functional anatomy (e.g. Bar- The skull forms a fundamental link between an animal rett & Upchurch 1994, 2005, 2007; Calvo 1994; and its environment, especially through feeding. Feeding Christiansen 1999, 2000; Upchurch & Barrett 2000) have © 2016 The Authors. doi: 10.1111/pala.12266 887 Palaeontology published by John Wiley & Sons Ltd on behalf of The Palaeontological Association. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. 888 PALAEONTOLOGY, VOLUME 59 provided insights into sauropodomorph ecology and evo- Institutional abbreviations. AMNH, American Museum of Natu- lution, although quantitative biomechanical analyses (Pre- ral History, New York City, USA; CMNH, Carnegie Museum of uschoft & Witzel, 2005; Witzel et al. 2011; Young et al. Natural History, Pittsburgh, USA; DINO, Dinosaur National € 2012; Button et al. 2014) have been rarer. Monument, Vernal, USA; MB.R, Museum fur Naturkunde, The ability to eat plants has been implicated in the Berlin, Germany. early diversification of Sauropodomorpha (Galton 1985a; Barrett & Upchurch 2007; Barrett et al. 2011) and the MATERIAL AND METHOD evolution of the sauropod feeding apparatus (Barrett & Upchurch 2007; Upchurch et al. 2007; Sander & Clauss Virtual osteological reconstruction 2008; Sander et al. 2011; Sander 2013). Adoption of obli- gate high-fibre herbivory and bulk-feeding are seen as CT scans of MB.R.1937, an adult individual of Pla- integral drivers of sauropod gigantism (Barrett & teosaurus engelhardti and CMNH 11338, a juvenile speci- Upchurch 2007; Upchurch et al. 2007; Yates et al. 2010; men of Camarasaurus lentus, were provided by Lawrence Sander et al. 2011; Sander 2013; Barrett 2014; Benson M. Witmer, with permission to use the Plateosaurus data et al. 2014). Whereas basally branching ‘prosauropod’ from R. Goessling on behalf of the Humboldt Museum taxa appear to have remained relatively morphologically fur€ Naturkunde. These scan data were imported into conservative (Barrett & Upchurch 2007; Young & Larvan Avizo (v6.3.1, v7 and v8.0; Visualization Sciences Group; 2010) the base of Sauropoda is associated with the http://www.vsg3d.com). Complete reconstructions were appearance of numerous cranial traits hypothesized to made of Camarasaurus and Plateosaurus by assigning each have led to greater bite force, jaw processing power and cranial element of CMNH 11338 and MB.R.1937 to a increased robustness of the skull and mandible. These separate label within the Avizo segmentation editor. Each include the dorsoventral expansion of the mandibular specimen has suffered taphonomic deformation, with symphysis, broadening of the snout, the development of missing and warped elements. Retrodeformation was per- ‘lateral plates’ of bone inferred to have braced the teeth, formed using translation, transformation and mirroring and gross changes to cranial shape (Upchurch & Barrett tools within Avizo (see Lautenschlager et al. 2014; Cuff & 2000; Barrett & Upchurch 2007; Upchurch et al. 2007). Rayfield 2015) to produce undistorted and complete 3D However, this hypothesized functional shift has not been osteological reconstructions of Camarasaurus and Pla- tested quantitatively using biomechanical methods. teosaurus. See Button et al. (2016) for more information. Button et al. (2014) employed finite element analysis (FEA) of the crania of Diplodocus and Camarasaurus to investigate ecological phenomena, and Lautenschlager Virtual muscle reconstruction et al. (2016) used similar methods to compare the mechanical performance of Plateosaurus with that of other Jaw adductor muscles were reconstructed from a combi- herbivorous dinosaurs. Here, this methodology is nation of first-hand observations and examination of CT extended to the mandible of Camarasaurus. In addition, scans of MB.R.1937 and CMNH 11338. Muscle origina- the jaw adductor musculature of Plateosaurus and Cama- tion and insertion sites were identified on the basis of rasaurus, restored digitally from CT scan data, are phylogenetic bracketing (Witmer 1995) and osteological described in a detailed, comparative context for the first correlates (see Holliday 2009), with topological con- time. Plateosaurus is similar in craniodental morphology straints provided by other tissues and comparison to to other ‘prosauropods’ whereas Camarasaurus exhibits more distant outgroups (Lepidosauria
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