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Shell Bone Histology of Solemydid Turtles (Stem Testudines): Palaeoecological Implications

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Shell Bone Histology of Solemydid Turtles (Stem Testudines): Palaeoecological Implications Zurich Open Repository and Archive University of Zurich Main Library Strickhofstrasse 39 CH-8057 Zurich www.zora.uzh.ch Year: 2014 Shell bone histology of solemydid turtles (stem Testudines): palaeoecological implications Scheyer, T M ; Pérez-García, A ; Murelaga, X DOI: https://doi.org/10.1007/s13127-014-0188-0 Posted at the Zurich Open Repository and Archive, University of Zurich ZORA URL: https://doi.org/10.5167/uzh-106169 Journal Article Originally published at: Scheyer, T M; Pérez-García, A; Murelaga, X (2014). Shell bone histology of solemydid turtles (stem Testudines): palaeoecological implications. Organisms Diversity Evolution:1-16. DOI: https://doi.org/10.1007/s13127-014-0188-0 Shell bone histology of solemydid turtles (stem Testudines): palaeoecological implications T. M. Scheyer, A. Pérez-García & X. Murelaga Organisms Diversity & Evolution ISSN 1439-6092 Volume 15 Number 1 Org Divers Evol (2015) 15:199-212 DOI 10.1007/s13127-014-0188-0 1 23 Your article is protected by copyright and all rights are held exclusively by Gesellschaft für Biologische Systematik. This e-offprint is for personal use only and shall not be self- archived in electronic repositories. If you wish to self-archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com”. 1 23 Author's personal copy Org Divers Evol (2015) 15:199–212 DOI 10.1007/s13127-014-0188-0 ORIGINAL ARTICLE Shell bone histology of solemydid turtles (stem Testudines): palaeoecological implications T. M. Scheyer & A. Pérez-García & X. Murelaga Received: 23 July 2014 /Accepted: 20 October 2014 /Published online: 6 November 2014 # Gesellschaft für Biologische Systematik 2014 Abstract Lately, solemydid turtles have been repeatedly re- gross anatomy of the fossil finds. Our results indicate that covered as stem Testudines, indicating that they belong to Solemydidae share unique histological features pertaining to neither one of the two major branches of crown turtles, the their strongly ornamented shell bones, which a) in cases allow Pancryptodira and Panpleurodira. Despite their wide temporal taxonomic identification of even small shell fragments and b) (Late Jurassic to Late Cretaceous) and spatial (North America unambiguously corroborate a terrestrial lifestyle of its mem- and Europe) distributions, solemydid turtles are not particu- bers. The latter further supports a terrestrial lifestyle prefer- larly well known, as exemplified by the fact that only a single ence of most representatives of the turtle stem. skull has been described for the whole group so far. Further- more, the palaeoecology of solemydid turtles is still contested Keywords Solemydidae . Stem turtles . Bone with hypotheses ranging from semi-aquatic to terrestrial life- microstructure . Palaeoecology . Bone ornamentation . styles. However, the habitat preference of stem Testudines, Mesozoic such as solemydids, is important to understand the evolution and early radiation of the turtle crown, which is primitively aquatic. Here we describe the shell bone microanatomy and Introduction histological microstructures of solemydid turtles using a broad sample of taxa of different ages and localities, as well as Solemydidae are a group of stem Testudines whose members review previous histological accounts, to elucidate the palaeo- are distributed over Europe and North America from the Late ecology of the group independent of the geological setting and Jurassic (Tithonian) to the Maastrichtian at the end of the Cretaceous (Joyce et al. 2011; Pérez-García et al. 2013). Anquetin (2012) and Joyce et al. (2011) both recovered some T. M. Scheyer (*) representatives of this Laurasian clade of turtles, the North Paläontologisches Institut und Museum, Universität Zürich, American Naomichelys speciosa Hay, 1908 and the European Karl-Schmid-Strasse 4, CH-8006 Zürich, Switzerland e-mail: [email protected] Helochelydra nopcsai Lapparent de Broin and Murelaga, 1999 respectively, among other stem representatives of A. Pérez-García Testudines in their phylogenetic analyses, instead of within Centro de Geologia, Faculdade de Ciências da Universidade de the turtle crown (e.g. Lapparent de Broin and Murelaga 1999; Lisboa (FCUL), Edificio C6, Campo Grande, 1749-016 Lisbon, Portugal Danilov 2005). Only recently, a nicely preserved skull was e-mail: [email protected] described for Helochelydra nopcsai (Joyce et al. 2011), whereas skulls of other representatives have been found (cf. A. Pérez-García Naomichelys in North America, cf. Solemys in Europe) but Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, C/ Senda del Rey, 9, 28040 Madrid, Spain hitherto remain unpublished. Other solemydid taxa are known only from shell and other postcranial remains. X. Murelaga One of the most obvious diagnostic features of solemydid Departamento de Estratigrafía y Paleontología, Facultad de Ciencia y turtles is their characteristic shell bone surface ornamentation Tecnología, Universidad del País Vasco/EHU, Apartado 644, 48080 Bilbao, Spain (Lapparent de Broin and Murelaga 1996; 1999; Joyce et al. e-mail: [email protected] 2011). This ornamentation consists of high or low tubercles, Author's personal copy 200 T.M. Scheyer et al. and can also include isolated ridges or ridge networks in that most fossils from the Cambridge greensand deposits are which the sinuous ridges frequently anastomose. Recently, reworked and likely of Albian age. Joyce et al. (2011) proposed a new diagnosis for the so far In addition, the set of thin-sections of Solemydidae aff. identified as valid representatives of Solemydidae, only using Helochelydra sp. (MPG-725-3, a peripheral and MPG-725- the shell bone sculpturing. 4, a possible plastron fragment) used in Pérez-García et al. The palaeoecology of solemydid turtles is still contested and (2013) from Galve (Galve sub-basin, Maestrazgo Basin of the different hypotheses have been proposed. Marmi et al. (2009) Iberian Range, Teruel Province, Spain), together with addi- considered at least some species of the Late Cretaceous Euro- tional sections of four specimens (MNCN 59503, including a pean Solemys Lapparent de Broin and Murelaga, 1996 to have costal, peripheral and two plastral fragments probably belong- a semi-aquatic lifestyle based on palaeoenvironmental and ing to several specimens) from Barremian strata of another taphonomic interpretations of the finding, while also providing locality of the Maestrazgo Basin (Morella, Morella sub-basin, additional shell bone histological evidence. Joyce et al. (2011), Castellón Province, Spain) were added for comparison. All on the other hand, argued for terrestrial habits of solemydid taxa and specimens included in the present study are listed in turtles based on anatomical data, such as the presence of limb Table 1. ossicles. Unfortunately, the described solemydid turtles so far A few sections of the sampled bones of Solemys did not include articulated forelimbs, which could otherwise be vermiculata (MCNA-15047, a costal fragment; MCNA- used to elucidate their palaeoecology, as has been done previ- 15046, a shell fragment) and Solemys sp. (UPUAM-14001, ously for Triassic stem turtles Proganochelys quenstedti Baur, a costal fragment) were further modified into black and white 1887 and Palaeochersis talampayensis Rougier, de la Fuente images (Fig. 2) to perform a compactness analysis (Table 2) and Arcucci, 1995 (Joyce and Gauthier 2004). with the program Bone Profiler, Windows-based version 4.5.8 In addition to taphonomic and morphological data, analysis (Girondot and Laurin 2003), to indicate a potential lifestyle of the microstructure and microanatomy of bones, including based on shell bone microstructures. an increasing body of shell bone data, presents an independent Thin sections were studied and images were taken using a line of evidence to expound the palaeoecology of fossil turtles LEICA compound microscope DM 2500 M equipped with a (e.g. Scheyer 2007; Scheyer and Sander 2007; Scheyer et al. LEICA digital camera DFC 420C. Images were then modified 2014). Furthermore, even small (shell) bone fragments can be into figures using Adobe Creative suite 6 (Photoshop and used for histological analysis, even in cases where more Illustrator). complete fossils including limb bones are absent. In the present study, we thus describe and review the shell Institutional abbreviations bone histology of solemydid turtles with focus on shell bone material from the Late Cretaceous of several sites on the FM, The Field Museum, Chicago, Illinois, USA; IPS, Institut Iberian Peninsula, in comparison to previous histological ac- Català de Paleontologia, Barcelona, Spain; MCNA, Museo de counts of the group and to the histological study of material Ciencias Naturales de Alava, Vitoria-Gasteiz, Spain; from various countries and ages, to elucidate whether the NHMUK, Natural History Museum, London, UK; MNCN, solemydid taxa share microstructural and internal histological Museo Nacional de Ciencias Naturales, Madrid, Spain; MPG, details. These data are then used to elucidate the palaeoecol- Museo Paleontológico de Galve, Galve, Teruel, Spain;
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