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Comparative Bone Histology of the Turtle Shell (Carapace and Plastron)

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Comparative Bone Histology of the Turtle Shell (Carapace and Plastron) Comparative bone histology of the turtle shell (carapace and plastron): implications for turtle systematics, functional morphology and turtle origins Dissertation zur Erlangung des Doktorgrades (Dr. rer. nat.) der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität zu Bonn Vorgelegt von Dipl. Geol. Torsten Michael Scheyer aus Mannheim-Neckarau Bonn, 2007 Angefertigt mit Genehmigung der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn 1 Referent: PD Dr. P. Martin Sander 2 Referent: Prof. Dr. Thomas Martin Tag der Promotion: 14. August 2007 Diese Dissertation ist 2007 auf dem Hochschulschriftenserver der ULB Bonn http://hss.ulb.uni-bonn.de/diss_online elektronisch publiziert. Rheinische Friedrich-Wilhelms-Universität Bonn, Januar 2007 Institut für Paläontologie Nussallee 8 53115 Bonn Dipl.-Geol. Torsten M. Scheyer Erklärung Hiermit erkläre ich an Eides statt, dass ich für meine Promotion keine anderen als die angegebenen Hilfsmittel benutzt habe, und dass die inhaltlich und wörtlich aus anderen Werken entnommenen Stellen und Zitate als solche gekennzeichnet sind. Torsten Scheyer Zusammenfassung—Die Knochenhistologie von Schildkrötenpanzern liefert wertvolle Ergebnisse zur Osteoderm- und Panzergenese, zur Rekonstruktion von fossilen Weichgeweben, zu phylogenetischen Hypothesen und zu funktionellen Aspekten des Schildkrötenpanzers, wobei Carapax und das Plastron generell ähnliche Ergebnisse zeigen. Neben intrinsischen, physiologischen Faktoren wird die Mikrostruktur des Panzerknochens von einem Mosaik phylogenetischer and funktionaler Faktoren beeinflusst. Das Verhältnis beider Einflüsse variiert sehr stark unter den Schildkrötengroßgruppen. Nur wenn funktionelle Aspekte nur schwach ausgeprägt sind, können phylogenetische Signale abgeleitet werden. Die Knochenhistologie kann demnach zur Überprüfung bestehender (morphologischer, molekularer oder serologischer) Verwandtschaftshypothesen genutzt werden. Gruppen, die gut definierte Knochenmikrostrukturen aufweisen, sind die Bothremydidae, Pleurosternidae, Chelydridae, Plesiochelyidae und Thalassemydidae, Dermochelyidae, Dermatemydidae, Carettochelyidae, und Trionychidae. Weiterhin kann die systematische Position unsicher zugeordneter Taxa (z.B. aff. Platychelys sp., Platysternon megacephalum), sowie unzureichend bekanntes Materials bestimmt werden. Aff. Platychelys sp. sowie der Kirtlington Histomorph I werden beide den Pleurosternidae zugeordnet. Die Zuordnung des Histomorph I führt zu einer Ausdehnung des Fossilberichts der Pleurosternidae in den Mittleren Jura hinein. P. megacephalum zeigt einige histologische Gemeinsamkeiten mit den Chelydridae, was wiederum eine Unterstützung älterer morphologischer Hypothesen darstellt. In den restlichen Großgruppen ist kein klares phylogenetisches Signal vorhanden, oder es kommt zu einer Überprägung des Signals durch funktionelle Faktoren. Die Anpassung der Knochenmikrostruktur des Panzers an das aquatische Milieu gehört zu den stärksten funktionellen Faktoren. Hierdurch konnte eine Gruppierung aller untersuchten Schildkröten in vier Kategorien (I „terrestrischer Lebensraum“ bis IV „extremste Anpassung an das aquatische/marine Milieu) bezüglich ihrer Ökologie/Palökologie vorgenommen werden. Vergleiche der ältesten Vertreter der Schildkröten mit rezenten ‚aquatischen’ und ‚terrestrischen’ Vertretern belegen unabhängig die terrestrische Palökologie der basalen Testudinata. Die Knochenpanzermikrostrukturen wurden weiterhin zur Klärung des Ursprungs der Schildkröten genutzt. Basierend auf dem Vergleich von basalen Schildkröten und verschiedenen Außengruppenvertretern, welche Pareiasaurier, Placodontier, Mammalier, Archosauromorphe und Lepidosaurier beinhalteten, wird ein Ursprung innerhalb der Diapsida mit naher Verwandtschaft zu Archosauriern hypothetisiert. Für den Panzer der Placodontier wird weiterhin ein, in Osteodermen bisher unbekanntes knorpeliges Gewebe (´postkranialer faserknorpelhaltiger Knochen´), sowie ein generelles Modell der Osteogenese vorgestellt. Abstract—The bone histology of the turtle shell is valuable for addressing osteoderm and shell formation, reconstruction of fossil integumentary soft-tissue structures, phylogenetic hypotheses and functional aspects of the turtle shell, with both carapace and plastron showing similar results. Besides intrinsic physiological factors, the shell bones are proposed to be influenced by a mosaic of phylogenetic and functional factors influencing the microstructural properties. The ratio between phylogenetic and functional constraints is highly variable among the major turtle groups, and only where functional aspects are less dominant, phylogenetic signals can be deduced from the bone histology. The bone histology can thus be used to verify existing intra-specific phylogenetic (e.g., morphological, molecular and serologic) hypotheses among turtles. Groups that are well defined by bone histological characters are Bothremydidae, Pleurosternidae, Chelydridae, Plesiochelyidae and Thalassemydidae, Dermochelyidae, Dermatemydidae, Carettochelyidae and Trionychidae. Furthermore, the systematic position of uncertainly assigned taxa (e.g., aff. Platychelys sp., Platysternon megacephalum) and poorly known shell material (e.g., Kirtlington turtles) could be assessed. Aff. Platychelys sp., as well as Kirtlington histomorph I are both assigned to Pleurosternidae herein. Assignment of the latter taxon would indicate that the fossil record of Pleurosternidae has to be extended back into the Middle Jurassic. P. megacephalum was found to share some histological features with Chelydridae, thus supporting prior morphological hypothesis. In the other major turtle groups, the bone histology does not show clear phylogenetic signals or functional factors override existing phylogenetic signals respectively. One functional aspect that profoundly influences turtle shell bone microstructures is the adaptation to an aquatic habitat and life-style. In this respect, all turtles were grouped into four categories (I “terrestrial environment” to IV “extreme adaptation to aquatic/marine environments”), based on their ecology/palaeoecology. Comparison of the oldest known turtles with recent ‘aquatic’ and ‘terrestrial’ turtles independently revealed a terrestrial palaeoecology for the basal Testudinata. Shell bone microstructures can further elucidate the origin of turtles. Based on the comparison of basal turtles and several outgroup taxa including osteoderm-bearing pareiasaurs, mammals, placodonts, archosauromorphs and lepidosaurs, the origin of turtles is hypothesised to lie within Diapsida, with close relationships to archosaurs. In the case of placodont armour, a unique bone tissue (here termed ´postcranial fibro- cartilaginous bone`) is described and a general model of osteogenesis is proposed. Contents ZUSAMMENFASSUNG ABSTRACT 1. Introduction 1 1.1 GENERAL INTRODUCTION 1 1.2 MORPHOLOGY AND ANATOMY OF THE TURTLE SHELL 1 1.3 AIMS OF THE STUDY 5 1.3.1 Implications for turtle systematics 5 1.3.2 Implications for turtle shell functional morphology 8 1.3.3 Implications for turtle origins 9 1.3.4 Implications for the ecology of turtles 11 1.4 PREVIOUS WORK 14 1.4.1 Historical aspects on bone histology 14 1.4.2 Dermal bone histology and metaplastic bone formation 14 1.4.3 Reptile dermal bone histology 15 1.4.4 Turtle shell bone histology 16 1.4.5 Historical introduction to the development of the turtle shell 16 1.4.6 Current consensus on the development of the turtle shell 17 2. Material and Methods 20 2.1 SAMPLING STRATEGY 20 2.2 PREPARATION 21 2.2.1 Sampling of turtle shell elements 21 2.2.2 Sampling by core-drilling 21 2.2.3 Planes of Sectioning 23 2.2.4 Preparation of standard petrographic thin-sections 24 2.2.5 Analysis and documentation 24 2.2.6 Picture credits 25 2.3 TERMINOLOGY 25 2.4 INSTITUTIONAL ABBREVIATIONS 26 3. Morphological description of outgroup taxa 28 3.1 OUTGROUP 1: TEMNOSPONDYL AMPHIBIANS 28 3.1.1 Trimerorhachis sp. 28 3.1.2 Mastodonsaurus giganteus (Jaeger, 1828) 29 3.1.3 Gerrothorax pustuloglomeratus (Huene, 1922) 29 3.2 OUTGROUP 2: MAMMALIA 29 3.2.1 Folivora (Xenarthra) 30 3.2.2 Cingulata (Xenarthra) 30 3.3 OUTGROUP 3: NON-TESTUDINATAN REPTILIA 31 3.3.1 Parareptilia (Pareiasauria) 32 3.3.2 Eureptilia (Placodontia) 34 3.3.3 Eureptilia (Lepidosauria) 39 3.3.4 Eureptilia (Archosauromorpha) 39 4. Morphological description of Testudinata 41 4.1 BASAL TESTUDINATA 41 4.1.1 Proganochelyidae 41 4.1.2 Proterochersidae 42 4.1.3 Kayentachelyidae 43 4.1.4 Meiolaniidae 44 4.2 PLEURODIRA 45 4.2.1 Platychelyidae 46 4.2.2 Pelomedusidae 47 4.2.3 Bothremydidae 48 4.2.4 Podocnemidae 49 4.2.5 Chelidae 51 4.3 CRYPTODIRA 56 4.3.1 Cryptodira incertae sedis (Kirtlington turtle sample; Solemydidae) 61 4.3.2 Baenidae 62 4.3.3 Pleurosternidae 64 4.3.4 Eurysternidae 66 4.3.5 Plesiochelyidae and Thalassemydidae 67 4.3.6 Xinjiangchelyidae 68 4.3.7 “Sinemydidae” and “Macrobaenidae” 70 4.3.8 Cheloniidae “sensu lato” 71 4.3.9 Cheloniidae “sensu stricto” 74 4.3.10 Protostegidae 76 4.3.11 Dermochelyidae 77 4.3.12 Chelydridae 79 4.3.13 Testudinoidea indet. 82 4.3.14 Emydidae 82 4.3.15 Bataguridae/Geoemydidae 86 4.3.16 Testudinidae 93 4.3.17 Eucryptodira incertae sedis (aff. ?Trionychoidea) 98 4.3.18 Dermatemydidae 99 4.3.19 Kinosternia 101 4.3.20 Kinosternidae 102 4.3.21 Adocidae 104 4.3.22 Nanhsiungchelyidae 105 4.3.23 Carettochelyidae 106 4.3.24 Trionychidae (Plastomeninae, Cyclanorbinae and Trionychinae) 109 5. Bone histological results of outgroup
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