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Identification, Distribution, and Function of Gastroliths in Dinosaurs and Extant Birds with Emphasis on Ostriches (Struthio Camelus)

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Identification, Distribution, and Function of Gastroliths in Dinosaurs and Extant Birds with Emphasis on Ostriches (Struthio Camelus) IDENTIFICATION, DISTRIBUTION, AND FUNCTION OF GASTROLITHS IN DINOSAURS AND EXTANT BIRDS WITH EMPHASIS ON OSTRICHES (STRUTHIO CAMELUS) Dissertation zur Erlangung des Doktorgrades (Dr. rer. nat.) der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn vorgelegt von Oliver Wings aus Sangerhausen Bonn 2004 Diplodocid sauropod accidentally ingesting gastroliths while feeding on a cycad. IDENTIFICATION, DISTRIBUTION, AND FUNCTION OF GASTROLITHS IN DINOSAURS AND EXTANT BIRDS WITH EMPHASIS ON OSTRICHES (STRUTHIO CAMELUS) Dissertation zur Erlangung des Doktorgrades (Dr. rer. nat.) der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn vorgelegt von Oliver Wings aus Sangerhausen Bonn 2004 Angefertigt mit Genehmigung der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn 1. Referent: Privat-Dozent Dr. Martin Sander 2. Referent: Professor Dr. Jes Rust Tag der Promotion: 02.12.2004 Diese Dissertation ist auf dem Hochschulschriftenserver der ULB Bonn http://hss.ulb.uni- bonn.de/diss_online elektronisch publiziert This dissertation is published electronically on the ULB Bonn server for university publications: http://hss.ulb.uni-bonn.de/diss_online Dedicated to Claudia. Thanks for everything. French proverb: “Il a un estomac d’autuche!” (literally: He has the stomach of an ostrich!) means: He can tolerate everything! Geheimnisvoll am lichten Tag Läßt sich Natur des Schleiers nicht berauben, Und was sie deinem Geist nicht offenbaren mag, Das zwingst du ihr nicht ab mit Hebeln und mit Schrauben. JOHANN WOLFGANG VON GOETHE, Faust. Erster Teil (Szene Nacht) TABLE OF CONTENTS PREFACE............................................................................................................................... 11 ABSTRACT ............................................................................................................................ 13 ZUSAMMENFASSUNG ....................................................................................................... 15 GENERAL ACKNOWLEDGEMENTS.............................................................................. 17 CHAPTER I: Gastroliths – An overview............................................................................. 19 References ............................................................................................................................ 21 CHAPTER II: A review of gastrolith function and a revised classification..................... 23 Abstract ................................................................................................................................ 23 Introduction ......................................................................................................................... 23 Terminology ......................................................................................................................... 24 Existing definitions for gastrolith....................................................................................... 24 Suggested new definition for gastrolith.............................................................................. 25 Categories of gastrolith regarding their origin ................................................................... 26 Introduction of the term “exolith” ...................................................................................... 27 Definition of geophagy and lithophagy.............................................................................. 29 Gastrolith function .............................................................................................................. 30 Trituration of ingesta .......................................................................................................... 30 Mixing of foodstuff ............................................................................................................ 31 Mineral supply.................................................................................................................... 32 Secretion of stomach juices ................................................................................................ 33 Mineral storage................................................................................................................... 33 Hydrostatic function ........................................................................................................... 34 Other ballast arguments...................................................................................................... 36 Ingestion due to pathological reasons................................................................................. 37 Nutritional diseases............................................................................................................. 37 Destruction of parasites ...................................................................................................... 37 Establishment of a normal intestinal microbial flora.......................................................... 38 Alleviation of hunger and preservation of stomach shape ................................................. 38 Accidental ingestion ........................................................................................................... 39 Stones mistaken as prey...................................................................................................... 40 Playing................................................................................................................................ 40 Nest building....................................................................................................................... 41 Thermoregulation ............................................................................................................... 41 Memorial of events............................................................................................................. 42 Admiration of semi-precious gemstones ............................................................................ 42 Conclusions .......................................................................................................................... 43 Acknowledgements.............................................................................................................. 43 References ............................................................................................................................ 44 Tables.................................................................................................................................... 50 CHAPTER III: Amounts and composition of gastroliths in farm ostriches (Struthio camelus)................................................................................................................................... 51 Abstract ................................................................................................................................ 51 Introduction ......................................................................................................................... 51 Anatomy and physiology of the digestive tract.................................................................. 52 Diet and digestion............................................................................................................... 52 Existing data about gastroliths in ratites .......................................................................... 53 Material and Methods......................................................................................................... 54 Results................................................................................................................................... 56 Ostriches from Remagen, Germany ................................................................................... 56 Rock type composition ..................................................................................................... 57 Grain size distribution and number of gastroliths............................................................. 58 Ostriches from the Klein Karoo, South Africa................................................................... 58 Discussion and Conclusions................................................................................................ 59 Occurrence of gastroliths.................................................................................................... 59 Amounts of gastroliths........................................................................................................ 59 Rock type composition ....................................................................................................... 61 Grain size composition ....................................................................................................... 61 Foreign objects ................................................................................................................... 62 Usability of the data as reference values ............................................................................ 63 Acknowledgements.............................................................................................................. 63 References ............................................................................................................................ 64 Tables...................................................................................................................................
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