Evolution of Body Size of Extinct Endemic Small Mammals from Mediterranean Islands
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PhD Thesis Universitat Autònoma de Barcelona Doctorate in Biodiversity 2016 Evolution of body size of extinct endemic small mammals from Mediterranean Islands BLANCA MONCUNILL SOLÉ SUPERVISORS o MEIKE KÖHLER p o SALVADOR MOYÀ SOLÀ p o XAVIER JORDANA p Departament de Biologia Animal, Biologia Vegetal i d’Ecologia PhD Thesis - 2016 Evolution of body size of extinct endemic small mammals from Mediterranean Islands Blanca Moncunill Solé DissertaƟ on presented by Blanca Moncunill Solé in fulfi llment of the requirements for the degree of Doctor in the Universitat Autònoma de Barcelona, doctorate program in Biodiversity of the Departament de Biologia Animal, Biologia Vegetal i d’Ecologia. Under the supervision of: • Dra. Meike Köhler, ICREA at InsƟ tut Català de Paleontologia Miquel Crusafont and teacher of the Departament de Biologia Animal, Biologia Vegetal i d’Ecologia at Universitat Autònoma de Barcelona. • Dr. Salvador Moyà-Solà, ICREA at InsƟ tut Català de Paleontologia Miquel Crusafont and teacher of the Departament de Biologia Animal, Biologia Vegetal i d’Ecologia at Universitat Autònoma de Barcelona. • Dr. Xavier Jordana, Postdoctoral Researcher at InsƟ tut Català de Paleontologia Miquel Crusafont. Doctoral candidate B½Ä MÊÄçÄ®½½ Sʽ Supervisors Dr. S½òÊÙ MÊù Sʽ Dra. M®» KÏ«½Ù Dr. Xò®Ù JÊÙÄ All the work (text, tables, fi gures, illustrations, design and typesetting edition) comprised in this PhD Thesis is the result of research and tasks carried out by the author. When information has been derived from another sources, it is accurately indicated in the text. B M S Bellaterra, June 2016 qr© Rights Reserved Where does it come from, this quest, this need to solve life’s mysteries when the simplest of ques ons can never be answered? Why are we here? What is the soul? Why do we dream? Perhaps we’d be be er off not looking at all. Not delving, not yearning. But that’s not human nature. Not the human heart. That is not why we are here. Mohinder Suresh (Genesis, Heroes) CONTENTS p PROLOGUE p 6 p ABBREVIATIONS p 7 p CHAPTER 1: Introduction p 9 p 1.1. Body size p 11 p 1.1.1. Introduction to body size p 11 p Retrospective review of history in body size’s research p 12 p 1.1.2. Scaling: principles and mechanisms of funcionality of live animals p 13 p Isometry and allometry p 13 p Body mass: estimations in fossil record p 14 p 1.1.3. Broad patterns in diversifi cation of body size: time and space p 15 p Variation in time: Cope’s Rule p 15 p Variation in space: Bergmann’s Rule and Island Rule p 15 p 1.1.4. Biology and ecology of body size p 16 p Selection pressures operating on body size: benefi ts and costs of large and small sizes p 16 p Allometric relationship among life history and body size p 17 p 1.2. Islands p 19 p 1.2.1. Introduction to islands: Mediterranean Islands p 19 p 1.2.2. Insular faunas: Island Rule and Island Syndrome p 20 p 1.2.3. Insular ecosystems: a new ecological regime p 23 p 1.2.4. Optimal body size p 27 p CHAPTER 2: Aims & Objectives p 29 p CHAPTER 3: Materials & Methods p 35 p 3.1. Database of extant and extinct fauna p 37 p 3.2. Skeletal measurements and body mass data p 41 p 3.3. Statistical methodology p 44 p Allometry for predicting body mass p 44 p Statistical regression models: Ordinary Least Squares p 45 p Assessment of the regression models p 46 p Predicting the body mass of fossil species p 47 p 3.4. Supplementary data p 48 p CHAPTER 4: How large are the extinct giant insular rodents? New body mass estimations from teeth and bones p 77 p Abstract p 79 p Introduction p 79 p Materials and methods p 81 p Results p 83 p Discussion p 87 1 p Acknowledgments p 90 p References p 91 p Supporting information p 94 p CHAPTER 5: The Island Rule and the native island of Mikrotia magna (Muridae, Rodentia) from Terre Rosse deposits (Gargano, Apulia, Italy): inferences from its body mass estimation p 101 p Abstract p 103 p Introduction p 104 p Abbreviations p 105 p Materials and methods p 106 p Results p 107 p Discussion p 108 p Conclusions p 113 p Acknowledgments p 113 p Bibliography p 114 p CHAPTER 6: The weight of fossil leporids and ochotonids: body mass estimation models for the order Lagomorpha p 119 p Abstract p 121 p Introduction p 121 p Materials and methods p 122 p Results p 124 p Discussion p 127 p Acknowledgments p 128 p References p 128 p Supporting information p 130 p CHAPTER 7: Comparing the body mass variations in endemic insular species of Prolagus (Ochotonidae, Lagomorpha) in the Pleistocene of Sardinia (Italy) p 145 p Abstract p 147 p Introduction p 147 p Abbreviations p 148 p Material p 149 p Methods p 149 p Results p 150 p Discussion p 150 p Conclusions p 155 p Acknowledgments p 156 p References p 156 p CHAPTER 8: First approach of the life history of Prolagus apricenicus (Ochotonidae, Lagomorpha) from Terre Rosse sites (Gargano, Italy) using body mass estimation and paleohistological analysis p 161 p Abstract p 163 p Introduction p 164 2 p Material and methods p 165 p Results p 166 p Discussion p 170 p Conclusions p 171 p Acknowledgments p 171 p References p 172 p CHAPTER 9: How common is gigantism in insular fossil shrews? Examining the “Island Rule” in soricids (Mammalia: Soricomorpha) from Mediterranean Islands using new body mass estimation models p 175 p Abstract p 177 p Introduction p 177 p Material and methods p 178 p Results p 183 p Discussion p 188 p Conclusion p 193 p Acknowledgments p 193 p References p 193 p Supporting information p 196 p CHAPTER 10: Discussion p 205 p 10.1. Body mass regression models for small mammals p 207 p Bivariate and multiple regression models p 207 p 10.2. Body mass estimation of fossil species of small mammals p 211 p Body masses of fossil small mammal species: comparison with previous body mass estimations p 214 p Body masses of fossil small mammal species: comparison with extant relative species p 215 p 10.3. Body mass evolution and Island Rule in small extinct mammals p 219 p Direction and magnitude of body size shift in extinct insular small mammals p 221 p Factors that trigger the Island Rule in small mammals: a paleontological view p 222 p Evolutionary trends in body size shifts under insular regimes in extinct small mammals p 228 p Body size shifts and Life History strategy in extinct small mammals p 230 p CHAPTER 11: Conclusions p 235 p CHAPTER 12: References p 241 p CHAPTER 13: Acknowledgments p 269 3 PROLOGUE Body size (or its proxy: body mass) has a central posi on in the colossal web of interdependent biological variables of an organism.