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Evolution of the Hand in Miocene Apes: Implications for the Appearance of the Human Hand

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Evolution of the Hand in Miocene Apes: Implications for the Appearance of the Human Hand Evolution of the hand in Miocene apes: implications for the appearance of the human hand Universitat Autònoma de Barcelona Facultat de Ciències Departament de Biologia Animal, Biologia Vegetal i d'Ecologia Unitat d'Antropologia Biològica PhD Thesis Evolution of the hand in Miocene apes: implications for the appearance of the human hand Sergio Almécija 2009 Institut Català de Paleontologia Memòria presentada per Sergio Almécija per optar al títol de Doctor per la Universitat Autònoma de Barcelona, programa de doctorat en Biologia (opció Antropologia) del Departament de Biologia Animal, Biologia Vegetal i d'Ecologia. Recerca dirigida pel Professor Salvador Moyà-Solà, ICREA a l'Institut Català de Paleontologia i a la Unitat d'Antropologia Biològica del Departament de Biologia Animal, Biologia Vegetal i d'Ecologia, Universitat Autònoma de Barcelona. Doctorand: Sergio Almécija Director: Prof. Salvador Moyà-Solà 1 Sergio Almécija 2009 To the Big Bang, for everything 2 Evolution of the hand in Miocene apes: implications for the appearance of the human hand Contents Pròleg [Prologue, in Catalan]---------------------------------------------------------------------- 6 Agraïments [Acknowledgements] ---------------------------------------------------------------- 9 Chapter 1 A brief introduction to the hands of the primates -------------------------------- 15 Hand anatomy ------------------------------------------------------------------------------- 15 Prehension------------------------------------------------------------------------------------ 18 Hand function and capabilities ------------------------------------------------------------ 19 Hand and manipulation ----------------------------------------------------------------- 20 Hand and locomotion -------------------------------------------------------------------- 22 Objectives of this work --------------------------------------------------------------------- 24 Chapter 2 Pierolapithecus catalaunicus: Insights from the hand phalanges of a stem great ape -------------------------------------------------------------------------------------------- 26 Introduction ---------------------------------------------------------------------------------- 27 Materials and methods --------------------------------------------------------------------- 28 Measurements ---------------------------------------------------------------------------- 28 The comparative fossil sample --------------------------------------------------------- 29 The comparative extant sample -------------------------------------------------------- 29 Statistical methods ----------------------------------------------------------------------- 30 Description of the manual phalanges of Pierolapithecus ----------------------------- 34 Proximal phalanges ---------------------------------------------------------------------- 34 Intermediate phalanx -------------------------------------------------------------------- 35 Distal phalanges -------------------------------------------------------------------------- 36 Morphological comparisons --------------------------------------------------------------- 37 Proximal phalanges ---------------------------------------------------------------------- 37 Intermediate phalanges ------------------------------------------------------------------ 38 Distal phalanges -------------------------------------------------------------------------- 38 Morphometric comparisons --------------------------------------------------------------- 39 Discussion ------------------------------------------------------------------------------------ 45 Phalangeal morphology in Miocene apes -------------------------------------------- 45 Phalangeal curvature -------------------------------------------------------------------- 46 Robusticity and intrinsic relative phalangeal length -------------------------------- 51 Suspensory behaviors in Pierolapithecus -------------------------------------------- 55 3 Sergio Almécija 2009 Conclusions ---------------------------------------------------------------------------------- 57 Chapter 3 Hispanopithecus laietanus: The first steps towards the orangutan suspensory behaviors ------------------------------------------------------------------------------------------- 62 Introduction ---------------------------------------------------------------------------------- 63 Materials and methods --------------------------------------------------------------------- 65 Measurements and comparative sample ---------------------------------------------- 65 Statistical techniques -------------------------------------------------------------------- 65 Results ---------------------------------------------------------------------------------------- 66 Reconstruction and morphological description -------------------------------------- 66 Morphometric comparisons ------------------------------------------------------------ 70 Discussion ------------------------------------------------------------------------------------ 73 Functional interpretation ---------------------------------------------------------------- 73 Paleobiological reconstruction and evolutionary inferences ----------------------- 78 Chapter 4 Locomotor inferences from Pierolapithecus and Hispanopithecus phalanges ------------------------------------------------------------------------------------------------------- 96 Introduction ---------------------------------------------------------------------------------- 97 Locomotor inferences based on manual phalanges ------------------------------------ 98 On locomotor categories ---------------------------------------------------------------- 98 Phalangeal curvature based on included angle (IA) measurements -------------- 99 Index of relative curvature (IRC) ----------------------------------------------------- 100 Relative phalangeal length (RPL) ---------------------------------------------------- 102 Phalangeal morphology ---------------------------------------------------------------- 105 Discussion ----------------------------------------------------------------------------------- 105 Conclusions --------------------------------------------------------------------------------- 107 Chapter 5 Homo habilis or Paranthropus? --------------------------------------------------- 118 Introduction --------------------------------------------------------------------------------- 119 Materials and methods -------------------------------------------------------------------- 122 Methodology and variables employed ----------------------------------------------- 122 Statistical comparisons ----------------------------------------------------------------- 124 Comparative sample -------------------------------------------------------------------- 127 Results --------------------------------------------------------------------------------------- 127 Discussion ----------------------------------------------------------------------------------- 140 Taxonomic attribution of the Swartkrans hominin phalanges -------------------- 140 Taxonomic attribution of the OH 7 phalanges -------------------------------------- 143 4 Evolution of the hand in Miocene apes: implications for the appearance of the human hand Hand function in Paranthropus ------------------------------------------------------- 148 Implications for the study of the origin of the hominid hand and culture ------ 154 Summary and conclusions ---------------------------------------------------------------- 158 Chapter 6 OH 7: The original handy man? --------------------------------------------------- 160 Introduction --------------------------------------------------------------------------------- 161 Materials and methods -------------------------------------------------------------------- 163 Results --------------------------------------------------------------------------------------- 165 Morphological comparisons ----------------------------------------------------------- 165 Morphometric comparisons ----------------------------------------------------------- 166 Discussion ----------------------------------------------------------------------------------- 167 Conclusions --------------------------------------------------------------------------------- 169 Chapter 7 Orrorin tugenensis: Early origins of refined manipulation -------------------- 171 Introduction --------------------------------------------------------------------------------- 172 Materials and methods -------------------------------------------------------------------- 173 Results --------------------------------------------------------------------------------------- 174 Discussion ----------------------------------------------------------------------------------- 176 Regarding the taxonomic and anatomic attribution of the OH 7 hand remains 179 Conclusions --------------------------------------------------------------------------------- 179 Summary ------------------------------------------------------------------------------------------ 182 Discussion and conclusions--------------------------------------------------------------------- 186 Epilogue ------------------------------------------------------------------------------------------- 195 Glossary ------------------------------------------------------------------------------------------- 196 References ----------------------------------------------------------------------------------------- 200 5 Sergio Almécija 2009 Pròleg [Prologue, in Catalan] Aquesta treball de tesi és el resultat de la investigació duta terme des
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