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Unravelling the Positional Behaviour of Fossil Hominoids: Morphofunctional and Structural Analysis of the Primate Hindlimb

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Unravelling the Positional Behaviour of Fossil Hominoids: Morphofunctional and Structural Analysis of the Primate Hindlimb ADVERTIMENT. Lʼaccés als continguts dʼaquesta tesi queda condicionat a lʼacceptació de les condicions dʼús establertes per la següent llicència Creative Commons: http://cat.creativecommons.org/?page_id=184 ADVERTENCIA. El acceso a los contenidos de esta tesis queda condicionado a la aceptación de las condiciones de uso establecidas por la siguiente licencia Creative Commons: http://es.creativecommons.org/blog/licencias/ WARNING. The access to the contents of this doctoral thesis it is limited to the acceptance of the use conditions set by the following Creative Commons license: https://creativecommons.org/licenses/?lang=en Doctorado en Biodiversitat Facultad de Ciènces Tesis doctoral Unravelling the positional behaviour of fossil hominoids: Morphofunctional and structural analysis of the primate hindlimb Marta Pina Miguel 2016 Memoria presentada por Marta Pina Miguel para optar al grado de Doctor por la Universitat Autònoma de Barcelona, programa de doctorado en Biodiversitat del Departamento de Biologia Animal, de Biologia Vegetal i d’Ecologia (Facultad de Ciències). Este trabajo ha sido dirigido por el Dr. Salvador Moyà Solà (Institut Català de Paleontologia Miquel Crusafont) y el Dr. Sergio Almécija Martínez (The George Washington Univertisy). Director Co-director Dr. Salvador Moyà Solà Dr. Sergio Almécija Martínez A mis padres y hermana. Y a todas aquelas personas que un día decidieron perseguir un sueño Contents Acknowledgments [in Spanish] 13 Abstract 19 Resumen 21 Section I. Introduction 23 Hominoid positional behaviour The great apes of the Vallès-Penedès Basin: State-of-the-art Section II. Objectives 55 Section III. Material and Methods 59 Hindlimb fossil remains of the Vallès-Penedès hominoids Comparative sample Area of study: The Vallès-Penedès Basin Methodology: Generalities and principles Section IV. The Femora 87 Chapter 1. External morphology of the femur 89 Descriptions Comparative sample, measurements and statistical analysis Comparisons Chapter 2. Femoral neck cortical bone distribution 105 Comparative sample, measurements and methods Results Chapter 3. Cross-sectional structural properties of the shaft 121 Comparative sample, measurements and methods Results Section V. The Tibia 133 Chapter 4. External morphology of the tibia 135 Description Comparative sample, measurements and statistical analysis Comparisons Section VI. The Patella 143 Chapter 5. External morphology of the patella 145 Description Comparative sample, measurements and statistical analysis Comparisons Chapter 6. Patellar biomechanics during knee !exion 155 Comparative sample The FE method Experimental FE analyses Results Section VII. Discussion 171 Functional inferences The positional behaviour of the fossil great apes from the Vallès-Penedès Section VIII. Summary and Conclusions 201 Section IX. References 207 Section X. Appendix A 237 ACKNOWLEDGEMENTS [in Spanish] Me enfrento a una hoja en blanco con el reto y la responsabilidad de mostrar mi agradecimiento a todas aquellas personas que han hecho posible esta tesis, directa o indirectamente, y espero saber hacerlo lo mejor posible. Lo hago en español para que este agradecimiento llegue a todas ellas. Sino lo consigo a través de estas líneas, espero haber sido capaz de expresarlo o demostrarlo de una manera u otra a lo largo de estos años, desde que pisé por primera vez un yacimiento con mi amiga María de Bellas Artes hasta hoy que escribo estas palabras. Isaac Newton dijo una vez “If I have seen further it is by standing on the shourlders of giants” [Si he logrado ver más lejos, ha sido porque me he alzado sobre hombros de gigantes]. Yo no sería capaz de expresar mejor con palabras mi agradecimiento y admiración por los directores de esta tesis, el Dr. Salvador Moyà-Solà y el Dr. Sergio Almécija. Gracias a vuestra ayuda y consejos hoy tengo entre mis manos una tesis que casi me parecía imposible realizar, ¡gracias!. Me gustaría agradecerle a Salvador que contestara a ese primer email que le mandé y que tardé casi dos semanas en escribir. Agradecerle también que sin prácticamente conocerme pusiera en mis manos la responsabilidad de estudiar los fósiles que forman parte de este trabajo, por enseñarme que no hay mejor manera de lidiar con esta responsabilidad que desde la pasión, la pasión por la paleontología, y las ganas de aprender y descifrar el pasado. Debo agradecerle también pronunció su nombre mientras conversábamos sobre el proyecto de la tesis sentados en el despacho de los antiguos módulos. A Sergio me gustaría agradecerle su gran exigencia y el continuo reto intelectual al que me ha hecho enfrentarme durante estos años. Aunque duro y difícil en algunos momentos, me atrevo a decir que ha valido la pena. Agradecerle también todo lo que se ha preocupado por mí y por mi carrera, así como porque las cosas salieran bien, por todos esos mails que ha mandado (los que sé y los que creo desconozco) para que consiguiera CTs, datos o lo que haya ido necesitando. Me tomo además la licencia de decir que para mí ha sido un placer poder crecer de manera paralela en nuestros recién estrenados papeles de director y doctoranda. 13 Quiero dar también las gracias a todos los investigadores con los que he compartido estos años y/o algunos trabajos, y que de alguna manera u otra me han dejado aprender de ellos y han compartido algo de manera especial al Dr. David M. Alba porque me gustaría agradecerle todos los consejos, charlas y Agradecerle también esos manuscritos corregidos “completamente en rojo”, un reto, pero siempre un aprendizaje. Destacar igualmente mi agradecimiento al Dr. Daniel DeMiguel, por su paciencia, su ayuda incansable y por tener siempre una palabra amable y una respuesta para todas esas dudas y miedos resultados expuestos en esta tesis forman parte de dos artículos ya publicados y otros manuscritos todavía en preparación. Por este motivo me gustaría darle las gracias al resto de coautores que no he nombrado hasta el momento: Dr. Josep Fortuny, por su ayuda durante mis primeros pasos (y no tan primeros...) en el mundo de la paleontología virtual; al Dr. Jordi Marcé-Nogué y Dr. Francesc Puigvert, por su paciencia y ayuda para sacar adelante el trabajo en algo tan nuevo para mí como eran los Elementos Finitos; al Dr. Matthew C. O’Neill, por su ayuda con las regresiones PGLs; a la Dra. Melissa Tallman, por su ayuda propiedades biomecánicas de los fémures. Gracias también al Dr. Isaac Casanovas-Vilar por todo lo que me ha contado sobre la geología y los yacimientos de esta fascinante cuenca (¡y por los mapas!); al Dr. Joan Madurell por esas charlas sobre los yacimientos cuaternarios, carnívoros y otros bichos varios; y a la Dra. Ashley Hammond por sus consejos y comentarios sobre algunas partes de esta tesis. Igualmente me gustaría agradecerle de manera muy especial a Marta Palmero todos esos consejos estéticos y de diseño a lo largo de estos años, así como su gran ayuda en la maquetación de este trabajo. Darle también las gracias a los investigadores internacionales que amablemente se han prestados a revisar la tesis para que opte a la mención internacional: Dr. William I. Sellers y Dr. Christopher B. Agradecerle también de manera especial toda la ayuda prestada con los temas relacionados con esta tesis o con la FPU a la coordinadora del doctorado, la Dra. Assumpció Malgossa. Quisiera también dar las gracias a los investigadores que amablemente han compartido con nosotros datos, modelos 3D, imágenes o CTs para realizar los trabajos que conforman esta tesis: C. Owen Ibáñez, Masato Nakatsukasa, Daisuke Shimizu, Lorenzo Rook y Roberto Machiarelli. Gracias también a las empresas CESPA Gestión de Residuos y FOSSILIA Serveis Paleontològics i Geològic, S.L. por las fotografías del ACM que en algún momento de la tesis nos han cedido amablemente. Agradecer también a la empresa Transmitting Science S.L. y a Soledad de Esteban-Trivigno la organización de los cursos a los que he asistido y que han sido de gran ayuda en algunas partes de esta tesis. 14 Dar las gracias a todas aquellos centros e instituciones que me han permitido visitar y trabajar en sus colecciones o usar sus equipos: Institut Català de Paleontologia Miquel Crusafont (ICP, Sabadell). Hospital Universitari Mútua de Terrassa (Terrassa). Unitat de Antropologia (Dpto. de Biologia Animal, de Biologia Vegetal i d’Ecologia) de la Universitat Autònoma de Barcelona (UAB, Bellaterra). Universidad de Burgos (UBU, Brugos). American Museum of Natural History (AMNH, Nueva York): Mammalogy Collection y Microscopy and Imaging facilities. ‘Abdus Salam’ International Centre for Theoretical Physics (ICTP, Triestre). [Multidisciplinary Laboratory of the International Centre for Theoretical Physics in the frame of the ICTP/Elettra EXACT Project, funded by the Regione Friuli-Venezia Giulia]. Royal Museum of Central Africa (RMCA, Tervuren). Stony Brook University (SBU, Stony Brook). Smithsonian National Museum of Natural History (USNM, Washington). Museum of Comparative Zoology (MCZ), PaleoLab, Peabody Museum of Archaeology and Ethnology (PMAE) y Center for Nanoscale Systems (CNS) pertenecientes a Harvard University (Cambridge). [The CNS is a member of the National Nanotechnology Infrastructure Network (NNIN), which is supported by the National Science Foundation under NSF award no. ECS-0335765. CNS is part of Harvard University]. Naturischorisches Museum (NHMW, Viena) y MicroCT-Lab de la Universität Wien (UNIVIE, Viena); Museo Nacional de Ciencias Naturales-CSIC (MNCN, Madrid). Igualmente, agradecer su ayuda y amabilidad a todos
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