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 NORMAL AND PATHOLOGICAL FOOT BONES VARIABILITY IN HISTORICAL AND MODERN SERIES EDUARDO SALDÍAS VERGARA DOCTORAL THESIS, 2019 DR. ASSUMPCIÓ MALGOSA MORERA DR. ALBERT ISIDRO LLORENS DIRECTORS UNITAT D’ANTROPOLOGIA BIOLOGICA DEPARTAMENT DE BIOLOGIA ANIMAL, BIOLOGIA VEGETAL I D’ECOLOGIA DOCTORAT EN BIODIVERSITAT NORMAL AND PATHOLOGICAL FOOT BONES VARIABILITY IN HISTORICAL AND MODERN SERIES EDUARDO SALDÍAS VERGARA DOCTORAL THESIS 2019 DR. ASSUMPCIÓ MALGOSA MORERA DR. ALBERT ISIDRO LLORENS DIRECTORS 2 This research received support from the Comisión Nacional de Investigación Científica y Tecnológica (CONICYT), from the Ministry of Education of the Government of Chile (CONICYT PAI/2014-73130722). In addition, the author of this thesis is member of 2017SGR1630 of the Generalitat de Catalunya, Spain. 3 NORMAL AND PATHOLOGICAL FOOT BONES VARIABILITY IN HISTORICAL AND MODERN SERIES Thesis presented by Eduardo Saldías Vergara, in fulfillment of the requirements for the Doctorate in Biodiversity, international mention of Departament de Biologia Animal, Biologia Vegetal i d’Ecologia, Universitat Autònoma de Barcelona, directed by: Dr. Assumpció Malgosa Morera, Chair Professor at Unitat d’Antropologia Biològica, Departament de Biologia Animal, Biologia Vegetal i d’Ecologia, Universitat Autònoma de Barcelona. Dr. Albert Isidro Llorens, PhD in Medicine and Surgery, and specialist in Orthopedic Surgery and Traumatology at Servei COT, Hospital Universitari Sagrat Cor, Barcelona, Spain. _________________________ _________________________ Assumpció Malgosa Morera Albert Isidro Llorens Director Director ______________________________ Eduardo Saldías Vergara PhD candidate 4 In memoriam to Eduardo Vergara and Domingo Saldías, my grandparents, who passed away during my PhD process. My best memories and their legacy always will be in my heart. 5 Acknowledgements Special thanks to my family, who supported me in every one of my academic steps. Despite the distance, circumstances and difficulties, they were always present. Their support, principles and teachings have been fundamental to achieve my goals. In addition, I want to thank all my friends who were my second family in Barcelona. Here, I would like to mention my friends from Chile, Colombia, Italy and Spain. Specially, I want to thank to Gonzalez- Bahamonde and Calle-Reyes families, who welcomed me as one of them. My gratitude to the Government of Chile (CONICYT Becas Chile) for the financial support. The opportunity to study abroad would be impossible with my own resources. Academically, I would like to thank my PhD directors, who guided me in this long career to get important results and grow up as a professional. Their advice and suggestions will be very useful in the future, especially when I return to Chile and transfer their knowledge. In the same line, I am thankful to everyone who supported this project and allowed me to study their collections. Taking in consideration the above, thanks to José Antonio Sánchez (Universidad Complutense de Madrid), Miguel Botella and Inmaculada Alemán (Universidad de Granada), Alfonso Rodríguez (Universitat Autònoma de Barcelona), Antonio Moro (Museu de Terrasa), Mauro Rubini (Servizio di Antropologia della Soprintendenza del Lazio), Alfredo Coppa (Sapienza Università di Roma), Gianluca Zanzi (Sovrintendenza Capitolina ai Beni Culturali di Roma Capitale), Cristina Martínez-Labarga and Marica Baldoni (Tor Vergata Università degli studi di Roma), Carlos Garcia Mallo, Rodolfo Marín and Anna Gallart (Hospital Universitari Sagrat Cor de Barcelona) and Lucrezia Marchiandi who helped me with the Italian language and Roman life. Finally, to Bernardo Vila, Patricia Chandía, Joesephina Saager, Niklas Li, Alexandra Salas, Eduardo Becker, Eugenia Voronova, Cristian Ramos, Alex Rodríguez and Ana María Martínez for their academic support. 6 “They called us resurrectionists, grave robbers.” “The miracle of life is granted, and how that miracle can be defective, is a nuance that I am most interested in understanding.” “I must know why five fingers are intended before I can discover the cause of six.” E.B. Hudspeth, “The Resurrectionist: The Lost Work of Dr. Spencer Black” 7 “Sapere aude”… (Horatius, Epistularum liber primus) 8 Index 1. Introduction .................................................................................................................... 10 1.1. Abstract ...................................................................................................................... 11 1.2. Hominization and bipedalism .................................................................................... 13 1.3. Foot evolution in hominids ........................................................................................ 22 1.4. Foot morphology in modern humans ......................................................................... 33 1.5. Differences in foot structure: human variability and pathologies .............................. 37 1.6. Aims ........................................................................................................................... 42 1.7. Structure of the thesis ................................................................................................ 44 1.8. References .................................................................................................................. 46 2. Results .............................................................................................................................. 53 2.1. Sex estimation from the navicular bone in Spanish contemporary skeletal collections .......................................................................................................................................... 54 2.2. Morphological and biomechanical implications of cuboid facet of the navicular bone in the gait .......................................................................................................................... 61 2.3. Pathological and normal variability of the foot bones in osteological collections .... 69 2.4. A new methodology to estimate flat foot in skeletal remains. The example of Mediterranean collections ............................................................................................... 103 2.5. Bipartite cuneiforms in osteological and scanner series .......................................... 136 2.6. Earliest probable case of Mueller-Weiss Disease from Ancient Egypt ................... 159 3. Discussion ...................................................................................................................... 163 3.1. General considerations ............................................................................................. 164 3.2. The study of the foot ................................................................................................ 165 3.2.1. Normal variability ............................................................................................. 165 3.2.2. Pathological variability ..................................................................................... 170 3.3. Final considerations ................................................................................................. 177 3.4. References ................................................................................................................ 178 4. Conclusions ................................................................................................................... 184 9 1. INTRODUCTION “The human foot is a masterpiece of engineering and a work of art”. Leonardo da Vinci 10 1.1. Abstract Introduction: This thesis was devoted to the study of the foot, mainly in ancient collections. The foot is a forgotten bone set in many anthropological, paleopathological, and forensic studies. However, being the key structure that maintains the upright position, it is basic to know the life of a specific individual and the destiny of a population. To understand the structural and biomechanical changes of the modern foot, it is necessary to appeal to Paleoanthropology; the unique science that studies in detail the foot anatomy of our ancestors. Thus, through an interdisciplinary study, this analysis meets a series of scientific needs regarding foot morphology and the changes in its structure between populations. Aim: We analyzed the normal and pathological variability of the foot bones, emphasizing mainly on factors that alter its structure. Moreover, we considered the particular features of each collection and their anatomical changes across time. Materials and methods: To carry out our study, we used 890 samples that correspond to pairs of skeletonized feet of contemporary and ancient osteological collections from Spain, Italy, and Oman with different dating. We applied morphological and metric methods to check their differences, with different purposes. In addition, we used exploratory exams