Associations between Skeletal Fractures and Locomotor Behavior, Habitat Use, and Body Mass in Nonhuman Primates DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Heather Jarrell, B.A., MSc Graduate Program in Anthropology The Ohio State University 2011 Dissertation Committee: W. Scott McGraw, Advisor Clark Larsen Jeffrey McKee Sam Stout Charlotte Roberts Copyright by Heather Jarrell 2011 ABSTRACT Injuries sustained during falls from heights may be the most frequent cause of long bone trauma among nonhuman primates. Yet the interrelatedness of trauma and positional behaviors is poorly understood. The purpose of this study is to assess how common locomotor behaviors, broad habitat use tendencies, and body mass are associated with skeletal fracture frequencies in primates. Primates exhibiting a higher degree of arboreality should exhibit greater fracture frequencies than more terrestrial primates due to their greater risk of obtaining injuries in falls from heights. Similarly, primates whose locomotor repertoire includes more specialized behaviors should exhibit higher frequencies than those of more generalized quadrupeds. Falls sustained by primates commonly active higher in the canopy and larger primates should be more likely to show severe repercussions (as depicted by higher fracture frequencies) than primates active closer to the ground and smaller primates. Fractured long bones from primates encompassing twenty-two taxonomic groups housed at The Ohio State University, Cleveland Museum of Natural History, American Museum of Natural History, National Museum of Natural History, and the Caribbean Primate Research Center were examined macroscopically and radiographically. Locomotor and habitat use profiles for each taxonomic group were developed based upon quantitative field observations in the primatological literature. ii Fracture patterns appear to be most closely associated with locomotor mode, followed by arboreality, vertical distribution, and body mass, although each of these variables are interconnected. Locomotor mode preferences strongly correlate with fracture frequencies. As expected, suspensory primates exhibit the highest fracture frequencies, although leapers have the lowest frequencies. The locations of fractures are significantly correlated with some aspect of every variable examined. When suspensory primates break a bone, it tends to be either the humerus or femur. Small- or medium- sized, arboreal quadrupeds are more likely to fracture their tibia or fibula than another long bone, whereas large and very large quadrupeds tend to fracture any long bone preferentially except for the tibia or fibula. Fracture occurrences in leaping primates tend either to involve the clavicle preferentially or are independent of location. Factors contributing to intraspecific variation in fracture frequencies, including the impact of sex, age, and changing population pressures, may contribute to differential fracture distributions. Combining skeletal samples with behavioral observations from individuals‟ life histories highlights the under-representation of fractures in field studies as well as over-estimates of the degree to which fractures impair mobility. The fact that fracture frequencies and patterns revealed in this study are associated with locomotor and positional behaviors appears to highlight the importance of risk avoidance in primate evolution. Skeletal trauma may affect reproductive fitness either directly or incidentally. Consequently, primates should be under selective pressure to avoid the risk of obtaining fractures, developing behavioral and anatomical mechanisms to reduce the number and severity of falls from heights. Further analyses of the anatomical distribution of fractures in primates may reveal selective factors shaping iii locomotor anatomy. Ultimately, the analysis of fractures in primates which have healed under natural conditions may lead to a greater understanding of the origins of medical intervention in humans. iv Dedicated to Mom and Dad. v ACKNOWLEDGMENTS I would like to express my appreciation to all the individuals who have helped make the completion of this research possible, especially the members of my committee: Drs. Scott McGraw, Clark Larsen, Sam Stout, and Jeff McKee from the Ohio State University and Dr. Charlotte Roberts from the University of Durham, United Kingdom. Dr. Paul Sciulli‟s input regarding statistical analyses has been invaluable. Dr. Donna McCarthy-Beckett was my Graduate Faculty Representative and I thank her for her time and imput. Special thanks go to my advisor, Dr. McGraw, for all of his guidance throughout my tenure at OSU. Thank you for everything. Financial support was partially provided by the Ohio State University Alumni Grant for Graduate Research and Scholarship (AGGRS). I would like to thank Dr. Larsen for use of the NOMAD Portable X-ray system, provided through an equipment purchase via the Larsen Research and Travel Fund. Research at the Laboratory for Primate Morphology and Genetics was supported by the National Institutes of Health (grant # P40 RR003640) to the Caribbean Primate Research Center. I would like to thank all of the institutions which have housed the skeletal collections analyzed in this study. Lyman Jellema (Department of Physical Anthropology at the Cleveland Museum of Natural History [CMNH]), Darren Lunde and Eileen Westwig (Department of Mammalogy at the American Museum of Natural vi History [AMNH]), Linda Gordon (Department of Mammals at the National Museum of Natural History [USNM]), Dr. Edmondo Kraiselburg (Director of the Caribbean Primate Research Center [CPRC]), Dr. Donald Dunbar (Director of the Laboratory for Primate Morphology and Genetics [LPMG]), and Dr. Scott McGraw (OSU) all provided access to their respective collections. Dr. Christopher Ruff (Johns Hopkins University School of Medicine) and Evan Garofalo‟s assistance in transporting the NOMAD X-ray system and providing instruction in its use has been much appreciated. Lyman Jellema (CMNH), Eileen Westwig (AMNH), Dr. Barbara Brown (Department of Ichthyology at the AMNH), and Jeremy Jacobs (Division of Mammals at the USNM) all provided use of and/or assistance with radiographic equipment and darkroom facilities at their respective institutions. I am grateful for the efforts of everyone involved in the running and maintenance of all the divisions of the Caribbean Primate Research Center. I am especially thankful both to Dr. Dunbar, for all of the assistance he provided both during and after the application process, and to Dr. Terry Kensler (Lab Manager of the LPMG), for going above and beyond in making sure not only that I had everything I needed at the lab but also that I was fitting in well in Puerto Rico – even gifting me with a birthday cake! I would like to thank Myriam Vi ales, Dr. Bob Kensler, Dr. Jean Turnquist, and Pizarro for their support at the LPMG. I also would like to thank Dr. Janis Gonzalez and Dr. H ctor P rez for the information they provided at the Sabana Seca Field Station. Thanks go to James Ayala (Colony Manager of the Cayo Santiago Field Station [CSFS]) and Dr. Adaris Mas-Rivera (Resident Scientist at the CSFS), for allowing my visit to the Cayo vii Santiago breeding colony. Census Takers Edgar Davila and Giselle Caraballo provided valuable insight into the behaviors of the current and past CSFS colony. Finally, thanks go to my family and friends, most of all to my parents, J.W., Shelly, and Tommy, who took it in turns to placate and berate, empathize and question as the situation called for it, and who believe in me, always. viii VITA 2001 ...............................................................B.A. Scholars Anthropology, Northwestern State University 2003 ...............................................................MSc Human Osteology & Palaeopathology, University of Bradford, United Kingdom 2000 to 2002 .................................................Field Crew and Geophysical Technician, Louisiana National Guard and National Forest Surveys 2001 to 2002 .................................................Laboratory Supervisor, Cultural Resource Office, Northwestern State University 2004 to 2009 .................................................Graduate Teaching Associate, Department of Anthropology, The Ohio State University 2009 to 2010 .................................................Lecturer, Department of Anthropology, The Ohio State University ix PUBLICATION Jarrell H, Hailey TI. 2002. The Fort Jesup Hospitals: Historical, Geophysical, and Archaeological Investigations. In: Morgan N, Shatwell J, editors. Archaeology, Interpretation, and Management in Cane River National Heritage Area. Natchitoches: Cane River National Heritage Area Commission. p 32-44. FIELD OF STUDY Major Field: Anthropology Minor Field: Anatomy x TABLE OF CONTENTS ABSTRACT……………………………………………………………………....……………….ii DEDICATION………………..………………...……………………………………………….v ACKNOWLEDGEMENTS…………………………………….……………………………………………………….vi VITA………………………………………..………………..………………….…………………………….ix LIST OF TABLES…………………………………………...…………………………………………….xiv LIST OF FIGURES…………………………………..…………………………………………………….xvi 1 CHAPTER 1: Introduction……………………………………………………………….1 1.1 Project summary……………………………………………………………….1 1.2 Hypotheses……………………………………………………………………...……………….4 2 CHAPTER 2: Research Background……………………………………………………………….11 2.1 Bone and fracture biology……………………………………………………………….11 2.1.1 Biomechanical characteristics