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Patterns of Morphological Integration in Modern Human Crania: Evaluating Hypotheses of Modularity Using Geometric Morphometrics

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Patterns of Morphological Integration in Modern Human Crania: Evaluating Hypotheses of Modularity Using Geometric Morphometrics PATTERNS OF MORPHOLOGICAL INTEGRATION IN MODERN HUMAN CRANIA: EVALUATING HYPOTHESES OF MODULARITY USING GEOMETRIC MORPHOMETRICS DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Adam Kolatorowicz Graduate Program in Anthropology The Ohio State University 2015 Dissertation Committee: Jeffrey K. McKee, Advisor Paul W. Sciulli Samuel D. Stout Mark Hubbe Copyrighted by Adam Kolatorowicz 2015 ABSTRACT This project examines patterns of phenotypic integration in modern human cranial morphology using geometric morphometric methods. It is theoretically based in the functional paradigm of craniofacial growth and morphological integration. The hypotheses being addressed are: 1) cranial form is influenced by secular trends, sex, and phylogenetic history of the population and 2) integration patterns wherein the basicranium is the keystone feature best explains the relationships among in cranial modules. Geometric morphometric methods were used to collect and analyze three- dimensional coordinate data of 152 endocranial and ectocranial landmarks from 391 anatomically modern human crania. These crania are derived from temporally historic and recent groups in the United States spanning both sexes and across several ancestral groups. Landmark data were subjected to generalized Procrustes analysis and then areas of shape variation were identified via principal components analysis of shape coordinates. Discriminant function analysis and canonical variate analysis identified regions that can be used to separate groups. Temporal period, ancestry, and sex all have significant effects on mean shape. Age-at-death accounts for a small proportion of the total variation. Modern individuals have higher, narrower vaults with highly arched palates ii and historic individuals have short, wider vaults with shallower palates. The forehead, brow ridges, and cheek shape were closely associated with sexual dimorphism. Variation in both the vault and face allowed for separation of ancestral groups with concomitant inferior movement of the anterior basicranium in the median plane. Three major hypotheses of modularity were tested based on functional demands of cranial modules, functional-developmental fields, and the basicranium. Comparing covariance structures of partitions of landmark subsets revealed that the cranium is more integrated when considering functional demands of cranial components origins of cranial components and is less modular when considering developmental origins. Special sensory modules are the most independent units in the cranium. Depending on the definition of cranial modules, results may be quite different and not comparable across studies. This project integrates anthropology, evolutionary anatomy, and developmental biology. It makes a significant contribution to our understanding of integration patterns in the modern human cranium and highlights differences among theoretical frameworks of integration. The findings can be used for individual identification in medicolegal contexts and clinical applications for surgical treatment of craniofacial-related disorders and injuries. Future research will include examining patterns of morphological integration in non-human primates. iii Dedicated to my parents. iv ACKNOWLEDGEMENTS I would like to acknowledge the faculty and staff at multiple institutions for the role they played in making this dissertation possible. First and foremost, my dissertation committee chair Dr. Jeff McKee and committee members Dr. Sam Stout, Dr. Paul Sciulli, and Dr. Mark Hubbe in the Department of Anthropology at The Ohio State University provided thoughtful comments and constructive criticism throughout the process. Dr. Mike Warren and Mr. Carlos Zambrano from the C.A. Pound Human Identification Lab at the University of Florida – Gainesville graciously let me borrow their lab’s digitizer. It is good to know people in high places when funding does not come through. Dr. Amanda Agnew in the Division of Anatomy at the Ohio State University granted access to the skeletal teaching collection and workspace used for the observer error study. Dr. Tim Gocha was instrumental in establishing working relationships in the Division of Anatomy. Thanks to Mr. Lyman Jellema, Collections Manager of the Hamann-Todd Osteological Collection at the Cleveland Museum of Natural History, for being a most gracious host while I collected data. Dr. Sabrina Curran helped to break up the monotony of data collection with some entertaining and fruitful conversations about anthropology, evolution, and methodology. Dr. Dawnie Wolfe Steadman facilitated access to the W. M. Bass Donated Skeletal Collection at the University of Tennessee – Knoxville and Ms. v Heli Maijanen provided logistical support while I collected data. Finally, I appreciate the patience and knowledge base of staff members in the Department of Anthropology at Ohio State. Ms. Elizabeth Freeman, Mr. Wayne Miller, and Ms. Jean Whipple offered administrative and logistical support during many steps while I completed this project. I am indebted to my family and friends who supported me through this journey. In particular, I am grateful for the comradery fostered by The Juices of Fire, the collegiality of fellow graduate students at Ohio State, and the support system created by Ms. Cheryl Lynn Wyckoff. Above all, I would like to thank my Dad. He was a true renaissance man who sought to better understand the world around him. He had broad interests in art, science, and technology with formal and informal study in astronomy, biology, culture, chemistry, computer science, engineering, history, mathematics, paleontology, and physics. As a jack of all trades and master of many, he inspired me to be curious about the world. Although he is no longer with us to read this dissertation I know that he would appreciate my contribution to our understanding of the pale blue dot we inhabit. vi VITA 2000................................................................A.A. Anthropology, College of Lake County 2002................................................................B.S. Anthropology, Northern Illinois University 2003 to 2004 .................................................Graduate Teaching Assistant, Department of Biology, University of Indianapolis 2004 to 2006 .................................................Adjunct Instructor, Department of Biology, University of Indianapolis 2006................................................................M.S. Human Biology, University of Indianapolis 2006 to 2007 ..................................................Graduate Teaching Associate, Department of Anthropology, The Ohio State University 2008................................................................Graduate Research Associate, Department of Anthropology, The Ohio State University 2008 to 2013 ..................................................Graduate Teaching Associate, Department of Anthropology, The Ohio State University 2011................................................................Fellow, Forensic Science Academy, Joint POW/MIA Accounting Command – Central Identification Laboratory, Joint Base Pearl Harbor – Hickam 2010 to present ...............................................Adjunct Instructor, Department of Social Science, Department of Biological and Physical Sciences, Columbus State Community College 2014................................................................Lecturer, Department of Anthropology, The Ohio State University vii 2014 to present ..............................................Adjunct Instructor, Department of Sociology and Anthropology, Ohio University PUBLICATIONS Gocha TP, Ingvoldstad ME, Kolatorowicz A, Cosgriff-Hernandez MTJ, Sciulli PW. 2015. Testing the applicability of six macroscopic skeletal aging techniques on a modern Southeast Asian sample. Forensic Sci Int 249:318.e1-7. FIELDS OF STUDY Major Field: Anthropology Minor Field: Anatomy Minor Field: Statistical Data Analysis viii TABLE OF CONTENTS ABSTRACT ........................................................................................................................ ii DEDICATION ................................................................................................................... iv ACKNOWLEDGEMENTS .................................................................................................v VITA ................................................................................................................................. vii TABLE OF CONTENTS ................................................................................................... ix LIST OF TABLES ............................................................................................................ xii LIST OF FIGURES ......................................................................................................... xiii CHAPTER 1: INTRODUCTION .......................................................................................1 1.1 PURPOSE ............................................................................................................4 1.2 SIGNIFICANCE OF RESEARCH ......................................................................5 1.3 HYPOTHESES ....................................................................................................7 1.4 ORGANIZATION OF DISSERTATION ............................................................8 1.5 SUMMARY .........................................................................................................9
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