FUNCTIONAL MORPHOLOGY, VARIATION, AND NICHE SEPARATION IN THE LARGE-BODIED FOSSIL COLOBINES by MONYA ANDERSON A DISSERTATION Presented to the Department of Anthropology and the Graduate School of the University of Oregon in partial fulfillment of the requirements for the degree of Doctor of Philosophy June 2019 DISSERTATION APPROVAL PAGE Student: Monya Anderson Title: Functional Morphology, Variation, and Niche Separation in the Large-Bodied Fossil Colobines This dissertation has been accepted and approved in partial fulfillment of the requirements for the Doctor of Philosophy degree in the Department of Anthropology by: Stephen Frost Chairperson Frances White Core Member Nelson Ting Core Member Samantha Hopkins Institutional Representative and Janet Woodruff-Borden Vice Provost and Dean of the Graduate School Original approval signatures are on file with the University of Oregon Graduate School. Degree awarded June 2019 ii © 2019 Monya Anderson iii DISSERTATION ABSTRACT Monya Anderson Doctor of Philosophy Department of Anthropology June 2019 Title: Functional Morphology, Variation, and Niche Separation in the Large-Bodied Fossil Colobines Colobines are ecological referents that can be used to assess and reconstruct paleoenvironmental conditions. Colobine taxa discussed here include Paracolobus mutiwa, P. Cercopithecoides williamsi, C. kimeui, and Rhinocolobus turkanensis. All of these taxa except for R. turkanensis display postcranial adaptations consistent with terrestrial locomotion in contrast to their exclusively arboreal African counterparts. At sites like these of the Upper Burgi Member of Koobi Fora, these four large colobines, up to four species of hominins, and three large-bodied cercopithecine taxa are known. This level of sympatry in primate communities is unmatched in modern habitats emphasizing the importance of niche separation for understanding diversity in the fossil record. The descriptions of two previously undescribed specimens: P. mutiwa specimen KNM-WT 16827 and the informally designated P. mutiwa specimen L895-1are presented here with detailed qualitative and quantitative postcranial analyses. Postcranial measurements from fossil colobines, fossil cercopithecines, and a large extant sample for comparative purposes. Measurements include 112 linear metrics and 54 functional indices. Body mass estimates for the fossil descriptions were calculated based on published equations for postcranial estimation. Dietary proxies for the fossil taxa are based on dental iv morphology and taken from the literature. Substrate preference is estimated using qualitative description of skeletal elements, quantitative comparison to extant taxa, and observational data of substrate use of extant species. Paracolobus mutiwa specimen KNM-WT 16827 shows postcranial morphologies consistent with terrestriality distinct from Cercopithecoides and Theropithecus and is distinct from P. chemeroni to a degree warranting generic reassessment of the species. Specimen L895-1 shows postcranial morphology consistent with terrestriality, is distinct from other contemporaneous cercopithecids, and is most similar in size and morphology to P. mutiwa. The niche separation analyses show C. williamsi to be the most terrestrial of the large colobines with the largest consumption of leaves. P. mutiwa overlaps with C. williamsi in size, but is less terrestrial and R. turkanensis is the least. There is also clear separation among the large colobines and Theropithecus in diet and substrate use. The sympatry and diversity of these large-bodied colobines suggests a much more diverse and dynamic primate environment than seen in East Africa today. v CURRICULUM VITAE NAME OF AUTHOR: Monya Anderson GRADUATE AND UNDERGRADUATE SCHOOLS ATTENDED: University of Oregon, Eugene, OR Purdue University, West Lafayette, IN DEGREES AWARDED: Doctor of Philosophy, Biological Anthropology, 2019, University of Oregon Master of Science, Biological Anthropology, 2012, University of Oregon Bachelor of Arts, Anthropology, 2010, Purdue University AREAS OF SPECIAL INTEREST: Biological anthropology, non-human and human primate fossil record, paleoanthropology, functional morphology, paleoenvironmental reconstruction, vertebrate paleontology PROFESSIONAL EXPERIENCE: Graduate Employee, Department of Anthropology, University of Oregon, 2010-2019 Instructor of Record, Department of Anthropology, University of Oregon, 2017-2019 GRANTS, AWARDS, AND HONORS: Graduate Teaching Fellowship, Department of Anthropology, University of Oregon, 2010-2019 Johnston Scholarship Graduate Research Fellowship, Department of Anthropology, University of Oregon, 2018 Departmental Nominee for Dissertation Research Fellowship, Department of Anthropology, University of Oregon, 2018 Doctoral Dissertation Improvement Grant, National Science Foundation, 2017 Graduate School Research Award, University of Oregon, 2013 vi Johnston Scholarship Graduate Research Fellowship, Department of Anthropology, University of Oregon, 2013 Departmental Travel Award, Department of Anthropology, University of Oregon, 2012 Departmental Travel Award, Department of Anthropology, University of Oregon, 2011 Outstanding Senior in Anthropology, Department of Anthropology, Purdue University, 2010 O. Michael Watson Anthropology Senior Capstone Award, Department of Anthropology, Purdue University, 2010 Dean's List, Purdue University, 2006-2010 University Honors, Purdue University, 2006-2010 College of Liberal Arts Honors, Purdue University, 2006-2010 College of Liberal Arts Dean's Scholar, Purdue University, 2006 National Honor Society, Terre Haute South Vigo High School, Terre Haute, IN, 2006 PUBLICATIONS: Anderson, M., 2019. An assessment of the postcranial skeleton of the Paracolobus mutiwa (Primates: Colobinae) specimen KNM-WT 16827 from Lomekwi, West Turkana, Kenya. Journal of Human Evolution (In Review) Anderson, M., Frost, S.R., 2016. Pedal Functional Morphology of Paracolobus chemeroni. American Journal of Physical Anthropology 159(S62), 80. Anderson, M., Frost, S.R., 2015. A Reassessment of the Phylogeography of Sulawesi macaques based on 3D geometric morphometrics, American Journal of Primatology 77, 49-50. Anderson, M., Frost, S.R., Gilbert, C.C., Delson, E., 2014. A reassessment of dental variation in the genus Cercopithecoides and its implications for species diversity. American Journal of Physical Anthropology. 153, 67. Anderson, M., Frost, S.R., Gilbert, C.C., Delson, E., 2013. Morphological diversity and species recognition in South African Cercopithecoides williamsi. American Journal of Physical Anthropology. 150, 68. vii Anderson, M., Frost, S.R., Gilbert, C.C., Delson, E., 2015. Cranial shape and intrageneric diversity in the genus Cercopithecoides. 156(S60), 69. Arenson, J.L., Anderson, M., Eller, A.R., Simons, E.A., White, F.J., Frost, S.R., 2015. Bergmann’s rule in skull size of wild vs. captive fascicularis group macaques. American Journal of Physical Anthropology. 156 (S60), 71. Arenson, J.L., Anderson, M., White, F.J., Frost, S.R., 2016. Morphological assessment of a putative hybrid species, Trachypithecus pileatus, based on a 3D geometric morphometric analysis of cranial morphology. American Journal of Physical Anthropology 159(S62), 82. Clarke, K.S., Anderson, M., Eller, A.R., Simons, E., White, F.J., Frost, S.F., 2015. Morphological signals of stress and socioendocrinology: Comparing measures of cranial fluctuating asymmetry and second to fourth digit ratio in cercopithecids. American Journal of Physical Anthropology 156, 106. Clarke, K.S., McNulty, K.P., Eller, A.R., Arenson, J.L., Anderson, M., Simons, E.A., White, F.J., Frost, S.R., 2015. Morphological signals and mating systems: Comparing measures of cranial fluctuating asymmetry and second-to-fourth digit ratio in anthropoid primates. American Journal Primatology. 77(S1), 56. Gilbert, C.C., Pugh, K., Anderson, M., Frost, S.R., Delson, E., 2018. Evolution of the modern baboon (Papio hamadryas): A reassessment of the African Plio-Pleistocene record. Journal of Human Evolution 122, 38-69. Glenzer, R.R., Clarke, K.S., Anderson, M., Eller, A.R., White, F.J., Frost, S.R., 2015. Comparing morphometric methods in Macaca mulatta crania. American Journal of Physical Anthropology. 156(S60), 146. Reda, H., Frost, S.R., Simons, E., Anderson, M., Haile-Selassie, Y., Preliminary Study of the Cercopithecidae from Leado Didoa locality, Woranso-Mille (central Afar), Ethiopia. American Journal of Physical Anthropology 162(S64), 328-329. viii ACKNOWLEDGMENTS This dissertation is the result of many years of patience, support, and collaborative effort. I have been very fortunate over the years to have had the encouragement of dozens of teachers, colleagues, friends, and family. I would first like to thank my amazing advisor Stephen Frost for his intellectual guidance, fieldwork opportunities, an availability over the past nine years. Thank you for introducing me the amazing world of monkeys; I have enjoyed giving them their due and look forward to discovering more! A special thanks also to all the member of my committee: Frances White, Nelson Ting, and Samantha for your feedback, suggestions, and support over the years. Thank you. This project could not have been done without extensive fieldwork for which I would like to thank the curators, staff, and researcher as the Kenya National Museum (Emma Mbua, Frederick Manthi, Mzalendo Kibunjia, Rose Nyaboke, & Cecilia Kanyua), National Museum of Ethiopia (ARCCH, Getachew Senishaw, Tomas Getachew, Yohannes Haile-Selassie, Jean-Renaud