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EARLY HOMININ ENVIRONMENTS in SOUTHERN AFRICA: a MICROMAMMALIAN PERSPECTIVE by JENNIFER NICOLE LEICHLITER B.A, Colorado College

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EARLY HOMININ ENVIRONMENTS in SOUTHERN AFRICA: a MICROMAMMALIAN PERSPECTIVE by JENNIFER NICOLE LEICHLITER B.A, Colorado College EARLY HOMININ ENVIRONMENTS IN SOUTHERN AFRICA: A MICROMAMMALIAN PERSPECTIVE by JENNIFER NICOLE LEICHLITER B.A, Colorado College, 2008 M.A. University of Colorado Boulder, 2011 A thesis submitted to the Faculty of the Graduate School of the University of Colorado Boulder in partial fulfillment of the requirement for the degree of Doctor of Philosophy Department of Anthropology 2018 ii SIGNATURE PAGE This thesis entitled: Early Hominin Environments in Southern Africa: A Micromammalian Perspective Written by Jennifer Nicole Leichliter has been approved for the Department of Anthropology ____________________________________________ Dr. Matt Sponheimer, Committee Chair ____________________________________________ Dr. Nico Avenant ____________________________________________ Dr. Herbert Covert ____________________________________________ Dr. Jaelyn Eberle ____________________________________________ Dr. Joanna Lambert Date ___________ The final copy of this thesis has been examined by the signatories, and we find that both the content and the form meet acceptable presentation standards of scholarly work in the above mentioned discipline. iii ABSTRACT Leichliter, Jennifer Nicole (Ph.D., Anthropology) Early Hominin Environments in Southern Africa: A Micromammalian Perspective Thesis directed by Professor Matt Sponheimer Environmental change, especially the expansion of open C4 savanna grasslands at the expense of more heavily wooded C3 habitats, is frequently cited as a driver of hominin evolution. The disappearance of the genus Australopithecus, the rise of the cranio-dentally robust Paranthropus, and the emergence of Homo have all been linked to environmental change during the period from 3 and 2 Ma. The large mammal communities which co-existed with hominins are often used to reconstruct paleoenvironments during this period, but small mammals have been relatively underutilized despite their potential to provide information about hominin ecosystems at a fine scale. A primary goal of this dissertation was to determine whether patterns of evolutionary change observed in the large mammal fossil record of southern Africa during the Plio-Pleistocene are also evident in the small mammal record. A combination of stable carbon isotope analyses and taxon-dependent analyses were deployed, and in some cases significantly developed, in re-assessing the micromammalian fossil record associated with early hominins. Large gaps remain in our knowledge of small mammal isotopic ecology, especially in African ecosystems, where C4 plants are abundant and many landscapes relevant to hominin evolution persist. This dissertation focuses, in part, on understanding the relationship between small mammal stable carbon isotope composition and their habitats in a modern southern African C4 savanna and applying this information to the fossil record. It was determined that small mammal d13C compositions record habitat composition. iv However, insectivores appear to track proportions of C3/C4 vegetation better than rodents. Preliminary analyses of the d13C composition of fossil tooth enamel of small mammals from early hominin-bearing deposits in southern Africa suggest that C4 resources may have been more abundant in the past. Taxonomic-dependent faunal analyses indicate no clear shift from more mesic, closed habitats to more open-grassy habitats between 3 and 1 Ma in southern Africa, but instead suggest that the environment in this region was predominantly open with patches of mesic C3 woodland habitat. Australopithecus africanus and Paranthropus robustus appear to have been associated with both habitat types, perhaps taking advantage of resource-rich C3 habitat patches whenever they were available. v TABLE OF CONTENTS 1 Introduction ........................................................................................................................ 1 Background ................................................................................................................. 1 Chapter Summaries ..................................................................................................... 9 2 Small mammal insectivore stable carbon isotope composition as a proxy for habitat ......... 12 Introduction ............................................................................................................... 12 Materials and Methods .............................................................................................. 19 2.2.1 Field Collection Methods ....................................................................... 19 2.2.2 Stable Isotope Analyses .......................................................................... 22 Results ...................................................................................................................... 24 Discussion ................................................................................................................. 30 Conclusion ................................................................................................................ 36 3 Stable carbon isotope ecology of small mammal tooth enamel from barn owl roosts in the Sterkfontein Valley ................................................................................................................... 37 Introduction ............................................................................................................... 37 Materials and Methods .............................................................................................. 40 3.2.1 Isotopic Analyses ................................................................................... 46 Results and Discussion .............................................................................................. 49 3.3.1 Incisor-Molar Pairs................................................................................. 49 3.3.2 Modern Roost Site Comparisons ............................................................ 52 3.3.3 Relative Abundance for Modern Roost Sites .......................................... 58 3.3.4 Fossil Analyses ...................................................................................... 62 Conclusions ............................................................................................................... 67 4 Micromammal Fossil Descriptions .................................................................................... 69 Introduction ............................................................................................................... 69 Site Background Information ..................................................................................... 71 4.2.1 Kromdraai B .......................................................................................... 71 4.2.2 Sterkfontein Member 4 ........................................................................... 73 4.2.3 Swartkrans Member 1 (Hanging Remnant) ............................................. 73 4.2.4 Gladysvale External Deposit .................................................................. 74 Systematic Paleontology ............................................................................................ 75 5 African Small Mammal Taxonomic Habitat Index .......................................................... 101 Introduction ............................................................................................................. 101 Development of the taxonomic habitat index ........................................................... 102 Evaluating the taxonomic habitat index ................................................................... 107 5.3.1 Biozones .............................................................................................. 107 5.3.2 Roost Sites ........................................................................................... 112 6 Paleoecological Analyses ................................................................................................ 120 Introduction ............................................................................................................. 120 6.1.1 Assumptions ........................................................................................ 124 Materials and Methods ............................................................................................ 126 6.2.1 Species Richness and Diversity ............................................................ 127 6.2.2 Similarity Indices ................................................................................. 129 6.2.3 Taxonomic Ratios ................................................................................ 129 6.2.4 Correspondence Analysis ..................................................................... 130 vi 6.2.5 Taxonomic Habitat Indices ................................................................... 131 Results and Discussion ............................................................................................ 132 6.3.1 Rarefaction ........................................................................................... 132 6.3.2 Shannon-Weiner Index, Evenness, Simpson’s Dominance ................... 134 6.3.3 Similarity Indices ................................................................................. 140 6.3.4 Taxonomic Ratios ................................................................................ 142 6.3.5 Correspondence Analysis ..................................................................... 144 6.3.6 Habitat Reconstructions
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