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The Pennsylvania State University The Graduate School College of Earth and Mineral Sciences MOLECULAR AND ISOTOPIC PERSPECTIVES ON EARLY HUMAN HABITATS AT OLDUVAI GORGE, TANZANIA A Dissertation in Geosciences and Biogeochemistry by Clayton R. Magill © 2013 Clayton R. Magill Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy May 2013 The dissertation of Clayton R. Magill was reviewed and approved* by the following: Gail M. Ashley Professor of Earth and Planetary Sciences, Rutgers University Special Signatory Katherine H. Freeman Professor of Geosciences Dissertation Advisor Co-Chair of Committee Nina G. Jablonski Distinguished Professor of Anthropology Lee R. Kump Professor of Geosciences Head of the Department of Geosciences Co-Chair of Committee Mark E. Patzkowsky Associate Professor of Geosciences *Signatures are on file in the Graduate School ii ABSTRACT Early human (hominin) evolution was linked to dramatic changes in regional hydrology and ecosystem composition. Emerging hypotheses linking hominin evolution and the environment present specific, testable predictions for hominin evolutionary responses to different modes of environmental change. Yet, discontinuous terrestrial sediment sequences coupled with indirect proxy indicators for plants and water obscure environmental perspectives on the hominin fossil record. Opportunity to overcome these challenges is presented by lake and soil sediments from a preeminent hominin archaeological locality – Olduvai Gorge, Tanzania. These extensive, well- constrained sediments have abundant organic matter, allowing for unparalleled resolution of biomarker data through time and space for an individual locality. To link the hominin archaeological record to local environmental conditions at Olduvai Gorge, I characterized distributions and stable isotope compositions (δ13C and δD) for a diverse suite of modern and sedimentary biomarkers. Towards this end, I established a quantitative framework for reconstructing plant functional type (PFT) relative abundances in tropical ecosystems based on biomarker δ13C values from living plants by compiling previously published data relating (a) PFT to tropical ecosystem structure, (b) PFT to soil organic matter δ13C values, and (c) soil organic matter δ13C values to biomarker δ13C values. Separation of previously published δD values for biomarkers from living plants according to PFT can elucidate differences in the biochemical and physiological influences on apparent fractionation between source-waters and biomarkers (εlipid/water). By extension, ‘landscape’ apparent fractionation factors are calculated 13 from isotopic mass-balance of εlipid/water values via biomarker δ C estimates for relative PFT iii abundances. Using this novel approach to establish fractionation factors in dynamic environments, I then reconstructed δD values for precipitation and lake waters at Olduvai Gorge during the past. The early Pleistocene is associated with a key adaptive juncture in hominin evolution – the emergence of our direct ancestor Homo erectus/ergaster. To constrain local environmental conditions during this juncture, I measured δ13C and δD values for sedimentary biomarkers from a continuous sequence of lake sediments deposited between about 2.0 and 1.8 million years ago (i.e., the early Pleistocene) at Olduvai Gorge. Biomarker δ13C values correlate strongly with changes in orbital geometry and tropical sea-surface temperatures during this interval, and reveal cyclic catchments-scale ecosystem shifts between closed woodlands and open grasslands. After correcting measured biomarker δD values for ‘landscape’ apparent fractionation factors, reconstructed source-water δD values indicate lower amounts of precipitation fell on open grasslands (about 250 mm yr-1) as compared to closed woodlands (about 700 mm yr-1). The scale and pace of environmental changes at Olduvai Gorge contrast with long-held views of directional or step-wise aridification in eastern Africa during the early Pleistocene. Faunal evolutionary responses to environmental change are strongly influenced by microhabitat (<100 m2) features. To reconstruct fine-scale spatial heterogeneity in plants and water associated with hominin habitation, we measured the distributions and δ13C values for plant biomarkers (leaf-waxes, lignin and 5-n-alkyresorinols) preserved in time-equivalent soil sediments across the iconic FLK Zinjanthropus archaeological Level 22 (FLK Zinj) locality at Olduvai Gorge. Central trenches of FLK Zinj are associated with low δ13C values for leaf-waxes and lignin iv monomers indicative of closed woodland habitat. About 200 m to the north of the central trenches, abundant aquatic-plant and sedge biomarkers occur in conjunction with mound-like tufa deposits, suggesting wetland habitat near freshwater springs. In contrast, southern trenches contain high δ13C values for leaf-waxes and lignin monomers indicative of open grassland habitat. Taken together, these data delimit a heterogeneous microhabitat mosaic that is obscured in catchment-scale records of environmental change. v TABLE OF CONTENTS LIST OF TABLES ..................................................................................................................... xi LIST OF TABLES ..................................................................................................................... xiv ACKNOWLEDGMENTS .......................................................................................................... xv CHAPTER 1: Introduction 1.1. Introduction ............................................................................................................ 1 1.2. Environmental hypotheses of hominin evolution ................................................... 2 1.3. Records of environmental change for eastern Africa ............................................. 3 1.4. The rise of arid ecosystems in Africa ..................................................................... 3 1.5. Organic matter signals for ecosystem ..................................................................... 4 1.6. Organic geochemical proxies for hydroclimate ...................................................... 5 1.7. Isotopic notation ..................................................................................................... 6 1.8. Research objectives and dissertation outline .......................................................... 7 1.9. Communications and publications from this dissertation ...................................... 9 1.10. References ............................................................................................................. 10 CHAPTER 2: Ecosystem variability and early human habitats in eastern Africa 2.1. Abstract ................................................................................................................... 12 2.2. Introduction ............................................................................................................. 13 2.3. Background ............................................................................................................. 14 2.3.1. Site descriptions ............................................................................................ 14 2.3.2. Sedimentary organic matter .......................................................................... 16 2.3.3. Carbon isotopes in leaves, biomarkers and soil organic matter ................... 16 2.3.4. Structural classification for ancient ecosystems ........................................... 18 2.4. Results and discussion ............................................................................................ 19 2.4.1. Ecosystem change and woody cover ............................................................ 19 2.4.2. Biogeochemical variability at the ecosystem scale ...................................... 21 2.4.3. Mechanisms of ecosystem change ................................................................ 22 2.4.4. Mechanisms of hydroclimate change ........................................................... 23 2.4.5. Ecosystems and hominin evolution .............................................................. 25 2.5. Conclusions ............................................................................................................. 25 2.6. Figures .................................................................................................................... 27 2.7. References ............................................................................................................... 33 vi CHAPTER 3: Water, plants and early human habitats in eastern Africa 3.1. Abstract ................................................................................................................... 39 3.2. Introduction ............................................................................................................. 40 3.3. Background ............................................................................................................. 41 3.3.1. Sample locality ............................................................................................. 41 3.3.2. Precipitation patterns in eastern Africa ........................................................ 41 3.3.3. Leaf-lipid apparent fractionation factors ...................................................... 42 3.3.4. Algal-lipid apparent fractionation factors ..................................................... 45 3.4. Results ....................................................................................................................