Isotopic records of ecological variability in modern and ancient environments in Kenya

Stable isotope analysis of biological and geological materials has provided important information on environmental change over many timescales. Such records rely on an interpretive framework of large isotopic datasets of recent (i.e., modern and/or Holocene-aged) material in order to define paleoecological interpretations. This dissertation will contribute to a better understanding of ancient and historical ecological change in East Africa by (1) providing a modern interpretive framework for a megaherbivore, Hippopotamus amphibius, in East Africa and (2) investigating the relationship between climate and ecology in Kenya during the Holocene using multiproxy records, including stable isotopes of large mammalian herbivores and leaf wax biomarker isotopes. Although H. amphibius (hippo) tooth enamel isotopes are widely used to understand paleoenvironment, little is known about their dietary variability across time and space. In this dissertation, I present an extensive study on stable isotope serial samples (δ13C and δ18O) of modern hippos in order to understand the breadth of their dietary and behavioral flexibility in Kenya. This large dataset (10 mm interval samples of canine tooth enamel for 30 hippo canines; 1,410 samples in total) reveals the remarkable dietary diversity of hippos: they are indiscriminate feeders and can consume both C3 and C4 herbaceous forage. Furthermore, certain profiles with known death dates have captured isotopic indicators of ecological perturbations, such as drought and C3 plant encroachment following elephant extirpation. Multiproxy records of ecology can provide even more paleoecological information than single-mammal records. Assumptions about environmental change in the fossil record are often based on inference from known global climate and presumed changes in ecology. Using a multiproxy approach (tooth enamel isotopes and leaf wax biomarkers), I present Holocene (11.8 Kya - present) paleoecological data from two basins: the arid Turkana Basin in northern Kenya and the more mesic Victoria Basin in southwestern Kenya. These data indicate that as the monsoon was weakening due to global climatic change (i.e, Milankovitch forcing), there was an increase in C3 resources (bush, shrub, trees, and herbs) in Turkana, whereas Lake Victoria remained predominately C4 (tropical lowland grasses). This interbasinal record reveals that we cannot make assumptions about changes in ecology in an entire region based on climatological forcing mechanisms alone.

ISOTOPIC RECORDS OF ECOLOGICAL VARIABILITY IN MODERN AND ANCIENT ENVIRONMENTS IN KENYA by Kendra Lorraine Chritz A dissertation submitted to the faculty of The University of Utah in partial fulfillment of the requirements for the degree of Doctor of Philosophy Department of Biology The University of Utah December 2015 Copyright © Kendra Lorraine Chritz 2015 All Rights Reserved The University of Utah Graduate School STATEMENT OF DISSERTATION APPROVAL The dissertation of Kendra Lorraine Chritz has been approved by the following supervisory committee members: Thure E. Cerling , Chair 8/27/2015 Date Approved James Ehleringer , Member 8/27/2015 Date Approved Katherine H. Freeman , Member 10/14/2015 Date Approved M. Denise Dearing , Member 8/27/2015 Date Approved Francis H. Brown , Member 8/27/2015 Date Approved and by M. Denise Dearing the Department/College/School of and by David B. Kieda, Dean of The Graduate School. , Chair/Dean of Biology ABSTRACT Stable isotope analysis of biological and geological materials has provided important information on environmental change over many timescales. Such records rely on an interpretive framework of large isotopic datasets of recent (i.e., modern and/or Holocene-aged) material in order to define paleoecological interpretations. This dissertation will contribute to a better understanding of ancient and historical ecological change in East Africa by (1) providing a modern interpretive framework for a megaherbivore, Hippopotamus amphibius, in East Africa and (2) investigating the relationship between climate and ecology in Kenya during the Holocene using multiproxy records, including stable isotopes of large mammalian herbivores and leaf wax biomarker isotopes. Although H. amphibius (hippo) tooth enamel isotopes are widely used to understand paleoenvironment, little is known about their dietary variability across time and space. In this dissertation, I present an extensive study on stable isotope serial samples (δ13C and δ18O) of modern hippos in order to understand the breadth of their dietary and behavioral flexibility in Kenya. This large dataset (10 mm interval samples of canine tooth enamel for 30 hippo canines; 1,410 samples in total) reveals the remarkable dietary diversity of hippos: they are indiscriminate feeders and can consume both C3 and C4 herbaceous forage. Furthermore, certain profiles with known death dates have captured isotopic indicators of ecological perturbations, such as drought and C3 plant encroachment following elephant extirpation. Multiproxy records of ecology can provide even more paleoecological information than single-mammal records. Assumptions about environmental change in the fossil record are often based on inference from known global climate and presumed changes in ecology. Using a multiproxy approach (tooth enamel isotopes and leaf wax biomarkers), I present Holocene (11.8 Kya – present) paleoecological data from two basins: the arid Turkana Basin in northern Kenya and the more mesic Victoria Basin in southwestern Kenya. These data indicate that as the monsoon was weakening due to global climatic change (i.e, Milankovitch forcing), there was an increase in C3 resources (bush, shrub, trees, and herbs) in Turkana, whereas Lake Victoria remained predominately C4 (tropical lowland grasses). This interbasinal record reveals that we cannot make assumptions about changes in ecology in an entire region based on climatological forcing mechanisms alone. iv TABLE OF CONTENTS ABSTRACT……………………………………………………………..……………….iii LIST OF FIGURES…………………………….…………………….….………....……vii LIST OF TABLES.……………………..……………………………………..……..……x ACKNOWLEDGMENTS………………………………………..…..………….…...…xiii Chapters 1. INTRODUCTION…………………………..…………….…………………….....1 1.1. Background…………………………………………..………..……..1 1.2. Chapter Summaries…………………………………..…..….……….6 1.3. References……………………………..………………..…..………..9 2. ISOTOPIC COMPOSITION OF WILD H. AMPHIBIUS CANINES REVEALS NONSELECTIVE FEEDING BEHAVIOR IN KENYA………..………16 2.1. Abstract……………………………………………..………..……..16 2.2. Introduction…………………………………...…...………………..17 2.3. Materials and Methods……………………….....…………………..18 2.4. Results………………………...………...…………………………..24 2.5. Discussion…………………………………………..…………...….25 2.6. Conclusions…...………………………………..…...…………..…..35 2.7. Acknowledgments.…………………..……………...…….……..….35 2.8. References…………………………..…..……………….……….....36 3. DECADAL DIET CHANGE IN HIPPOPOTAMUS AMPHIBIUS IN QUEEN ELIZABETH NATIONAL PARK, UGANDA……………..….………..56 3.1. Abstract………………………………………………..……..……..56 3.2. Introduction……………………………………….…………..…….57 3.3. Materials and Methods………………………………………..…….63 3.4. Results…………………….………………………………….….….64 3.5. Discussion………………………………………..……………...….65 3.6. Conclusions………...………………………..…………………..….68 3.7. Acknowledgments.…………………………..…………………..….68 3.8. References………………………………………………….……….69 4. HOLOCENE PALEOENVIRONMENT IN KENYA, WITH PERSPECTIVES ON HOLOCENE ARCHAEOLOGY AND PLEISTOCENE ENVIRONMENTAL CHANGE……………………...……………….…79 4.1. Abstract………….…………………………………………..…..…..79 4.2. Introduction………………………….…………….…………..…….80 4.3. Study Design and Location…….……………………...…………….85 4.4. Materials and Methods………………….…………………..……….86 4.5. Results……………….……………………..…………………….….88 4.6. Discussion……………………….…………..………………........…91 4.7. Conclusions………...………….………………..……………….…100 4.8. Acknowledgments.……….………………………..………...…..…101 4.9. References………………………………………...…..…….…...…101 5. ENVIRONMENTS AND TRYPANOSOMAISIS RISKS FOR EARLY HERDERS IN THE LATER HOLOCENE OF THE LAKE VICTORIA BASIN, KENYA………….………….………….……………...………130 5.1. Abstract……….……………………………………..……………..130 5.2. Introduction…………….…………………………...………..…….130 5.3. Materials and Methods….……………………………………....….133 5.4. Results…………….………………..…………………………...….132 5.5. Discussion…………………….………..……………………......…132 5.6. Conclusions………………………………..…………………….…133 5.7. Acknowledgments.…….……………………..……………...…..…134 5.8. References…………………………….…………………….…...…134 5.9. SI Appendix…………..………………………………….….…...…136 Appendices A: RAW HIPPO CANINE ISOTOPE DATA AND FIGURES………………...…..….147 B: RAW ISOTOPE DATA FROM QUEEN ELIZABETH NP HIPPOS…….…..…….227 C: RAW ISOTOPE VALUES OF ARCHAEOLOGICAL TOOTH ENAMEL AND LEAF WAX BIOMARKERS…...…………………………….………….…....232 vi LIST OF FIGURES Figures 1.1. Axial precession and location of the perihelion throughout the Holocene……..…..15 2.1. Map of localities where hippos were collected.……..………………………...……43 2.2. Boxplots of δ13C values from hippo canine enamel profiles. Boxplot colors correspond to environmental groupings given in Figure 2.3..…………...45 2.3. Boxplots of enamel isotope values grouped by environment. Boxplot colors correspond to statistically significant ecological groupings (P-values presented in Table 2.3)..…………………………………..………......….46 2.4. Comparison of hippo profile isotope values (blue) vs. other mammal enamel isotope values (data from Cerling 2014)…………………………..………......…52 2.5. Comparison of two hippos from Lake Naivasha – Crayfish Camp vs. Crater Lake.53 2.6. Isotope profiles of three Tsavo hippo canines showing differential responses to drought in the park based on locality.………………….………..…....…54 2.7. High resolution δ13C and δ18O enamel data from KF-11……………………….….55 3.1. Map of Queen Elizabeth Park, Uganda.………………………..…………….….....76 3.2. Carbon isotope profiles of the three Queen Elizabeth hippo serial samples (1970 tusk in black, 1991 tusk in dark grey, 2000 tusk in light grey) and number of hippos (dotted line) and elephants (solid line) present in the park over time (Olivier 1991; Plumptre et al. 2010a)………………………………..…..78 4.1. Map of the study area.………..…………………………………………...….....…112 4.2. Holocene paleoenvironment and archeology in Kenya………..…………….….....113 4.3. Boxplots of ẟ13C values and dietary designations for herbivores in the Lake Victoria Basin………………………………………………..……………..……118 4.4. Boxplots of ẟ13C values and dietary designations for herbivores in the Lake Turkana Basin……………………………………………………………..…..…121 4.5. Leaf wax biomarker isotope values for terrestrial lacustrine deposits in Area 103, Turkana (dates recalibrated using OxCal 4.2 using IntCal13 from Owen and Renaut, 1986)………….………………………………..……….....122 4.6. Boxplots of ẟ13C values and dietary designations for herbivores in the Ele Bor …124 4.7. Holocene paleoenvironment at Lake Victoria, Kenya……………………………..126 4.8. Holocene paleoenvironment at LakeTurkana and Ele Bor, Kenya..........………….128 5.1. Location of Gogo Falls in relation to the other Holocene archaeological sites (●) and towns (○) in the Lake Victoria Basin……………………………...……131 5.2. (A) Holocene pottery traditions present at the archaeological sites in the Victoria Basin. (B) Leaf wax δ13C and modeled %C4 from lacustrine sedimentary cores in Lake Victoria (35). (C) Moraceae (tropical mesic trees and shrubs) pollen counts from lacustrine sedimentary cores in Lake Victoria (data from ref. 34). (D) Poaceae (grass) pollen counts from lacustrine sedimentary cores in Lake Victoria (data from ref. 34)…………...…....131 5.3. Box plots of δ13C tooth enamel values from archaeological tooth enamel, Trench III, Gogo Falls…………………………………………..…………………..132 5.4. Box plots of δ13C1750 tooth enamel values from modern comparative fauna from Kenya and Tanzania……………………………………...……………..132 5.S1.Boxplots of δ13C tooth enamel values of modern goat, Capra hircus (Table 5.S4), modern sheep, Ovis aries (Table 5.S4), and Gogo Falls (Table 5.S3)…144 A.1. δ13C and δ18O canine enamel profile plot of Aberdares NP hippo………..…..….148 A.2. δ13C and δ18O canine enamel profile plot of Amboseli #1 hippo (0mm = death)..149 A.3. δ13C and δ18O canine enamel profile plot of Amboseli #2 hippo………...…....…150 A.4. δ13C and δ18O canine enamel profile plot of Arabuko Sokoke #1 hippo.……...…151 A.5. δ13C and δ18O canine enamel profile plot of Arabuko Sokoke #2 hippo.……...…152 A.6. δ13C and δ18O canine enamel profile plot of Arabuko Sokoke #3 hippo………....153 A.7. δ13C and δ18O canine enamel profile plot of Arabuko Sokoke #4 hippo……..…..154 A.8. δ13C and δ18O canine enamel profile plot of Buffalo Springs hippo…………..…155 A.9. δ13C and δ18O canine enamel profile plot of Chyulu hippo……………..………..156 viii A.10. δ13C and δ18O canine enamel profile plot of Kisumu hippo ……………….....…157 A.11. δ13C and δ18O canine enamel profile plot of Laikipia hippo………………….…158 A.12. δ13C and δ18O canine enamel profile plot of Mara hippo……………..…………159 A.13. δ13C and δ18O canine enamel profile plot of Meru NP hippo…………..……..…160 A.14. δ13C and δ18O canine enamel profile plot of Minjila hippo…………..………….161 A.15. δ13C and δ18O canine enamel profile plot of Mokowe hippo………..…………...162 A.16. δ13C and δ18O canine enamel profile plot of Mpeketoni hippo………..…………163 A.17. δ13C and δ18O canine enamel profile plot of Mwea – Gitaru Dam hippo……..…164 A.18. δ13C and δ18O canine enamel profile plot of Mwea #1 hippo………………...….165 A.19. δ13C and δ18O canine enamel profile plot of Mwea #2 hippo…………..………..166 A.20. δ13C and δ18O canine enamel profile plot of Mwea #3 hippo…………..…….….167 A.21. δ13C and δ18O canine enamel profile plot of Naivasha hippo………………..…..168 A.22. δ13C and δ18O canine enamel profile plot of Naivasha hippo………..…………..169 A.23. δ13C and δ18O canine enamel profile plot of Lake Nakuru hippo……………..…170 A.24. δ13C and δ18O canine enamel profile plot of Lake Ol Bolossat hippo……..…….171 A.25. δ13C and δ18O canine enamel profile plot of Turkana – Koobi Fora 2 hippo..…..172 A.26. δ13C and δ18O canine enamel profile plot of Witu hippo………..……………….173 ix LIST OF TABLES Tables 2.1. UNESCO vegetation classification scheme (from White, 1983)……………....…...44 2.2. Hippo tusks sampled by locality………………..………………………..…………47 2.3. P-values of relationships between environmental groupings of δ13C enamel values from pairwise comparisons using a Tukey-Kramer (Nemenyi) test…..…50 3.1. Mean and standard deviation δ13C values for the three hippo canines. P-values from Kruskal-Wallis with Tukey and Kramer (Nemenyi) post-hoc test (executed in R) verify that the three hippos have distinctive feeding niches.………77 4.1. δ13C of herbivore tooth enamel from the Lake Victoria Basin.……………...…...116 4.2. δ13C of herbivore tooth enamel from the Lake Turkana Basin.…………………..120 4.3 δ13C of herbivore tooth enamel from Ele Bor...……….…....…..……………....…124 5.S1. δ13C tooth enamel values (average and standard deviation) from archaeological tooth enamel, Trench III, Gogo Falls..……….……..……..………....…136 5.S2. δ13C1750 tooth enamel values (average and standard deviation) from modern comparative fauna from Kenya and Tanzania……………………….....137 5.S3. δ13C and δ18O tooth enamel values from Gogo Falls……….………………..…...138 5.S4. Modern caprine δ13C1750 and δ18O tooth enamel...……………………..………...141 5.S5. Comparison of wild vs. domestic fauna at Neolithic sites in Kenya……...……...143 A.1. Aberdares NP hippo isotope values.....................................………...………....…174 A.2. Adhi Dam (Boni NR) hippo isotope values...................................…...………..…175 A.3. Amboseli #1 hippo isotope values...................................…...……....………....…177 A.4. Amboseli #2 hippo isotope values..…...…………...……………………..…....…179 A.5. Arabuko Sokoke #1 hippo isotope values..….……….……………………......…181 A.6. Arabuko Sokoke #2 hippo isotope values..….……….……………………......…182 A.7. Arabuko Sokoke #3 hippo isotope values…….……….………..…..………....…183 A.8. Buffalo Springs hippo isotope values..….……….…………....……………....…184 A.9. Chyulu hippo isotope values..……....………………..……………………......…186 A.10. Kisumu (Lake Victoria) hippo isotope values…...….…………….………......…187 A.11. Laikipia hippo isotope values….……….………………………....………......…188 A.12. Maasai Mara hippo isotope values…….……….…………………….….…….…189 A.13. Meru NP hippo isotope values..….……….…………………………..….…....…190 A.14. Minjila hippo isotope values..….……….………………..…………….…...……191 A.15. Mokowe hippo isotope values..……..………………………………….….......…193 A.16. Mpeketoni hippo isotope values..…………………………………..….…...….…195 A.17. Mwea – Gitaru Dam hippo isotope values………….…................................……197 A.18. Mwea #1 isotope values..………….…..........................................................……198 A.19. Mwea #2 hippo isotope values.......................................................................……200 A.20. Mwea #3 hippo isotope values.......................................................................……202 A.21. Naivasha – Crater Lake hippo isotope values.............................…………..….…204 A.22. Naivasha – Crayfish hippo isotope values............................………………….…206 A.23. Lake Nakuru hippo isotope values………............................…………..……...…208 A.24. Lake Ol Bolossat hippo isotope values.………............................…………….…209 A.25. Tsavo #1 hippo isotope values..………............................…………………….…210 A.26. Tsavo #2 hippo isotope values...………............................…………..………..…212 A.27. Tsavo #3 hippo isotope values..……….............................…………..………..…214 xi A.28. Turkana – Koobi Fora 1 hippo isotope values.………..….……...………………216 A.29. Turkana – Koobi Fora 2 hippo isotope values..………..………...………..…..…221 A.30. Witu hippo isotope values...………..………...……………………..………....…223 A.31. Bonferroni corrected P-values, Wilcoxan rank-sum test………………………...225 B.1. Raw isotope values of QEP hippos through time.………………….....………….228 C.1. Isotope values of Luanda (9.7 – 8.5 Kya) fauna.………………….…….……….233 C.2. Isotope values of Wadh Lang’o (~3 Kya) fauna.……………...……...…...…….235 C.3. Isotope values of Wadh Lang’o (~2 Kya) fauna.………..………......….....…….236 C.4. Lothagam faunal isotopes (8.3 Kya)……………………..……….……...…...….237 C.5. Later Holocene faunal isotopes (8.3 Kya)………………………………..…...….238 C.6. EBA, Horizon C faunal isotope values…………………………………….…….240 C.7. EBA, Horizon B faunal isotopes……………………………………………..…..241 C.8. EBA Horizons A1 and A2 isotope values…………………………..…………....242 C.9. C28 FAME leaf wax biomarker values……………………………………….…..243 xii ACKNOWLEDGEMENTS This dissertation is dedicated to the many people whose support, constructive criticism, and care contributed to its ultimate success. Foremost, I wish to thank my advisor, Thure Cerling, for his guidance and mentorship throughout my Ph.D. He took me on as a student in paleoecology and started me off running in my first year, pushing me to manage the lab and take advanced geology courses (even though I came from a Biology degree and couldn’t map a section to save my life). Thure always challenged me to do things in a thoughtful, conscientious manner, and he encouraged me to ask difficult questions and think about my research in lateral ways. He always reminded me to approach complicated problems using the adage “do all you know and try all you don’t”. He encouraged me to hold many jobs at once - graduate student, lab manager, technician, my own field project director – and though occupying these roles simultaneously seemed overwhelming at the time, these responsibilities have prepared me for a career in academia better than I could have imagined. I wish to thank the members of my committee, who have each provided immense support and guidance throughout my Ph.D. Firstly, I wish to thank Kate Freeman, who generously opened her lab to me to learn an entirely new field of research – organic geochemistry – and let me struggle through 1.5 semesters of challenging geochemical analysis in her world-class laboratory. Jim Ehleringer always challenged me to think critically and look at my data closely, and through constructive conversations together, always gave me new perspectives on data analysis and experimental design. I owe my knowledge of field geology, geomorphology of the Turkana Basin, and other invaluable topics, such as East African flora and fauna, to Frank Brown. He provided logistical support and timely advice for my early fieldwork in Turkana. I always left conversations with him learning new and invaluable things, whether they were passable roads to take in West Turkana or local lore. Denise Dearing was an incredible source of career and logistical advice on how to navigate graduate school and beyond, and was an amazing mentor on teaching and advising. I also had other wonderful mentors who gave so much of their time, advice, and constructive criticism to my research. Elisabeth Hildebrand spent many evening and weekend hours with me on Skype, giving me a crash course in African archaeology and pastoralism while tirelessly assisting me with the language and writing of my Doctoral Dissertation Improvement Grants. Fiona Marshall has been a wonderful source of support and mentorship during the final year of my Ph.D. It is with a heavy heart that I acknowledge my colleague, Samuel Andanje, who passed away earlier this year. We collaborated and planned many exciting future projects on wildlife isotope ecology, and his wealth of knowledge on African wildlife and the maze of navigating research within the Kenya Wildlife Service got me through my first few years of fieldwork. I also wish to acknowledge the mentorship and guidance of other researchers while in East Africa: F. Kyalo Manthi, Purity Kiura, Emma Mbua, Stan Ambrose, Louise Leakey, and Jessica Rothman. These people assisted me with the permit process, provided local connections and collaborators, loaned me vehicles, gave me rides in their cars and airplanes, and provided advice on tropical maladies and research life. xiv Furthermore, I wish to thank the cohort of graduate students who were my constant companions for research and friendship in Nairobi, Turkana, and Uganda: Kirsten Jenkins, Phil Slater, Anneke Janzen, Elizabeth Sawchuk, Sarah Pilliard, Margret Bryer, Caley Johnson, Nick Blegen, Hilary Duke, and Steven Goldstein. Here’s to many more years of fieldwork and life in East Africa together. There are many other graduate students and friends who were sources of support and friendship in my life during my Ph.D., including (but not limited to): Heather Graham, Sarah Knutie, Kevin Kohl, and Shannon Gaukler. I would not have made it through this project without them. Furthermore, I wish to express my gratitude and love to those who gave me support and care, and forced me to rest, even when I protested: Brittany Dame, who has been my good friend and climbing partner throughout graduate school, and Lila Leatherman, whose companionship has been a font of strength, happiness, and tranquility in my life. I wish to also thank my family for their patience throughout graduate school: my mother Kathy, my father Jeff, my brother Jeremy, and my sister-in-law Steffanie. I will never forget their patience and understanding for the missed family gatherings and times I had to work during holidays. They have always supported me throughout this entire process and have been incredible wells of strength during long and exhausting field seasons and months of tedious lab work. Finally, I wish to thank my partner, my friend, and my closest collaborator, Scott Blumenthal. Throughout the entirety of my degree, his support, caring, and critical eye has made me both a better scientist and a better person. He closely read drafts of manuscripts, assisted me with complicated lab maintenance, and cared for me when I was helpless and ill from malaria in the field. Thank you, all, for believing in me. xv INTRODUCTION 1.1 Background Stable isotope records of ecology from biological and geological materials have provided otherwise inaccessible knowledge of the ways in which mammals, ecosystems, and ancient people respond to perturbations in climate and ecology (Cerling et al. 2004; Cerling et al. 2005; Magill et al. 2013a; Magill et al. 2013b). Some isotopic records, such as those from certain biological tissues (e.g., hair and ever-growing teeth), can provide near-continuous records of how animals responded to disturbances in the natural world. In a paleoecological context, isotope records from fossil and archaeological mammalian tooth enamel have provided snapshots of their ecology, relating to changes in vegetation (Ambrose and Sikes 1991; Passey et al. 2002; Uno et al. 2011). These data, in concert with other isotope records from geological samples, particularly leaf wax biomarkers, can provide further context for paleoenvironment that is unbiased by mammalian behavioral characteristics, such as selective diets and migration. These records are of special interest to researchers working in eastern Africa, where direct application of stable isotope techniques can address wide-ranging questions in climate-environment-human interactions over millions of years. Such questions involve the particular landscapes in which humans evolved, and the sorts of challenges and opportunities those landscapes may have presented to them. In a more recent context (<10,000 years), paleoecological records from stable isotopes can help clarify the 2 environments that shaped modern human behavior, such as early food production. These two issues, though working on different temporal scales, go hand-in-hand, as recent sedimentological records can provide a model for understanding deep time climateenvironment interactions in Eastern Africa. Isotope records from biological material can also be used to address complicated questions of environmental change in historical ecology. Such data have assisted ecologists and paleoecologists in assessing dietary, behavioral, and environmental changes (Koch et al. 1995; Hirons et al. 2001; Ayliffe et al. 2004; Dalerum and Angerbjörn 2005; West et al. 2006; Zazzo et al. 2010). These data not only have utility in modern environments, but also in ancient environments, where they can provide a framework for how to interpret the past. Isotopic data from Hippopotamus amphibius canines can provide decadal-scale records due to their length and their recently understood ecological flexibility (Boisserie et al. 2005; Passey et al. 2005; Cerling et al. 2008). Questions still remain, however, as to exactly how variable these records are and what hippo dietary isotopes tell us about their immediate environment. Over longer timescales, many researchers relate environmental changes in East Africa to climate changes. The climate of eastern African is driven primarily by Milankovitch cycles, periodic changes in the geometry of Earth's orbit that dictate the amount of solar insolation on Earth (Claussen et al. 1999; deMenocal et al. 2000; Stager et al. 2003; Laskar et al. 2004). Currently, precession of the equinoxes (Figure 1.1) is the driving orbital parameter that influences tropical climate (Kutzbach 1981; Berger and Loutre 1997; Clement et al. 2001; deMenocal 2004; Battisti and Naylor 2009; Clark et al. 2009). Insolation dictates the location and convective strength of the intertropical 3 convergence zone (ITCZ), an area where Northern and Southern Hemisphere winds meet. The ITCZ moves seasonally, following peak solar insolation about equatorial Earth. Seasonal rainfall occurs in the ITCZ due to warming sea surface temperatures (SSTs) and differential heating between land and sea, which pulls moisture-rich air over the continents. When Northern Hemisphere peak solar insolation occurs as the earth is closest to the sun (the perihelion, Figure 1.1), the ITCZ strengthens, forming a lowpressure zone over northern Africa that drags moist air over the continent and intensifies the monsoon system (Nicholson 1996). During the early Holocene, precessionally driven high Northern Hemisphere insolation created mesic conditions known as the African Humid Period (AHP) (Kutzbach 1981; Braconnot et al. 2000; Gasse 2000; Russell et al. 2003; Mayewski et al. 2004; Garcin et al. 2007; Verschuren et al. 2009; Tierney et al. 2011b). Lake level water budget models suggest that mean annual rainfall in the watersheds of Lakes Turkana, Victoria, Naivasha, and Nakuru-Elmenteita increased to 15–35% greater than 1980s levels during the AHP, filling the lakes to their highest extents and, in some cases, reaching overflow points (Hastenrath and Kutzbach 1983). As northern hemisphere insolation strength waned c. 5.5–5.0 Kya, a conspicuous arid period began in eastern Africa (Kutzbach 1981; deMenocal et al. 2000; Gasse 2000). In Lake Turkana, decreasing rainfall is demonstrated by falling lake levels observable via paleolake terraces, whereas in Lake Victoria, this trend is reflected in hydrogen isotopes from leaf waxes, albeit more gradually than in Turkana (Berke et al. 2012; Garcin et al. 2012). Inferences of paleo- and/or historical ecology based on climatological data rely on the concept that rainfall is a major driver of the proportion of C3 (woody plants, trees, and 4 shrubs) vs. C4 (tropical lowland grasses) biomass in African ecosystems, and particularly in savanna ecosystems. Woody cover can increase or decrease in response to climatological factors, such as changes in overall precipitation or seasonal distribution of rainfall (Marshall et al. 2007; Tierney et al. 2011a). However, the assumption that rainfall is the only driver of woody cover in savannas is problematic for many reasons, the first of which is that the term “savanna” is a broad ecological designation. A savanna is defined as a “mixed tree-grass system characterized by a discontinuous tree canopy in a conspicuous grass layer” (Ratnam et al. 2011). Savannas have 5–80% fractional woody cover; they include the structural categories grassland, wooded grassland, and woodland/ bushland/shrubland (Sankaran et al. 2005; Cerling et al. 2011; Good and Caylor 2011; Lehmann et al. 2011; Murphy 2012). However, many other ecological factors also influence the relative proportions of grass and woody cover (C4 vs. C3 plants): soil nutrients, grass and shrub competition with trees, underground biomass, fire frequency, and herbivory. Given all this, inferring ecological changes solely based on rainfall changes is tenuous (Sankaran et al. 2005). Stable isotope analysis of mammalian tooth enamel is a useful indicator of modern and ancient environments in that dietary isotopes can differentiate between C3 browsers and C4 grazers (Lee-Thorp and Van der Merwe 1987; Ambrose and DeNiro 1989; Cerling et al. 1997). Tooth enamel resists diagenetic alteration and preserves well in the archaeological record, and is therefore preferred for paleoecological reconstructions (Lee-Thorp et al. 1989; Kohn and Cerling 2002; Uno et al. 2011). Isotopic measurements of enamel carbon (δ13C) and oxygen (δ18O) can be used to understand diet, habitat, and climate (Levin et al. 2006; Lee-Thorp et al. 2007). In many 5 instances, fauna provide the best estimates of environmental change over time, because many mammals adapt to changes in vegetation structure, and their isotopic signal is less temporally attenuated than other proxies (Kingston 2007). Modern and ancient sediments also provide important information about past environments. Bulk isotopic measurements of soil organic matter can indicate overall terrestrial ecosystem structure via representation of C3 vs. C4 plants, which correspond roughly to woody plants versus tropical lowland grasses in Africa (Cerling et al. 1989; Ambrose and Sikes 1991; Cerling et al. 2011). The isotopic signature of bulk plant material recovered at molecular level in the form of leaf wax biomarkers can convey more specific information about floral composition of paleoenvironments than bulk organic material or tooth enamel (Eglinton and Hamilton 1967; Brincat et al. 2000; Freeman and Colarusso 2009). Terrestrial leaf wax lipids preserved in sediments (medium-high molecular weight n-alkanes and n-alkanoic acids) derive from aquatic organisms and plant leaf waxes, respectively. Thus, compound specific isotopic analyses (δ13C and δD, deuterium) can convey highly specific information about paleoclimate, hydrology, and ecosystem structure (Xu and Jaffé 2008; Choudhary et al. 2009; Tierney et al. 2011b; Garcin et al. 2012). Thus, terrestrial biomarkers in lacustrine sediments represent a spatially integrated signal on a basinal scale (Talbot and Livingstone 1989). This dissertation will provide new isotopic records defining environmental variability in both modern and ancient ecosystems in Kenya and Uganda. Understanding the drivers for East African environmental change requires empirical data connecting ecology with known climatic and biological perturbations. In recent environments, changes in savanna structure in national parks have been attributed to changes in 6 herbivore biomass (especially elephants) and fire frequency (Dublin et al. 1990; Dublin 1995; Roques et al. 2001; Bradshaw et al. 2003; Western 2007; Valeix et al. 2011; Asner and Levick 2012). In Holocene paleoecological research, with the exception of a few studies (Ambrose and DeNiro 1989; Ambrose and Sikes 1991; Balasse and Ambrose 2005), most reconstructions of terrestrial paleoenvironments in Kenya in relation to archaeology have employed inferences from indirect paleoclimate records (Bower 1991; Gifford-Gonzalez 1998; Wright 2007; Prendergast and Lane 2010; Ashley et al. 2011). 1.2 Chapter Summaries 1.2.1. Chapter 2: Isotopic composition of wild H. amphibius canines reveals nonselective feeding behavior in Kenya Chapter 2 explores the relationship between individual hippo dietary histories and their local environment. Isotopic work in the last 20 years has revealed that hippos are not strict selective C4 grazers, as has been the long-standing assumption by ecologists. Hippos feed close to their aquatic environment, and thus likely reflect the proportion of C3/C4 herbaceous vegetation near their aquatic habitats, which can vary over the course of their lifetime due to changes in environmental conditions. To test this idea, we serially sampled (10 mm intervals) enamel from ever-growing hippo canines from wild hippos living in diverse environments across Kenya for δ13C and δ18O. Enamel δ13C values from these isotope profiles group significantly based on environmental classifications of the areas they come from, ranging from woodlands and forests to open grasslands. Similarly, radiocarbon dated profiles from hippos living in the same environment over the same time period and/or in the same geographic area reveal the variable nature of hippo diets among populations. These data show that enamel δ13C values from hippos can be 7 excellent recorders of local environment, though caution must be used when extrapolating hippo dietary isotope values to determine paleoecology of large areas. 1.2.2. Chapter 3: Decadal diet change in Hippopotamus amphibius in Queen Elizabeth National Park, Uganda. Chapter 3 builds upon the data presented in Chapter 2 and applies the same techniques to capture a signal of C3 succession in a savanna ecosystem in Uganda following a major ecological perturbation – the poaching and near extirpation of elephants from Queen Elizabeth Park (QEP), Uganda. Hippo canines from hippos living on the Mweya Peninsula in QEP that have either been radiocarbon dated or have known death dates were serially sampled for δ13C and δ18O. These hippos exhibit a >80% C4 diet throughout the 1960s, in agreement with vegetation transects from this time period that demonstrate an abundance of C4 grasses on the peninsula. Following heavy poaching in QEP throughout the 70s and early 80s, informal vegetation assessments indicated succession of C3 woody plants and forbs. Hippo dietary isotope values capture this increase in C3 on the peninsula during the 80s and 90s, and reveal the extent to which C3 herbaceous plants have encroached, decreasing grazing capacity on the peninsula. These results suggest that hippo canines can be excellent indicators of herbaceous vegetation change in savanna ecosystems and could be used in African parks to track C3 encroachment, an ongoing issue in modern African ecology. 1.2.3. Chapter 4: Holocene paleoenvironments in Kenya, with perspectives on Holocene archaeology and Pleistocene environmental change Changes in the strength of the East African monsoon due to orbital forcing have often been cited as drivers of terrestrial ecosystem change throughout the last 4 Ma. The 8 nature of how terrestrial ecosystems respond to orbital forcing has only been empirically demonstrated in a few studies, and rarely with an interbasinal perspective. It is expected that regional changes in monsoon strength would be reflected in locally distinctive ways. To test this hypothesis, paleoecological records using faunal tooth enamel and sedimentary leaf wax biomarkers were generated and compiled for Holocene records, a time of well-constrained paleoclimatolgical change from orbital forcing, for two distinct lake basins in Kenya – the northern Turkana Basin and the southwestern Lake Victoria basin. Where possible, pollen was also included. The data indicate that abrupt aridification in Turkana did indeed coincide with dietary change in mammal herbivores, whereas the gradual ardification in Lake Victoria did not. Likewise, an inland archaeological site, Ele Bor (250 km from Lake Turkana) reveals no dietary change in herbivores. Furthermore, leaf wax and tooth enamel data tend to agree in Victoria, though not in Turkana (where samples overlap). These data show that orbital forcing has differential effects in climate and vegetation change, and paleoecology must be assessed basin by basin. 1.2.4. Chapter 5: Environments and trypanosomiasis risks for early herders in the later Holocene of the Lake Victoria Basin, Kenya Herding was the earliest form of African food production, and transformed local populations of people and animals. Herders migrated from eastern to southern Africa around 2,000 years ago but only in small numbers. Zoonotic disease vectors, specifically the tsetse fly, which carries sleeping sickness, are thought to have impeded these movements. Archaeologists have argued that the presence of tsetse flies around Lake Victoria, Kenya, created a barrier preventing migration and forcing subsistence 9 diversification. 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Geochimica et Cosmochimica Acta 74:3571–3586. 15 Axial precession: ~23 Kyr -0.08 0.00 0.08 0 Kyr 100 200 shift of equinox relative to perihelion Today - perihelion during norther winter 5,500 YBP - perihelion during northern spring 11,000 YBP - perihelion during northern summer (Earth pictured in all three scenarios are in its relative position on December 21) Figure 1.1: Axial precession and location of the perihelion throughout the Holocene. Modified from Zachos et al. (2001). CHAPTER 2 ISOTOPIC COMPOSITION OF WILD H. AMPHIBIUS CANINES REVEALS NONSELECTIVE FEEDING BEHAVIOR IN KENYA 2.1. Abstract The common hippopotamus (Hippopotamus amphibius) is a semi-aquatic megaherbivore that plays a crucial role as an ecosystem engineer in the environment that it inhabits. Dietary isotopes in hippo canines have long been of interest to the paleoecological community, as their canines are ever-growing and can potentially record up to 10 continuous years of their ecology. Bulk stable isotope data from hippo cheek teeth has revealed that hippos are not strict C4 grazers, as previously thought. This necessitated further investigation into the ecological and dietary variability of hippos from different environments. To that end, hippo canine enamel was sampled serially (~10 mm) for δ13C along the length of the canine from 30 wild hippos. These isotope profiles have revealed the remarkable variability in hippo diets that relate, broadly, to the structure of the ecosystems they inhabit and major environmental changes, such as drought. Though C4 grasses and grassland exist in each of these localities, hippo dietary isotopes indicate that their C4 intake can be as low as 40% in more C3 environments, making them excellent indicators of C3 herbaceous groundcover. These dietary records only record environment on a very local scale, given that they forage less than 6 km away from their aquatic habitat. 17 2.2. Introduction Hippopotamus amphibius is a common megaherbivore (>1000 kg) found across Africa, living in a wide range of environments from arid savannas to forest (Eltringham 1999; Klingel 2008; Klingel 2013). They are semi-aquatic, spending the day in pools, rivers, and lakes, coming out of water at night to feed on herbaceous vegetation. Their feeding behavior is difficult to observe, as they feed nocturnally and tend to crop foods close to the ground, such that they create lawns of very short vegetation, deterring herbivores that prefer tall forage, such as elephants and buffalo (Field 1970; Arsenault and Owen Smith 2002; 2006; Kanga et al. 2011). They often stay close to their aquatic habitats to feed (1–6 km away) based on habitat structure and access to nutritious food (Eltringham 1974; Eltringham 1999; Lewison and Carter 2004). When necessary, they can wander quite far (up to 20 km) during rainy seasons due to lower trans-cutaneous water loss when moving on land (Vesey-FitzGerald 1960). Hippos occupy unique roles in their ecosystems and serve as ecosystem engineers in the habitats where they live. Their manner of walking along the same paths towards food sources causes narrow water channels to form, contributing to swamp development and irrigation (Owen-Smith 1992; Chansa et al. 2011). Hippos require as much as ~20 kgs of food per day; these high dietary requirements and repeated journeys on land can lead to trampled and compacted soil, killing vegetation and transforming soil properties, especially when populations are high (Lock 1972). An intermediate number of hippos, however, can increase herbivore diversity and environmental mosaics (Bourlière and Hadley 1970; Lock 1972; Owen-Smith 1992; Fritz et al. 2002; Verweij et al. 2006; Kanga et al. 2011). Hippos also facilitate nutrient cycling between lakes and terrestrial 18 environments (McCauley et al. 2015; Subalusky et al. 2015). Despite the important roles hippos occupy in their environment, basic aspects of their ecology are still uncertain. Understanding hippo diets is of particular interest, as they have been and are still classified as strict selective grazers by ecologists (Field 1970; Olivier and Laurie 1974; Kanga et al. 2011; McCauley et al. 2015), though recent isotopic samples on modern and fossil hippos has revealed the broad nature of their feeding behavior (Boisserie et al. 2005; Cerling et al. 2008; Harris et al. 2008). These data have raised intriguing questions about the extent of hippo dietary flexiblity, both among environments and within individuals themselves. Understanding how dietary isotopes of hippos relate to their environment will help us better interpret isotopic data of hippos from the geologic past, since their canines are ever-growing and can record up to a decade of their recent life (Passey et al. 2005; Uno et al. 2013). This study aims to understand ecological and dietary variability within individuals across different environments through sequential isotope sampling of modern wild hippo canine enamel in order to quantify the degree of dietary flexibility within the species. 2.3 Materials and Methods 2.3.1. Sample collection Hippo canines were collected from Kenya Wildlife Service (KWS) stockpiles; some of these had been shot by rangers for either herd management or had been identified as nuisance animals (i.e., crop raiding)(Figure 2.1). Tusk enamel powder was drilled at intervals, usually every 10 mm, using a diamond-tipped Dremel tool; these were sampled by Dr. Samuel Andanje (KWS) and trained students from Moi University, Kenya. To remove surface contaminants, the surface of the enamel was abraded off prior 19 to drilling. Because samples were obtained from recently modern hippos, enamel powders were not treated because various treatment procedures are not distinguishable for modern tooth enamel (Passey et al., 2002; Cerling 2014). In total, 1,410 samples were analyzed from 30 tusk profiles from national parks and protected areas across Kenya. Sample powders were weighed into silver capsules and stored in a drying cabinet until analysis. Samples were analyzed for δ13C and δ18O on a Finnigan MAT 252 coupled to a Carboflo dual-inlet carbonate device, which involves digestion in a common phosphoric acid bath at 90°C for 10 minutes. Stable isotope ratios are reported as δ values relative to the international carbon isotope standard, Vienna Pee Dee Belemnite (VPDB), using the standard permil (‰) notation, where δ13C = (Rsample/Rstandard -1) x 1000, and Rsample and Rstandard are the 13C/12C ratios of the sample and standard, respectively. Standard deviation of an internal carbonate standard (Carrara marble) was between +/- 0.1 and 0.2‰. Samples were corrected to 25ºC using modern tooth enamel samples analyzed at both 25º and 90ºC (Passey et al. 2007). 2.3.2. Environmental characterization Characterization of eastern African environments was done using the Potential Natural Vegetation map for Eastern Africa (also called the “Vegetation and Climate Change in East Africa” map; VECEA) (Lillesø et al. 2011). This map in an updated, interactive version of the classic vegetation map for East Africa developed by White (White 1983). This map is a higher spatial resolution than the White map, and both ecoregions and physiognomic vegetation groups include environmental data, such as rainfall, altitude, and temperature. The VECEA also includes a Google Earth overlay, which was used to define vegetation characteristics for hippo localities. The VECEA map 20 defines floristic vegetation types based on the degree of woody cover in an area, dominant woody species, and edaphic characteristics that may influence floral distribution within the vegetation type. Environmental characterizations used in this and subsequent chapters follow the UNESCO classification scheme, developed and defined by White (1983) (Table 2.1). Unfortunately, hippo localities are coarsely defined – typically only to the national park or reserve in which the animal was shot or the carcass was found. Thus, defining habitat localities for many hippos is challenging, and generalized to the most prevalent ecosystem types, which in some cases are compound. Further classification for forest types, accounting for climatic and edaphic characteristics, are explained in the detailed descriptions given here. The code “Bd” refers to the physiognomic vegetation type as “Somalia-Masai Acacia-Commiphora deciduous bushland and thicket”. This vegetation type is most commonly characterized as a woodland, with thickets that can often be impenetrable (Kindt et al. 2011c). Rainfall within this vegetation type ranges between 250 and 500mm/year (White 1983), and grasses are more abundant in places where the soil is sandy and otherwise well-drained. The most typical example of this vegetation type is found within Tsavo National Park (White 1983). The code “Wcd” refers to the physiognomic vegetation type “dry Combretum wooded grassland”. This vegetation type is a variant of a more major, widespread East African ecosystem type “Combretum wooded grassland” (White 1983; Kindt et al. 2011c). Wcd areas occur in places that receive between 600 and 1800 mm of rainfall per year. As the name implies, most of the herbaceous layer is composed of grasses and 21 herbs/forbs, with few thickets or bush present (Kindt et al. 2011b). The code “wd” refers to the physiognomic vegetation type “Edaphic wooded grassland on drainage-impeded or seasonally flooded soils”. This vegetation type is similar to riverine wooded grasslands, though wd typically contains primarily Acacia species (White 1983). These environments typically are comprised of tall grasses that are frequently subject to burning and conspicuous stands of trees (White 1983; Kindt et al. 2011b). The code “Fp” refers to the Zanzibar-Inhambane undifferentiated forest. This coastal forest variation is highly heterogeneous, with drier forest types covering a larger area on the coast (White 1983). This is the habitat type within the Boni National Reserve, from which the Adhi Dam hippo specimen came. This coastal forest is floristically diverse, though with a more closed canopy than Zanzibar-Inhambane woodland and scrub forest (Burgess et al. 1998; Tabor et al. 2010). The code “Fq” refers to the physiognomic vegetation type as “ZanzibarInhambane scrub forest”. This coastal forest delineates the coastal region forests and inland bushlands and occupies areas where rainfall is between 500 and 750 mm/yr (White 1983). This is a highly diverse forest type without dominance of a single tree species, and occurs at low elevations, below 250m above sea level. This vegetation type has abundant woody vegetation with a high degree of floral and faunal endemism, especially within Arabuko-Sokoke National Park (Burgess and Muir 1994). The coastal floristic region within which this vegetation type occurs can be generalized as an undifferentiated forest, meaning changes in the structure and composition of vegetation types occur over short distances (White 1983). 22 The code “g” refers to the physiognomic vegetation type “edaphic grassland on drainage-impeded or seasonally flooded soils”. This vegetation type is distinct from type wd in that it is a true grassland, with <10% woody cover (White 1983; Kindt et al. 2011a). These grasslands are associated with seasonally or permanently waterlogged soils at altitudes below 1,500m and a large range in rainfall (between 200 and 1,400mm/yr) (Kindt et al. 2011a). The code “R” refers to the physiognomic vegetation type “riverine wooded vegetation”. This is a compound vegetation designation encompassing several different physiognomic variants, including forest, woodland, and thicket. Unfortunately, finer resolution descriptions for hippos from these localities (Naivasha and Nakuru) were not available. This vegetation type is widely distributed across East Africa, and given that climate is not the primary driver of water availability, all variants of this vegetation type are edaphic. This code is a mosaic vegetation classification comprised of Acacia-Commiphora deciduous wooded grassland (Wd) and Combretum wooded grassland (Wc). Combretum wooded grassland is a major vegetation type in East Africa (White 1983), though more abundant outside of Kenya (Kindt et al. 2011b). Wd environments are typical of the greater Serengeti ecosystem, and what distinguishes them from deciduous bushland and thicket (Bd) is the relative lower abundance of bushy plants and higher representation of perennial grasses, which may be the influence of biotic factors, such as herbivory, rather than climate (White 1983). This compound vegetation type is prevalent in Amboseli National Park. The code We/Wd is a mosaic vegetation classification comprised of edaphic 23 wooded grassland on drainage-impeded or seasonally flooded soils (wd) and biotic Acacia wooded grassland (We). We is a floristic type that is closely related to evergreen bushland (Be), though We environments typically experience higher grazing pressure, and thus support fewer evergreen thickets (White 1983; Kindt et al. 2011b). We environments can have discontinuous groundcover in more arid areas and occur largely above 1,250 m above sea level (White 1983). This compound vegetation type is commonly found in the Mara and Laikipia district. The code “S” is a mosaic vegetation classification comprised of Somalia-Masai semi-desert grassland and shrubland (S). This distinctive vegetation type occurs in very arid regions and on sandy soils, particularly those with 100–250 mm/yr rainfall. In certain areas where this vegetation type exists, woody plants provide 2–20% of the ground cover, as either shrubs or bushy trees (White 1983). This vegetation type is present in the arid Turkana region and is most abundant along the lake margin. 2.3.3. Relationships between hippos from different environments To explore environmental constraints on δ13C dietary enamel values, carbon isotope values were grouped and tested for similarities between enamel δ13C and environment using a Kruskal-Wallis test (non-normality confirmed using a Shipro-Wilk test). This test was followed by a Tukey-Kramer (Nemenyi) post-hoc test using the Pairwise Multiple Comparison of Mean Ranks (PMCMR) package in R (Ihaka and Gentleman 1996). The Nemenyi post-hoc test uses mean rank sums on groups of unequal size (Tukey-Kramer method) and also applies a family-wise error method to control for false positives (Sokal and Rohlf 1994). These tests are necessary for this dataset, as the carbon isotopes values within these environmental groups come from profiles of 24 individuals, though still independent samples of diet. Certain profiles were excluded from the analysis, denoted by asterisks, with explanations given under Figure 2.2. 2.4. Results Isotope profiles are depicted graphically as boxplots in Figure 2.2 and grouped by environment in Figure 2.3, highlighting the remarkable range in δ13C values across all hippo profiles from different localities in Kenya. Overall, δ13C values from all hippo profiles exhibit a remarkable range of ~12‰. These values place hippo dietary isotopes within the dietary designation of mixed feeders (Figure 2.4; Table 2.2). Hippo enamel δ13C profile plots for individuals are presented in Appendix A, revealing the range and dietary breadth of this single species of megaherbivore across an entire country. Mean δ13C values range from 0.8‰ (the most enriched values) near Lake Turkana to -6.6‰ in Mwea Reserve (the most depleted). These results indicate that, at most, ~95% of the resources consumed by hippos in Kenya come from C4 plants, whereas at least ~40% of the resources are C4. Such a range in dietary flexibility in large African herbivores has only been observed in those with known mixed-feeding or seasonal diets, such as elephants or impala (Wronski 2002; Cerling et al. 2004; Owen Smith and Chafota 2012). This dataset reveals the diversity of hippo diets across environments in Kenya, and moreover that this dietary diversity is constrained within environments. The greatest ranges in carbon isotope values are from those that were from nuisance animals (Figure 2.2, denoted with an asterisk). Carbon isotope values from hippos separate statistically into four distinct groups, based on the broad physiognomic vegetation characterizations of the parks and reserves they came from: Bd, Wcd, wd, Fp (Tsavo, Mwea, Meru, Buffalo Springs, Chyulu, Olbolosat, Adhi Dam, and Kisumu); Fq, g, R, Wd/Wc (Mokowe, 25 Arabuko Sokoke, Mpeketoni, Minjila, Naivasha, Nakuru, Amboseli); We/wd (Laikipia, Maasai Mara); S (Turkana) (Figure 2.3; Table 2.3). In addition to exhibiting large ranges in δ13C values across profiles, certain hippo profiles also show large ranges in δ18O values across profiles. Such isotopic profiles include Chyulu (δ18O range: 4.7‰), Kisumu (δ18O range: 4.4‰), Mwea #2 (δ18O range: 4.8‰), Mwea – Gitaru Dam (δ18O range: 5.2‰), Meru (δ18O range: 4.2‰), K08-201 (δ18O range: 5.2‰), K01-TSW-291 (δ18O range: 4.8‰), Nakuru (δ18O range: 4.7‰) (see Appendix A). Oxygen isotopes in hippos reflect the isotopic value of their aquatic habitat due to their permeable skin that allows for high rates of cutaneous water loss (on land) and exchange (in water) (Luck and Wright 1964; Levin et al. 2006; Cerling et al. 2008). These ranges in δ18O values may represent changing habitat or changing values of the source water themselves, either from meteoric changes or other climatic forcing. 2.5 Discussion 2.5.1. Hippo dietary variability across Kenya Bulk hippo dietary flexibility, in both modern H. amphibius and throughout recent hippopotamus evolution, has been demonstrated through stable isotope analysis (Boisserie et al. 2005; Cerling et al. 2008; Harris et al. 2008). This dataset reveals the extent of hippo dietary variability within individuals, which was not previously known. Large ranges in isotope values are evident from nuisance animals, such as in Tsavo (K08201; δ13C range: 8.3‰), Naivasha - Crater Lake (NaivashaCL; δ13C range: 7.6‰), two Arabuko-Sokoke hippos (AS-168; δ13C range: 6.2‰ and AS-166; δ13C range: 4.4‰), but also in non-nuisance animals, such as Mwea #2 (δ13C range: 5.8‰), Ol Bolossat (δ13C range: 6.2‰), Meru (δ13C range: 6.9‰), and Tsavo #1 (δ13C range: 7.2‰) (Table 2.2). 26 Hippos are known crop raiders in many parts of Africa, and can cause expensive damages and loss of income for local famers (Weladji and Tchamba 2003; NaughtonTreves and Treves 2005; Kendall 2011; Kanga et al. 2012). This crop-raiding behavior is exacerbated by human impact and land-use change on their natural habitat, primarily for agriculture and livestock grazing (Lewison 2007; Kanga et al. 2011). Of the non-crop raiding individuals in this dataset, δ13C ranges across profiles can still be quite large. Of particular note are the four hippos from Mwea National Reserve, which all exhibit average δ13C enamel values between -5.7 and -6.6‰, with relatively large ranges in δ13C across the profiles (between 3-5.8‰) (Table 2.2). Mwea is categorized as Bd – “Somalia-Masai Acacia-Commiphora deciduous bushland and thicket”, and has dense bushy riparian corridors and woody vegetation (Chira and Kinyamario 2009). The depleted values for the four hippos from this reserve may indicate an abundance of C3 in the herbaceous understory, facilitated by abundant bush and woody vegetation (see Chapter 3). It is also important to consider dietary input from agricultural sources in this region. Mwea has been extensively developed in a large riceproduction scheme in Kenya (Kabutha and Mutero 2002), and though there have been reports of hippos raiding rice paddies in this area, the four hippos from Mwea were not known to have been killed for animal management purposes (S. Andanje, pers. comm.). Other profiles reveal divergences in ecology from populations in the same geographic area. For example, two tusks from the Lake Naivasha area (Crayfish Camp and Crater Lake) show how hippos record distinct environmental differences and disturbance in different parts of the lake basin (Figure 2.5). The profile from Crayfish Camp, a protected area with an abundance of grasses and herbaceous vegetation, has a mean δ13C value of - 27 1.7‰ +/- 1.3, a diet consisting of roughly 75% C4 resources. The hippo profile from the Crater Lake area, however, was shot for crop raiding, and lived in an area wherein much of the natural environment had been converted to agricultural land (S. Andanje, pers. comm.). This hippo profile exhibits a steady increase in δ13C over the course of its life until it was shot by KWS for crop raiding. Differences in δ18O between these two hippos reflect the difference in isotopic source waters in separate parts of the Naivasha lake basin (Ojiambo et al. 2001). These hippos reveal the ways in which these profiles capture heterogeneity within the same area, and how two hippos from the same geographic area can appear as though they come from two distinct ecological localities, though only 15 km apart. 2.5.2. Hippo canine isotopes as a record for environmental change Another way in which these isotope values can be used in modern environments is as a record of change within a given area using precisely dated canine enamel (Uno et al. 2013). Two profiles from Tsavo National Park reveal the utility of using canine enamel as a marker of environmental change (Figure 2.6). One radiocarbon-dated profile (K00-291) indicates that a hippo died in 1996, during a drought. Stable carbon isotopes from this profile indicate that the C3 component of this hippo’s diet increased over the last 3 years of its life, in addition to an enrichment in δ18O values. The radiocarbon-dated specimen K08-201 overlaps the end of the K00-291 profile and shows the same carbon isotope trend, followed by an increase in C4 resources to its diet as the drought improved. Both of these hippos came from a locality near a river in the northern part of Tsavo National Park – Mtito Andei. A third hippo profile (sample ages approximated using the average hippo growth rate of ~40 mm/yr) (Uno et al. 2013) complicates this 28 environmental record – K09-TSV, a hippo that lived in Mzima Springs, a naturally flowing spring that was unaffected by the drought. These hippos not only reveal dietary change forced by drought conditions in the national park, but variability across the park during a single ecological event. Serially sampled and dated hippo canines can be used as excellent records of environmental change, if from the same population. On a more local scale, it has been suggested that hippo tusk isotope profiles might record seasonal-scale variability (Passey et al. 2005; Souron et al. 2012). To addresses this question, one tusk (Turkana – Koobi Fora 1) was sampled at high resolution (2– 3mm) and radiocarbon dated in order to calculate its growth rate (~27 mm/yr). Even this resolution, however, fails to consistently capture the ~2‰ annual variability in δ18O recorded in Lake Turkana surface waters (Cerling 1996), except in a few instances, such as between 1978 and 1979 (Figure 2.7). Even at such high sampling resolution, hippos do not perfectly record local water conditions, pointing to the need to understand other factors that may determine δ18Oenamel, such as physiology. 2.5.3. Changing environments at the end of life Certain trends in both δ13C and δ18O are reflected across many isotope profiles in this dataset. Specifically, in the final 1–2 years of life (assuming a static growth rate of ~4 cm/yr, with a maximum growth rate of ~8cm/yr), many hippo profiles show a significant enrichment in δ18O in the last year of life relative to the rest of their life (Wilcoxan rank sum test with Bonferroni correction, P<0.05; Appendix A). Specific profiles which exhibit this trend include: (δ13C, δ18O), Nakuru (δ18O), Mwea #1 (δ13C, δ18O), Mwea #2 (δ13C), Chyulu (δ18O), Kisumu (δ13C), Meru (δ13C, δ18O), Mpeketoni (δ18O), Witu (δ18O), (δ13C), Buffalo Springs (δ18O), and Aberdares (δ13C, δ18O) (see Appendix A). Of these 29 individuals, the death circumstances are only known for Tsavo #1, Arabuko Sokoke #1, and the Aberdares tusks. All three of these hippos died in unusual environmental conditions – both Tsavo #1 and Arabuko Sokoke #1, died during a drought year, and the hippo from the Aberdares NP locality may have walked from somewhere else – possibly Lake Naivasha – as hippos are uncommon in the park and this individual was previously unkown in the park (S. Andanje, pers. comm.). These data indicate that, for some individuals, there may have been an environmental circumstance or change in behavior (such as migration or crop raiding) that may have contributed to their death. The onset of drought in the last year of life in the Tsavo #1 and Arabuko Sokoke #1 tusks resulted in steadily increasing δ18O values in tooth enamel recording their last year of life (Appendix A, Figure 2.6), and similarly, these conditions resulted in vegetation change, recorded in δ13C values in their final year of life, though the Arabuko Sokoke #1 hippo was shot for crop raiding (Appendix A). Without knowing the specific life histories of other hippo individuals in which these isotopic changes occurred in their final year of life, it is difficult to tell which scenarios are likely. In any case, these data highlight a uniform trend in enamel stable isotopes that may reflect larger environmental changes in the last year of life, and may provide evidence for local environmental change or drought in the fossil record. 2.5.4. Hippo dietary variability across environments Bd, Wcd, wd, Fp (Bushland/Wooded Grassland environments) The hippos that exhibit the most depleted isotope values throughout their dietary isotope profiles all come from distinctly bushy/wooded environments in Kenya. These areas include: Adhi Dam (within the Boni National Preserve), Tsavo National Park, 30 Mwea National Reserve, Meru National Park, Buffalo Springs National Reserve, Chyulu Hills National Park, Lake Olbolossat, and the Kisumu area (Lake Victoria). The ecological characterizations of these localities based on the potential natural vegetation map corresponds to on-the-ground ecological research in these areas. Generally, these localities have many C3 plants and woody vegetation. Chyulu and Tsavo West are ecologically similar to each other with similar representation of C4 grasses, flowering dicots, and shrubs in both parks (Jensen and Belsky 1989). Woody vegetation has increased in Tsavo since 1970 as the elephant population has decreased, similar to woody increases seen in Queen Elizabeth Park (Corfield 1973; Lock 1993; Leuthold 1996; Plumptre et al. 2010). Mwea National Reserve is similarly a wooded environment, comprised of bushland, woodland, and wooded grassland (Chira and Kinyamario 2009). Likewise, Buffalo Springs National Reserve is also a dry savanna woodland and wooded grassland ecosystem with riverine woodland along the Ewaso Ngiro River, where many hippos live (Wittemyer 2001; Ihwagi et al. 2010). Meru is likewise described as a dry wooded grassland/bushland environment (Neal 1984). The natural area around Lake Olbolossat is a forest, though hippos use only the 2–3 km near the lake itself (Njeri 2003; Kinyili 2014). Likewise, the environment in the Boni National Preserve is a dry coastal forest/woodland, diverse in both flora and fauna (Burgess et al. 1998; Tabor et al. 2010). The modern ecology of Lake Victoria Basin in the unaltered areas — Ruma National Park — is a wooded grassland/bushland, within which woody plants have encroached over the last 50 years with the loss of large browsers (Muriuki et al. 2003). Trees and other woody vegetation impart a strong influence on the floral composition of the herbaceous layer in Africa savanna environments. It is common for C3 31 herbs and dicots to flourish under isolated tree canopies in savannas, and increasing tree density in such savannas negatively influence grass density in those same localities through soil modification and competition for water (Scholes and Archer 1997; Hudak et al. 2003; Bond 2008; February and Higgins 2010). Some, though not all, of these localities have experienced increasing woody vegetation over the last 50 years as a result of large browser removal (Leuthold 1996; Muriuki et al. 2003), while other environments became more woody due to edaphic and/or climatological factors (Chira and Kinyamario 2009). The hippos from these environments all similarly record diets consisting of predominately C3 resources and are likely recording C3 herbaceous vegetation in these parks, which may have been facilitated by an abundance of woody plants. Fq, g, R, Wd/Wc (grassy woodland/grassland/riparian environments) Hippo dietary isotope values within this grouping largely reflect the more open, grassy nature of these environments. The most depleted average profile value within this group is the hippo from Arabuko Sokoke (-3.6‰), the most forested environment within these hippo habitats. The ecology of the Kenyan a coastal area is diverse, and now consists of grassland, scrub forest, and the biologically diverse Arabuko-Sokoke forest (Fq), which once covered much of the Kenyan coast (Moomaw 1960). Through logging and transformation of the coastal ecosystems to agricultural and rangeland purposes, much of the forest has been converted to grasslands, which are abundant on the coast (type g) (Oyugi et al. 2008). The basins of lakes Naivasha (Wd/Wc) and Nakuru (R), in the valley of the central rift, are both characterized as open to densely wooded Acacia woodland/more open savanna grasslands, transitioning to bushland/forests along the rift escarpment, with an abundant representation of Poaceae, Asteraceae, and Leguminosae in 32 areas where herbivores graze (Kutilek 1974; Ambrose and Sikes 1991; Maitima 1991; Mutangah 1994; Mutangah and Agnew 1996). The dominance of wooded grasslands in these immediate lake basins, though still a rich representation of woody vegetation, explains the more enriched δ 13C values of hippos from these lakes (Naivasha: -1.93‰, Nakuru: -0.8‰). Amboseli National Park, unlike many other Kenyan parks, has undergone a woodland/bushland reduction and a grassland expansion in recent years (Altmann et al. 2002; Western 2007). A more C4 inclusive diet is evident from the Amboseli hippo (-1.4‰) We/wd (edaphic/biotic wooded grassland) Hippo dietary isotopes from the Laikipia Plateau and Maasai Mara are statistically distinctive from other similar ecosystems in that isotope values from these profiles are more depleted than values from other similar savanna parks, the Fq, g, R, Wd/Wc hippos. Isotopic values from the Laikpia hippo tusk (-2.7‰) are difficult to explain, as the Laikipia Plateau is a large, diverse region with natural Acacia and Commiphora wooded grassland, xeric scrub bushland vegetation (Young et al. 1995; Kahindi et al. 2010). Though C4 grasses are dominant in wooded grasslands on the plateau, forbs are common, especially in well-drained soils or recently burnt grasslands (Taylor et al. 2005). These more depleted values could represent increased intake of C3 plants on the plateau, or this hippo could have been raiding crops or consuming resources from an overgrazed area, as land-use change for agricultural purposes is common in Laikipia (Young et al. 1998; Gadd 2005). Hippo habitats within the Mara River in Maasai Mara National Park are comprised of trees along the riparian corridor, grading into bushland and thicket, then into grassland (Olivier and Laurie 1974). This area – 1.5 km away from the water – is the 33 environment used by hippos for feeding. Hippos were noted to have consumed grass in 1974, though such dietary behavior may be out of date (see Chapter 3), as the Mara ecosystem was open and grassy due to high elephant browsing activity. Recently, woodlands have recovered due to over-grazing by livestock (Dublin 1986; Dublin et al. 1990; Reid and Ellis 1995; Lamprey et al. 2004). The more depleted carbon isotope values from the Mara hippo may reflect increasing C3 facilitated by woodland development. S (Somalia-Maasai semi-desert grassland/shrubland) The arid grasslands around the margins of Lake Turkana are distinctive and occur specifically in very arid environments (<200mm rainfall/yr) in northern Kenya and Somalia (White 1983). This distinctive halophytic C4 grassland around the lake margin consisting primarily of Sporobolus spp. is where lake-dwelling hippos feed. These dietary isotope values reflect this area and give a local signal of lake-side ecology (Cerling et al. 2003; Levin et al. 2011). 2.5.5. Hippo diets as a reflection of local environment Hippo dietary isotope values group into four distinct categories, which relate broadly to these physiognomic ecosystem types. It is important to note that hippos forage close to their aquatic habitats unless under ecological duress, such as overcrowded watering holes or insufficient herbaceous grazing lawns (Eltringham 1999; Klingel 2008). These hippo dietary profiles have revealed the flexible nature of their dietary strategies in greater detail than previously explored (Boisserie et al. 2005; Cerling et al. 2008), though dietary isotope profiles are still restricted and defined by food availability 34 within their environment. This result suggests that hippos are not selective feeders and do not seek out specific species of herbaceous plants, specifically C4 grasses as previously thought, as nearly all of these parks, reserves, and ecosystems have open grassy vegetation and grasslands within them. Likewise, these dietary profiles reveal distinctive differences even within the same ecosystem, such as the two hippos from Lake Naivasha (Figure 2.4). These data provide strong evidence that hippo dietary isotope values can serve as excellent markers of herbaceous C3/C4, but on a hyper-local scale. On an individual scale, these profiles record distinctive, specific records of local environmental change and stability. Though many profiles record isotope variability on the order of several permil across an individual’s lifetime, some isotope records showcase remarkable environmental stability in both δ13C and δ18O. These records come from a variety of environments, including lakes (the two Turkana hippos, Figures 2.7 and Appendix A), natural springs (Mzima Springs, K09-TSV; Figure 2.5), and rivers (Amboseli and Laikipia, Appendix A). These data are critical for informing paleoecologists on how to interpret isotopic data from fossil hippo enamel, especially from samples taken from hippo canine or molar enamel. Though it appears that, broadly, hippo δ13C values reflect those of local environment, there is still potential for bias by extreme ecological events, such as drought. Likewise, only in a few individuals is δ18O relatively stable over the entire course of hippo enamel growth period, meaning that fossil hippo enamel used for paleoaridity proxies may also be biased by extreme environmental events. With these caveats taken into consideration, hippo canine enamel in both modern and fossil environments are remarkable archives of environmental change, stability, and life history in these animals, and can serve as excellent records of local ecology, up to a 35 decade. 2.6 Conclusions This extensive dataset of serial enamel samples from hippo canines reveals the remarkable dietary variability within and among hippo individuals across environments in Kenya, ranging over 12‰ across all individuals. These data refine our thinking about modern hippo ecology and diets and their indiscriminate feeding nature within the confines of their environment. These data provide an important benchmark for natural ecology and dietary variability for hippos in modern environments that can help identify nuisance animals isotopically, as hippos can cause significant crop damage and income loss for agriculturalists (Naughton-Treves and Treves 2005; Kendall 2011; Nyirenda and Chansa 2011). Similarly, these hippo dietary isotope values provide a modern comparison for fossil and archaeological hippo isotope values. Previous studies on modern isotope values are derived from bulk enamel samples from molars (Cerling et al. 2008). These canine serial samples reveal the range of values within individuals. Though hippo dietary isotopes appear to reflect, to some degree, their environment of origin, isotopic indicators of diet still only reflect the environment immediately around their aquatic habitat, and thus reflect a local snapshot of their environment. Care must be taken when interpreting isotopic values of hippo enamel from the fossil record, and the extensive range and variability in their ecology and behavior must be considered. 2.7 Acknowledgements I wish to acknowledge the enormous contribution of the late Dr. Samuel Andanje of the Kenya Wildlife Service to this work, who facilitated harvest, collection, and 36 sampling of hippo canines across national parks and reserves. Without his work, this research and the resulting rich dataset would not have been possible. I also wish to thank T. Cerling, K. Uno, and S. Blumenthal for analytical assistance and constructive discussions about this dataset. 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Woodland open stand of trees at least 8m tall, with canopy cover of >40% and a grass-dominated herbaceous layer. Bushland open stand of bushes 3–7m tall, with canopy cover of >40%. Thicket closed stand of bushes 3–7m tall. Shrubland open or closed stand of shrubs up to 2m tall. Grassland dominated by grasses and other herbs and forbs, either with or without woody plants exceeding no more than 10% canopy cover. Wooded grassland dominated by grasses and other herbs and forbs, with woody canopy cover between 10 and 40%. 45 browsers (<25% C4 ) grazers (>75% C4 ) mixed C3 -C4 diet Chyulu Mwea 3 Lake Victoria - Kisumu Mwea 2 Mwea 1 Mwea - Gitaru Dam Adhi Dam OlOlbollosat Bolossat Meru NP *Tsavo 3 *Arabuko Sokoke 4 Arabuko Sokoke 3 Buffalo Springs Tsavo 2 Mara **Tsavo 1 *Naivasha Laikipia ***Aberdares NP *Arabuko Sokoke 2 Amboseli 2 Mpeketoni Minjila Naivasha Witu *Arabuko Sokoke 1 Amboseli 1 Nakuru Mokowe Turkana - Koobi Fora 2 Turkana - Koobi Fora 1 -10 -8 -6 13 -4 -2 0 2 δ C (V-PDB) Figure 2.2: Mean boxplots of δ13C values from hippo canine enamel profiles. Boxplot colors correspond to environmental groupings given in Figure 2.3. *denotes nuisance animals not included in statistical analysis. **died in drought. ***denotes animals with unknown provenance. 46 S We/Wd Wd/Wc R g Fq Wcd wd Fp Bd -10 -8 -6 -4 -2 0 δ13 C enamel Figure 2.3: Mean boxplots of enamel isotope values grouped by environment. Boxplot colors correspond to statistically significant ecological groupings (P values presented in Table 2.3). 2 47 Table 2.2: Hippo tusks sampled by locality. Starred hippos were excluded from analysis for reasons given below this table. Year of Length 𝛅13C Sample ID Locality Method SD death (mm) avg. ***Aberdares Aberdares NP 325 -2.7 1.0 KWSAdhidam Amboseli H1/H2 KWS-AMB0401 *AS168 *K00-AS-167 Arabuko Sokoke 2008 *AS166 Buffalo Springs Chyulu Kisumu K01-LAI-191 Adhi Dam Amboseli #1 Amboseli #2 Arabuko Sokoke #1 Arabuko Sokoke #2 Arabuko Sokoke #3 Arabuko Sokoke #4 Buffalo Springs Chyulu Kisumu Laikipia ca. 2008 known 495 -5.6 1.7 465 -1.4 0.6 ca. 2008 1996 known 425 -2.1 0.7 340 -1.7 1.6 1996 432 -2.3 1.3 2002 295 -3.6 0.5 1996 272 -4.2 1.3 known 465 -3.2 0.5 known 315 405 470 -7 -6.4 -2.7 0.9 1 0.3 345 275 525 -3.1 -5.1 -2.1 0.5 2.4 0.7 485 535 405 -0.1 -2.1 -5.7 0.4 0.4 0.6 2010 ca. 2000 Mara Meru KWS Minjila (Tana River) Maasai Mara Meru NP Minjila Mokowe Mpeketoni Mwea, Gitaru Dam Mwea 2009 Mwea NR MAR Mwea NP 2007 Mokowe Mpeketoni Mwea - Gitaru Dam Mwea #1 Mwea #2 2008 2005 known known 595 595 -5.9 -6.1 1.2 1.4 Mwea #3 2007 known 605 -6.6 1.0 *crop raider **died in drought ***unknown origin locality 48 Table 2.2 (continued) Sample ID Locality *KWSNaivasha KWS-NAIVCF-1108 KWSLNakuruKabutini-5 Olbolossat **K01-TSW291 Tsavo - 2009 (Mzima Springs) K08-201 Naivasha - CL Koobi Fora 2011 KEN-09-115 Turkana Koobi Fora 1 Turkana Koobi Fora 2 Witu Witu Naivasha - CF Nakuru Ol Bolossat Tsavo #1 Year of death ca. 2008 1996 Tsavo #2 Tsavo #3 *crop raider **died in drought ***unknown origin locality Method known 14 C/ known known Oct. 2007 1980 known ca. 2005 known 14 C Length (mm) 525 𝛅13C avg. -2.8 585 -1.9 1.3 315 -0.8 0.8 345 605 -5.5 -3.1 1.4 1.6 595 -3.2 0.6 535 -4.8 2.5 485 0.8 0.5 425 0.6 0.6 575 -1.9 0.4 SD 2.2 49 Table 2.2 (continued) Sample ID ***Aberdares KWS-Adhidam Amboseli H1/H2 KWS-AMB-0401 *AS168 *K00-AS-167 Arabuko - Sokoke 2008 *AS166 Buffalo Springs Chyulu Kisumu K01-LAI-191 Mara Meru KWS Minjila (Tana River) Mokowe Mpeketoni Mwea, Gitaru Dam Mwea 2009 Mwea NR MAR Mwea NP 2007 *KWS-Naivasha KWS-NAIV-CF1108 KWS-LNakuruKabutini-5 Olbolossat **K01-TSW-291 Tsavo - 2009 (Mzima Springs) K08-201 Koobi Fora 2011 KEN-09-115 Witu 𝛅18O avg. -2.6 -1.7 -5.7 -5.2 -4.5 -3.4 -3.2 SD Env. Lat. Long. 0.7 0.8 0.3 0.4 0.6 0.7 0.5 ? Fp wd/Wc wd/Wc Fq Fq Fq -0.4158 -1.5397 -2.7207 -2.7207 -3.3298 -3.3298 -3.3298 36.6667 41.4496 37.297 37.297 39.8772 39.8772 39.8772 -3 -3.9 -3.7 -3.9 -2.4 -4.1 -4.7 -2.3 0.6 0.3 1.5 0.4 1 0.4 1 0.8 Fq Bd Wcd wd We/Wd We/Wd Bd g -3.3298 0.4936 -2.549 -0.2209 0.3726 -1.5987 0.088 -2.2843 39.8772 37.6111 37.876 34.6387 36.7869 35.2782 38.1899 40.1392 -3.5 -1.4 -3.5 -4.1 -4.3 -3.5 -3.7 0.5 0.3 1.1 0.8 0.8 1.2 0.6 1 0.7 Fq Fq Bd Bd Bd Bd R R -2.232 -2.3904 -0.8259 -0.8259 -0.8259 -0.8259 -0.7729 -0.7876 40.8406 40.6968 37.6846 37.6846 37.6846 37.6846 36.3375 36.4107 -1.3 1.3 R -0.3834 36.1359 -2.3 -4.3 -4.4 1.1 0.8 0.5 wd Bd Bd -0.1556 -2.9642 -2.9835 36.4405 37.9104 38.0217 -2.2 1.4 1.2 -2.5 1.5 0.7 0.4 0.8 Bd S S g -2.1833 3.9472 3.9472 -2.3728 38.4166 36.1858 36.1858 40.3984 *crop raider **died in drought ***unknown origin locality 50 Table 2.3: P values of relationships between environmental groupings of δ13C enamel values from pairwise comparisons using a Tukey-Kramer (Nemenyi) test. Bd Fp Fq g R Fp 1 Fq <0.0001 <0.0001 g <0.0001 <0.0001 1 R <0.0001 <0.0001 0.99 0.99 S <0.0001 <0.0001 <0.0001 <0.0001 <0.0001 Wcd 0.23 0.73 <0.0001 <0.0001 <0.0001 wd 0.93 0.99 <0.0001 <0.0001 <0.0001 Wd/Wc <0.0001 <0.0001 1 1 1 We/Wd <0.0001 <0.01 0.05 <0.01 <0.01 51 Table 2.3 (continued) S Fp Fq g R S Wcd <0.0001 wd <0.0001 Wd/Wc <0.0001 We/Wd <0.0001 Wcd wd 0.94 <0.0001 <0.0001 <0.0001 <0.0001 Wd/Wc 0.02 52 n mammals (grey) = 1430 n hippos (blue) = 1410 250 C3 - diet C3 - diet closed canopy understory C4 - diet Mixed 200 150 100 50 0 - 25 - 20 - 15 - 10 δ13C enamel -5 0 Figure 2.4: Comparison of hippo profile isotope values (blue) vs. other mammal enamel isotope values (data from Cerling 2014). 5 δ13C%(VPDB)% 53 2$ 0$ !2$ !4$ !6$ !8$ !10$ !6$ !4$ !2$ 0$ 2$ 4$ δ18O%(VPDB)% Crater%Lake% Crayfish%Camp% 15%km% Figure 2.5: Comparison of two hippos from Lake Naivasha – Crayfish Camp vs. Crater Lake. 54 drought 5 0 δ13Cenamel 0 δ18Oenamel K00-291 δ18O K00-291 δ13C Mtito Andei K08-201 δ18O 13 K08-201 δ C K09-TSV δ18O Mzima Springs K09-TSV δ13C −5 −5 −10 1990 1995 2000 2005 Year Figure 2.6: Isotope profiles of three Tsavo hippo canines showing differential responses to drought in the park based on locality. 55 1960 1970 1975 1980 3 δ13C δ18O 4 2 3 1 2 0 1 −1 0 −2 −1 −3 1960 1965 1970 1975 1980 Year Figure 2.7: High resolution δ13C and δ18O enamel data from KF- δ18Oenamel δ13Cenamel 5 1965 CHAPTER 3 DECADAL DIET CHANGE IN HIPPOPOTAMUS AMPHIBIUS IN QUEEN ELIZABETH NATIONAL PARK, UGANDA 3.1 Abstract Megaherbivores (>1000kg) occupy important roles in African environments as ecosystem engineers, and changes to their ecology can result in major - and sometimes devastating - effects on their ecosystems. Despite their ecological importance, some basic aspects of their ecology (such as diet) remain poorly understood. Previous behavioral and gut/fecal content studies have categorized the common hippopotamus (Hippopotamus amphibius) as selective grazers, although isotopic analyses in East and Central African hippopotamus have suggested otherwise. Potential ecological drivers for dietary change remain poorly understood. We examine dietary flexibility in hippos over time within a single population using serial carbon isotope ratios (δ13C) in canine enamel from individuals that lived in Queen Elizabeth National Park, Uganda (QEP) near the Mweya Peninsula from 1960–1990s, a period characterized by significant expansion of bushy/woody vegetation following heavy poaching of elephants and other large mammals. Three hippo canines record a dietary switch within the Mweya hippo population from grazing (>80% C4) to a mixed C3/C4 diet (between 55–70% C4), likely driven by changes in the floral composition of the herbaceous vegetation layer. These data highlight the ability of hippos to cope with large-scale ecosystem change, and that 57 encroachment of C3 plants has occurred in both woody and herbaceous layers, as recorded in hippo canine enamel isotopes, further decreasing grazing capacity in the park. 3.2 Introduction 3.2.1 Hippopotamus ecology The common hippopotamus (Hippopotamus amphibius) is a large bodied herbivore found across Africa that lives in varied environments, ranging from arid savanna (see Chapter 2 for description) to forest (Eltringham 1999; Klingel 2008; 2013). They are semi-aquatic, spending the day resting in pools, rivers, and lakes, and feeding nocturnally on terrestrial vegetation (Vesey-FitzGerald 1960). Hippos generally feed less than 6 km away from their aquatic habitats and require as much as 20 kgs of foliage per day; these high feeding requirements and repeated journeys on land can lead to trampled and compacted ground, which can deter vegetation growth and transform soil properties (Lock 1972). Hippos can significantly restructure grazing areas, mowing tall grasses into short, closely-cropped lawns, which deters feeding by herbivores that prefer tall grasses, such as elephants and buffalo (Arsenault & Owen Smith 2002). Early studies of hippo ecology suggested that hippos selectively feed on C4 grasses (Field 1970; 1972). However, stable isotope ratios of carbon 13C/12C in hippo biological tissues reveal considerable variability in the consumption of C3 plants (herbs, forbs and shrubs) and C4 plants (tropical grasses <3000 m), indicating a range in diets from purely C3-based to purely C4-based (Boisserie et al. 2005; Cerling et al. 2008; see Chapter 2). Hippos have an unusual digestive anatomy, as they tend to masticate food to large particles that are inefficiently digested, relative to other ungulates, via nonruminating foregut fermentation, with long digesta retention times and low metabolic 58 rates (Clauss et al. 2004; Schwarm et al. 2006). Hippos are sometimes regarded as nuisance animals, as they frequently raid crops of maize, rice, and groundnut (Weladji & Tchamba 2003; Naughton-Treves & Treves 2005; Nyirenda & Chansa 2011); this nuisance status can impede conservation efforts. 3.2.2 Megaherbivores as ecosystem engineers Megaherbivores, such as hippos and elephants (Loxodonta africana), play critical roles in African ecosystems, including the maintenance of open grasslands, the modification of trophic guild structure (Owen-Smith 1992; Fritz et al. 2002; Duffy 2003; Asner & Levick 2012). Though hippos impact vegetation relatively near their aquatic habitats, elephants are important influencers of overall savanna habitat structure and function, and exert a higher level of influence (Beuchner & Dawkins 1961; Field 1971; Dublin et al. 1990; Barnes 2001). Elephants achieve this through selective browsing, predominantly on trees and thickets, which contributes to the persistence of open grasslands; high elephant populations can lead to woodland decline and can result in the extirpation of woody plants in areas of high elephant browsing (Dublin 1986). Elephants also facilitate other herbivore communities present by reducing predator cover in savannas (Laundré et al. 2001; Valeix et al. 2011). Elephant populations are declining across Africa due to poaching, and loss of these important megaherbivores will undoubtedly alter the stability and function of savanna ecosystems, and woody plant representation in savannas will increase (Wittemyer et al. 2014). In addition to this issue, climatological forcing and increasing global CO2 is facilitating bush encroachment (increasing numbers of unpalatable woody plants) across Africa (Ward 2005; Wiegand et al. 2005). We also have a poor 59 understanding of how elephant extirpation will affect other large-bodied keystone species, such as hippos, especially given the possibility for increasing C3 plants in areas where these animals live. We use ecological records and stable isotope proxy data from Queen Elizabeth National Park, Uganda (QEP) to understand the extent of ecological and behavioral change in a large population of hippos associated with a well-documented collapse in elephant populations. Our aims are to understand (1) the flexibility of hippo diets over time following a population collapse in elephants, (2) the ecological consequences of hippos feeding in disturbed environments, and (3) the extent of C3 herbaceous/bush encroachment that occurred following the elephant population collapse. We use stable isotope analysis of hippo canines to estimate the diet contributions of C3 and C4 plants to hippos inhabiting the Mweya Peninsula in QEP following an environmental disturbance (poaching and elephant culling in the 1970s) and associated large-scale environmental change (transformation of C4 grasslands to mixed C3/C4 herbaceous ground cover and wooded areas). 3.2.3 Stable isotope ecology in African mammals Stable isotope analysis is a powerful tool for understanding aspects of mammalian herbivore ecology in Africa, and is particularly useful for generating ecological records on time and space scales that are difficult or impossible to observe (Boutton et al. 1983; Ambrose & DeNiro 1986; Lee-Thorp & Van der Merwe 1987; Koch et al. 1990; Harris & Cerling 2002; Cerling et al. 2003; Cerling et al. 2004; 2005; 2008; Harris et al. 2008; Blumenthal et al. 2012; Cerling et al. 2013; Van Der Merwe 2013; Oelze et al. 2014). Stable carbon isotope analysis (δ13C) of herbivore tooth enamel can reveal the relative 60 proportions of C3 vs. C4 in the diets of African herbivores, and the relationship between δ13Cdiet and δ13Cenamel values is well-understood in African ungulates (Cerling & Harris 1999; Passey et al. 2005). Hippo canines represent excellent long-term archives of their ecology, since they are ever-growing and include isotopic input spanning 10 or more years of enamel growth (Laws 1968; Passey et al. 2005; Uno et al. 2013; see Chapter 2). Therefore, stable isotope analysis of tusks is ideal for quantifying primarily nocturnal feeding behavior of hippos, which is difficult to observe, particularly for generating multiyear to decadal-scale ecological records from deceased animals. 3.2.4 History of ecological research in Queen Elizabeth Park, Uganda Queen Elizabeth Park (Figure 3.1) is located in western Uganda in the Albertine Rift Valley, along the border of the Democratic Republic of Congo (DRC). The park is 1,979 km2, surrounding Lakes Edward and George, which are connected via the Kazinga Channel. Rainfall varies between 600 and 1400 mm/yr during March-May and in September- November, and is highest along the rift escarpment and along the Rwenzori Mountains, (Plumptre et al. 2010b). The soils in the park are rich in volcanic ash, leading to rich vegetation growth and high net primary production, with a high number of globally threatened and endemic plant and animal species (Field 1970; Plumptre et al. 2007a; 2010a). Detailed ecological research has been ongoing in QEP since the early 1950s due to the presence of the Nuffield Unit of Tropical Animal Ecology, which operated until the early 90s, focusing on issues of large herbivore and vegetation interactions with a wildlife management perspective (Plumptre et al. 2010b). This research has provided the foundation for our understanding of megaherbivore ecology in Africa (Eltringham 1999; 61 Klingel 2008), and also has documented two critical natural experiments, (1) a controlled hippo cull between the 1950- 60s and (2) extensive elephant poaching in the 70s, which can be used to understand long-term ecological change in African savannas (Petrides & Swank 1965). Throughout the park’s history, the effects of three major ecosystem drivers have been identified: elephant browsing, hippo grazing, and fire (Plumptre et al. 2010a). The hippo population was particularly abundant in the 1960s, leading to overexploitation of grasslands and competitive exclusion of other grazing ungulates. Considering the high dietary needs of wild hippos, the effects of significant overgrazing on the grass community by hippos at their peak population of about 14,000 individuals in the park was considerable, leaving bare-ground in areas where they fed (Field 1970; Plumptre et al. 2010b). The controlled hippo population culls began in 1958, stimulating new grass growth in areas around Lake Edward, particularly the Mweya Peninsula, that had been decimated by intense overgrazing and trampling by hippos (Thornton 1971). Subsequently, grasslands flourished and herbivore community diversity recovered (Eltringham 1974). In the 1960s and 70s, hippos were observed to feed predominately in grasslands on or near Mweya (i.e., within a few kilometers of Lake Edward) (Lock 1972) that were maintained by elephant browsing and fire (Field 1971; Dublin et al. 1990). Dominant grasses included Sporobolus pyramidalis, Bothriochloa inscultpa, Chloris gayana, Themeda triandra and Hyparrhenia filipendula, all C4 grasses that grow in heavily grazed areas (Lock 1972; Strugnell & Pigott 1978). Trees and shrubs in small thickets occasionally clustered around emergent Euphorbia candelabrum (Strugnell & Pigott 62 1978), and elephants preferred to feed in unburnt areas with Capparis tomentosa (thicket) and tall-grass S. pyramidalis (Field & Laws 1970). At the time of census in the mid-1970s, QEP had the highest large herbivore biomass (~20,000 kg/km-2) on Earth (Field & Laws 1970; Coe et al. 1976). From 1972 to 1980, during the reign of Idi Amin, management of all national parks essentially ceased, and widespread poaching decimated herbivore populations (Muwanika et al. 2003; Aleper & Moe 2006). Herbivore and vegetation surveys conducted in the mid-1970s on the Mweya peninsula revealed that, at the time, herbivore biomass was still relatively high and Hyparrhenia filipendula, Heteropogon contortus, and Bothriochloa insculpta grasslands were still present. Intensification of poaching activities, resulting in collapsing herbivore populations and a genetic bottleneck event (Eltringham & Malpas 1980; Muwanika et al. 2003) and C3 succession, occurred in the late 1970s (Yoaciel 1981). The population of elephants within the park fell from 4,139 to 150 individuals, and although hippo populations also contracted, they maintained genetic diversity during this poaching event, possibly due to influx of individuals from the DRC side of Lake Edward (Muwanika et al. 2003). Heavy wildlife poaching continued into the mid-1980s (Muwanika et al. 2003), by which time there was a significant expansion in the size and number of thickets and trees (predominantly Euphorbia candelabrum and Turraea robusta) due to the declining elephant population (Field 1971; Wyatt & Eltringham 1974; Dublin et al. 1990; Lock 1993). Hippo feeding intensified around the Mweya Peninsula, as low-level poaching kept hippos away from inland wallows and close to Lake Edward (Lock 1993; Plumptre et al. 2010a). Hippo range restriction resulted in a positive-feedback for thicket 63 encroachment: repeated trips onto land led to soil compaction, shunting rainwater into thickets and reducing fire fuel, further suppressing grass regrowth (Thornton 1971; Lock 1972; Eltringham 1974; Van Langevelde et al. 2003). The end result of these environmental changes was an increase in shrubs, trees, and thickets throughout much of the park, though most intensely around the Mweya Peninsula. 3.3. Materials and Methods 3.3.1 Sample collection and analysis Lower hippo canines were sampled from individuals who died on the Mweya Peninsula within Queen Elizabeth National Park. Date of death was assigned using recorded death dates (if known) or through radiocarbon dating of tusk enamel (Uno et al. 2013). Calendar years for hippos tusk samples were assigned using two methods. The 1960-1970 tusk was dated using bomb-curve radiocarbon dating (Uno et al. 2013). The death years for the other two tusks were known – 1991 and 2000. Using an average hippo lower canine growth rate of 4.4 cm/yr from five wild hippo tusks measured with bomb radiocarbon by Uno (2013), the calendar years were estimated for each sample in the profile assuming a constant growth rate. Samples of enamel were drilled at intervals along the length of the tusk using a diamond-tipped drill bit and Dremel tool. Enamel powders were treated with 2% H2O2 for 30 minutes to remove organics, then washed 3 times with distilled water. Enamel samples were reacted with 100% phosphoric acid in a common acid bath in a dual-inlet Carboflo carbonate device. Stable isotope ratios 13C/12C and 18O/16O of resulting CO2 were analyzed on an MAT 252, and stable isotope ratios are reported as delta (δ) values relative to the international carbon isotope standard, Vienna Pee Dee Belemnite (VPDB), 64 following the standard permil (‰) notation, where δ13C = (Rsample/Rstandard -1) x 1000, and Rsample and Rstandard. Enamel isotope values were corrected relative to an internal carbonate standard (Carrara marble) calibrated to VPBD and two in-house enamel standards. Dietary designations for hippos are given based on estimated dietary intake of C4 plants (lowland tropical grasses) and C3 plants (trees, shrubs, herbs). This is calculated using the isotope enrichment factor (ε*) between diet and herbivore tooth enamel of 14.1‰ (Cerling & Harris 1999), using the average isotopic value of modern C3 and C4 plants in eastern and central Africa (Cerling 2014; Cerling et al. 2015). 3.4. Results Stable isotope analysis revealed significant differences (Kruskal-Wallis and Nemenyi post-hoc test, P<0.05) in the isotopic composition of tusk enamel among the three Mweya hippos (Table 3.1). The 1970 tusk reveals a diet of greater than 80% C4 grass intake from ca. 1960 to 1970 (Figure 3.2 and Appendix B). The 1991 hippo profile is significantly more depleted in δ13C than the 1970 profile, indicating a mixed C3/C4 diet (ca. 65% C4) from 1982 to 1991, following collapse of the elephant population in Queen (Figure 3.2, Appendix B). The third tusk includes the time interval from 1985 to 2000 with an estimated diet ranging from ca. 55 to 70% C4. Both the 1991 and 2000 individuals show local minima in δ13C values for the ca. 1988–1989 interval. Previously published hippo molar enamel isotope values from the DRC side of Lake Edward during the 2000s and 1990s are similar those from serial samples in QEP, with 1990s DRC values ranging from -7 to -2‰, 2000s DRC values -7 to -1‰, though slightly more depleted than QEP values (Cerling et al. 2008). 65 3.5. Discussion 3.5.1 Hippos dietary change on the Mweya Peninsula, 1970–2000 Hippos have been characterized as selective C4-grazers that feed on short grasses and sedges (which comprises 95-99% of their diet), supplemented by forbs (Owen-Smith 1992). This dietary classification was established by stomach content analysis done by Field (1970), conducted on hippos in QEP during the 1960s, when grasses were abundant. Bulk stable isotope analysis of hippo tooth enamel values from individuals across East Africa have already revealed the diversity of hippo diets from different localities (Boisserie et al. 2005; Cerling et al. 2008). Changes in hippo diet within the Mweya Peninsula hippo population track changes in herbaceous groundcover on the peninsula. Environments where hippos are now abundant look strikingly different from those of the 1960s, where herbaceous groundcover on the Mweya Peninsula consisted predominately of grazing-tolerant C4 grasses (Lock 1972; Strugnell & Pigott 1978). In surveys conducted in 1992 and 2009, Plumptre and others (2010) found that Cynodon dactylon (a C4 grass), Commelina diffusa, C. africana (C3 herbs), Asystasia gangetica (C3 forb), Cyanotis foecunda (a C3 flowering herb), Achyranthes aspera (C3 herb), Ocimum suaveolens (C3 herb), Oplismenus hirtellus (C4 grass), and two Cyperus species have become dominant. Thus, hippos feeding areas from the 1960s have changed almost completely from C4 grasses to predominantly C3 herbaceous groundcover (Lenzi Grillini et al. 1996), which is reflected in hippo dietary isotopes. Hippo canine carbon isotopes record this herbaceous vegetation change within the Mweya Peninsula, and reveal the extent of encroaching C3 herbaceous plants in the area. The expansion of nongrassy ground vegetation further restricts grazing 66 capacity of the peninsula and decreases herbivore biodiversity and biomass. 3.5.2 Long-term effects of elephant poaching The effects of elephant poaching in the park have caused a restructuring of QEP’s ecology. Aerial photographs and photomosaic analysis of vegetation types within the park between 1950 and 2006 indicate an increase in woody cover of ~30% across QEP (Plumptre et al. 2010b). Civil war in the DRC in 1998 led to heavy poaching in Virunga National Park and other areas near QEP, resulting in herbivore migration to QEP and contributed to the partial recovery of herbivore communities, which may explain the delayed later succession of woody plants (Plumptre et al. 2007b). Although there is some lag time between early succession of woodlands and wooded grasslands in savannas, the effects of decreased vegetation maintenance by megaherbivores and fires have already resulted in bare-ground, scrubby/bushy environments where hippo feeding areas once were significant (Plumptre et al. 2010b). Elephant populations have been increasing in the park since 1990 (from ~500 to almost 3000 in 2005; Plumptre et al. 2010a) inhibiting only very recent woody cover encroachment in the park in some areas, but not in the Mweya peninsula (Plumptre et al. 2010a). Our findings also reveal that hippos are not such selective feeders that they will only feed upon short grass, but can shift their diets to accommodate increased C3 herbaceous groundcover when preferred grasses are no longer present. Therefore, hippos track herbaceous vegetation change, and carbon isotopic records from serially sampled hippos tusks may provide a much needed resource for investigating historical ecological change across tropical grassy biomes in Africa. This approach may also be useful for augmenting wildlife monitoring and management efforts of populations that are not being 67 regularly studied due to conflict or cannot easily be observed. 3.5.3 Elephant poaching and the future of African savannas Although the deleterious effects of human conflict and wildlife poaching as a mechanism for ecosystem destabilization are known, little has been studied on the longterm ecological effects of overharvesting on ground vegetation in savanna ecosystems (Dudley et al. 2002). Our findings demonstrate that in addition to bush encroachment (Prins & van der Jeugd 1993), elephant poaching results in C3 encroachment in the herbaceous ground layer of savannas as well. Therefore, the long-term and wide-reaching effects of bush encroachment, such as changes in nutrient composition of soil, depleting %C, and %N in heavily encroached areas (Hudak et al. 2003), are further compounded by suppression of C4 grass growth, a critical resource for numerous mesoherbivores in savanna ecosystems. The combined effects of bush and herbaceous encroachment forces certain herbivores outside of protected areas, compromising management and conservation efforts. The combined effects of bush encroachment, which has already occurred in many places in Africa (Scholes & Archer 1997; Roques et al. 2001; Ward 2005; Wiegand et al. 2005), and elephant poaching, which is occurring at unprecedented rates (Wittemyer et al. 2014), are likely to further exacerbate habitat degradation and suppress populations of grazing herbivores. Further characterization of herbaceous vegetation change in protected areas may be useful for understanding the rate of C3 encroachment and to devise management strategies for African savanna parks in a changing global environment. 68 3.6. Conclusions These isotopic data highlight the flexible nature of hippopotamus diets within populations over time, and show that hippos are able to adjust their diets after a major ecological shift. Decreasing elephant populations around Africa should not negatively inhibit hippo populations because hippos can adapt to the widespread changes in herbaceous vegetation. Hippos are key ecological engineers, consuming and spreading nutrients on land and transforming soil properties and landscapes that increase environmental heterogeneity. Hippo canine profiles can preserve a 10-year, or longer, record of diet history. As such, serial samples in hippo canines are long-term records of African ecology. In Queen Elizabeth Park, Uganda, these isotope records of hippo enamel indicate changes in C3-vegetation, not only in woody vegetation as a result of elephant culling, but also in herbaceous vegetation layers. These data are critical for wildlife management within African savanna parks, as increasing C3 vegetation in both woody and herbaceous layers of savanna parks decreases grazing capacity for grassfeeding herbivores. This outcome may result in pushing grazers outside of protected areas, posing serious threats for conservation in Africa. 3.7. Acknowledgments I would like to thank H. Klingel, S. Blumenthal, and T. Cerling who have all provided important, constructive comments on drafts of this manuscript. H. Klingel provided two of the three canines for this study. I would also like to thank the Uganda National Counsel for Science and Technology for granting permission to conduct this research and export samples. I thank the members of the Research Monitoring and Conservation Unit at the Uganda Wildlife Authority who granted us permission to 69 conduct this research, Aggrey Rwetsiba and Fred Kisame, and also Margaret Driciru, acting QEP park warden, and Nelson Guma, Conservation Area Manager. This study was funded by grants from the Geological Society of America (#9169111), the Global Change and Sustainability Center, NSF GRFP funding to KLC, the National Geographic Society (YEG #9349-13), a Leakey Foundation grant to SAB, the Wenner-Gren Foundation (#8694), NSF Doctoral Dissertation Improvement #1260535 Grant to SAB and NSF grant #0621542 to TEC. 3.8. References 1. Aleper, D. & Moe, S.R. 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The red circle highlights the Mweya Peninsula. 77 Table 3.1: δ13C values (mean and range) for the three hippo canines. P-values from Kruskal-Wallis with Tukey and Kramer (Nemenyi) post-hoc test (executed in R) verify that the three hippos have distinctive feeding niches. ID KL Queen VIC Q-09KL Length (cm) 37 Death Year 1970 35.5 1991 66.5 2000 δ13C mean -0.1 δ13C range -2.0 known -3.3 -2.5 <0.0001 - <0.01 known -3.0 -2.5 <0.0001 <0.0001 - Method 14 C P-value P-value (vs. 1970) (vs. 1991) <0.0001 P-value (vs. 2000) <0.0001 78 13 δ C enamel 0 -5 elephants in decline, regeneration and spread of Acacia and thickets; decreased fire frequency influx of migrant animals due to war DR Congo >80% C4 diet C3 /C4 diet (grazer) (mixed feeder) post 1958 hippo cull, highest herbivore biomass, politcal unrest, heavy poaching expanding grasslands 12000 8000 2000 4000 hippos elephants 4000 0 0 1960 1970 1980 Calendar Year 1990 2000 Figure 3.2: Carbon isotope profiles of the three Queen Elizabth hippo serial samples (1970 tusk in black, 1991 tusk in dark grey, 2000 tusk in light grey) and number of hippos (dotted line) and elephants (solid line) present in the park over time (Olivier 1991; Plumptre et al. 2010a). The chronolgy of major ecological and political events is listed above the graph. CHAPTER 4 HOLOCENE PALEOENVIRONMENT IN KENYA, WITH PERSPECTIVES ON HOLOCENE ARCHAEOLOGY AND PLEISTOCENE ENVIRONMENTAL CHANGE 4.1 Abstract Paleoenvironmental research in East Africa has long relied on the relationship between orbital forcing, regional climate, and vegetation response to reconstruct regional paleoecology, yet the connections between these three factors remains poorly understood. An interbasinal reconstruction (northern Lake Turkana and southwestern Lake Victoria) of Holocene paleoecology in Kenya may clarify these relationships. Previous climatological studies indicate that Lake Turkana responded abruptly to a weakening monsoon system at the end of the African Humid period (5.5 kya), forced by decreasing solar insolation (deMenocal et al., 2000; Garcin et al., 2012), whereas Lake Victoria responded more gradually to orbital forcing, as indicated by ẟDwax values (Berke et al., 2012a). Tooth enamel stable isotope records from Turkana also indicate a change in the diet of herbivores in the Turkana Basin with increasing C3 dietary components. Comparisons of early Holocene tooth enamel and leaf wax biomarker data reveal contrasting environments in Turkana, wherein herbivore diets include an abundance of C4 resources, but leaf waxes indicate fluctuating C3-C4 input to the sedimentary record. In the Lake Victoria Basin, both ẟ13Cwax values and tooth enamel dietary isotopes indicate 80 an abundance of C4 resources in the basin throughout the Holocene. This is in contrast to the pollen record, which indicates high (~40%) Moraceae (mesic tree) presence in the early Holocene, which decreases until present, and low (~10%) Poaceae (grass) pollen, which increases until present. These contrasting paleoecological records highlight disagreements noted in other studies (Feakins et al., 2013; Levin, 2015), calling for reevaluation of either pollen or leaf wax records, and how they may relate to carbonate proxies of paleoenvironment. Furthermore, these data do not support a uniform shift in ecology across Kenya, in accordance with insolation forcing, indicating that regional responses to orbital forcing must be assessed basin-by-basin. Finally, these data indicate that the same environmental factors which may have influenced the appearance of early herders in the Turkana Basin (i.e., abrupt climatological change and potentially ecological change) was not a factor in the Lake Victoria Basin, as gradual climate change did not result in abrupt, or even significant, ecological changes. 4.2 Introduction 4.2.1 Paleoecology and Holocene orbital forcing in East Africa Much research has been dedicated to understanding the relationships between climate and ecology in East Africa; specifically, what controls the distribution of woody and non-woody vegetation in modern (Coughenour and Ellis, 1993; Good and Caylor, 2011; Mills et al., 2012; Ratnam et al., 2011; Sankaran et al., 2005; Scholes and Archer, 1997), future (Bond and Midgley, 2000; Buitenwerf et al., 2012; Cramer et al., 2001; Moncrieff et al., 2013; Murphy and Bowman, 2012), and past environments (Berke et al., 2012a; deMenocal, 2011; Sinninghe Damsté et al., 2011; Willis et al., 2013). Establishing the connection between orbital forcing, climate change, and ecological response in the 81 geologic record has been central to this research (Ashley, 2007; Kingston, 2007; Kutzbach et al., 1996; Magill et al., 2013a; 2013b; J. D. Marshall et al., 2007; Maslin and Christensen, 2007; Verschuren et al., 2009). Many causal connections linking climate to ecology in African environments are missing and/or poorly understood (deMenocal, 2014; Kingston, 2007; Levin, 2015; Potts, 2013). Researchers have identified Holocene paleoenvironment as the key to understanding the connection between orbitally forced climate and ecology in East Africa (deMenocal et al., 2000; Gasse, 2000; Mayewski et al., 2004; Renssen et al., 2003; Tierney et al., 2011a). The early Holocene was a time of high insolation forcing, with evidence for high humidity and a stronger monsoon system across many parts of Africa, called the “African Humid Period” (“AHP”) (deMenocal et al., 2000; Tierney et al., 2011b). This event terminated sometime around 5.5 kya, forced by decreasing insolation and a weakening regional monsoon. Despite the abundance of climate records and modeling studies that have illuminated the nature of Holocene orbital forcing and terrestrial climate, associated ecological records for this time period are lacking. This is especially true in Kenya, where the anthropological and archaeological record is one of the richest in Africa. Studies linking changes in Holocene climate and ecology have revealed interbasinal differences in lacustrine records and have illuminated the complex ways in which ecology and environmental forcing are linked (Berke et al., 2012a; 2012b; Castañeda et al., 2009; Costa et al., 2014; Vincens et al., 2010). Currently, though, less attention has been given to records of ecology from these basins, and whether vegetation proxies indicate changes associated with climate dynamics during this time period. Such data would allow researchers to better interpret records of ecology from deeper time periods, and gain 82 insight into the ways in which vegetation dynamics in East Africa relate to orbital forcing on a regional scale. 4.2.2 Holocene archaeology and paleoenvironment The Holocene archaeological record documents the transition from foraging to food production, a fundamental shift in subsistence economy that is among the most significant cultural adaptations in human history. African food production began and spread in unusual ways: herding (i.e., keeping domestic animals without sole reliance on livestock for subsistence) preceded plant cultivation in many parts of Africa, whereas farming was the first form of food production in most parts of the world (Marshall and Hildebrand, 2002). Herding originated in the eastern Sahara around ~9,000 BP and spread through varied environments, reaching the southern tip of Africa seven millennia later. Herding spread into NW Kenya’s Turkana Basin around 4.5 ka (Barthelme, 1985), shortly after the termination of the AHP. The use of domestic livestock is not seen in the Central Rift, approximately 500 km south, until ~3 ka (Ambrose, 1998; GiffordGonzalez, 2000). Early herding in Kenya, particularly around Lake Turkana, coincides with social and cultural changes, such as construction of megalithic architecture and production of Nderit Ware, a distinctive and previously unknown pottery (Bower, 1991; Grillo and Hildebrand, 2013; Hildebrand et al., 2011). People may have begun herding in response to shifting climatic conditions that made it advantageous to guide animals to optimal environments and water sources (Marshall, 1990; Owen and Renaut, 1986; Smith, 1992). Many authors have suggested that changing climatic conditions may have been responsible, directly or indirectly, for the spread of herding as insolation-induced mesic conditions during the early Holocene 83 (11.7-8.2 ka; following Holocene formal stratigraphic designations of Walker et al., 2012) gave way to changing climates during the middle Holocene (8.2-4.2 ka) and more arid climate regimes during the late Holocene (4.2 ka-present) in many parts of eastern Africa (deMenocal et al., 2000; Tierney et al., 2011b; Walker et al., 2012). For example, early herding around Lake Turkana (the lower portions of the Turkana Basin, Figure 4.1) coincides with a sharp decrease in regional humidity (Owen et al., 1982), as indicated by the position of paleo-lake terraces (Garcin et al., 2012). In contrast, the Victoria Basin (or more specifically, the Winam Gulf area, Figure 4.1) sees livestock slightly later, c. 3.5 ka, and leaf wax deuterium isotopes indicate a more gradual decrease in humidity, in contrast to abrupt climatological changes in the Turkana Basin (Berke et al., 2012a; Prendergast, 2010). Although there is a general chronological association between the end of the AHP and the southward spread of herding, connections between changes in climate, terrestrial ecology, and human behavior are complex and often not synchronous between basins (Blome et al., 2012; Tierney et al., 2011a). Establishing causal links between regional climate change and the spread of herding is complicated due to issues of resolution and interpretation of archaeological and paleoenvironmental datasets, and also due to dilemmas over whether chronological correlations imply causal association. Empirical records for terrestrial paleoecological conditions directly associated with archaeological material are surprisingly limited. Paleoecological data from stable isotopes can shed light on changes in the distribution of woody cover, an important determinant of overall ecosystem structure in the tropics (see Chapter 2 for a thorough description of woody cover and African ecosystem classification). Woody cover distribution is influenced by 84 mean annual precipitation and seasonal distribution of rainfall, which, in turn, affect plant and herbivore biomass (Bell, 1982; Good and Caylor, 2011; Murphy and Lugo, 1986; Scholes and Archer, 1997). Because of these complicated environmental relationships, inferring ecosystem change from regional climate data is insufficient and limits our understanding of the ecosystems in which early herders lived, either by choice or circumstance. Three possible predictions for paleoecological changes through the Holocene include: 1) C3 plant dominance during the later Holocene due to rainfall seasonality and anthropogenic influences, such as amplified prevalence of grazing animals after 5 Ka (Sankaran et al., 2005); 2) increased warm season rainfall in the early Holocene causing C4 grasses to out-compete C3 woody plants prior to 5 Ka; 3) the system may also oscillate between more C3 and C4 during high insolation periods, as there may be a destabilizing effect of rainfall excess on savanna ecosystem structure (Sankaran et al., 2005). Furthermore, floral change seen in northern localities may represent a local pattern distinct from the more southerly localities, and manifest sooner and more abruptly, because changes in insolation are dampened at lower latitudes (Laskar et al., 2004). Comparing paleoecological proxies for C3 and C4 cover from archaeological sites in NW Kenya (Turkana) and SW Kenya (Victoria) will evaluate these regional differences and provide insight into the distinctive archaeological records of early food producers from these basins. This chapter will take a multiproxy approach to paleoecology by combining isotopic data from archaeological mammalian herbivore teeth, lacustrine leaf wax biomarkers, and pollen to understanding how basinal-scale vegetation dynamics relate to 85 climate forcing. This approach will help account for differences in spatial and temporal integration between these two proxies, and will provide a deeper, more nuanced understanding of the relationship between climate and vegetation change in East Africa. Records will be explored in two lake basins with two distinctive archaeological histories for early herding in eastern Africa. These lake basins exhibited differing responses to Holocene climate forcing: the Lake Turkana Basin responded abruptly to termination of the AHP (deMenocal et al., 2000; Garcin et al., 2007) and the Lake Victoria Basin responded more gradually to the termination of the AHP (Berke et al., 2012a). Resulting data will indicate what environmental changes, if any, occurred at Holocene archaeological sites in Kenya within the context of solar forcing and changing climate dynamics (Claussen et al., 1999; deMenocal et al., 2000; Garcin et al., 2012; Murphy and Lugo, 1986; Stager et al., 2003). 4.3 Study Design and Location 4.3.1 Study design Understanding how ecosystems changed over time is assisted by δ13C analysis of herbivorous mammalian tooth enamel as a proxy for changes in local plant biomass and animal behavior. Tooth enamel samples taken via identical sampling strategies from similar taxa found at Holocene archaeological sites in Lake Turkana and Lake Victoria are analyzed for δ13C and δ18O. Isotope data from archaeological specimens are compared to previously published modern values (Cerling et al., 2008; Kohn et al., 1996; 1998; Levin et al., 2006; Schoeninger et al., 2002; Cerling et al., 2015), leaf wax biomarkers for Lake Turkana (this study), and Lake Victoria (Berke et al., 2012b) in order to assess how site-specific ecology relates to basinal scale records. Three-hundred 86 thirty-three tooth enamel samples were analyzed (93 from six Turkana sites, 49 from Ele Bor, and 189 from three Lake Victoria sites) to answer these questions (Figure 4.2), generating a dataset that substantially exceeds previous Holocene isotope studies in eastern Africa (Ambrose and DeNiro, 1989; Ambrose and Sikes, 1991; Balasse and Ambrose, 2005a). δ13C data from enamel and leaf wax biomarkers provide direct isotopic information relating herbivore ecology and terrestrial ecosystem structure, respectively, to orbitally-mediated climate change. In particular, herbivore fossil enamel isotopes are useful for interpreting local, terrestrial ecological changes associated with calculated insolation and evaluating inferred changes in monsoon strength based on previously published records of lake levels, sedimentary biomarkers, and aridity. These data are key for understanding the local manifestations of changing monsoon strength across basins and provide quantitative data for ecological changes that occurred before, during, and after the spread of herding throughout Kenya. 4.4 Materials and Methods 4.4.1 Archaeological tooth enamel Teeth from archaeological sites in Lake Turkana (including the site of Ele Bor, as somewhat higher elevation, ~800 m a.s.l.) and Lake Victoria (primarily from the Winam Gulf) were sampled for δ13C and δ18O from collections in National Museum of Kenya (NMK). A new Turkana site was found (“Equid Site”) and existing collections were supplemented by field collected specimens during survey with the Later Prehistory of West Turkana project led by Drs. Elisabeth Hildebrand and John Shea (Stony Brook University). Samples from Ele Bor (Site M), 200 km east of Lake Turkana, were drilled from collections stored in the NMK. Samples were photographed and drilled with a 87 diamond tipped drill bit using a Dremel tool and stored in 1.7 ml centrifuge tubes following destructive sampling procedures outlined by the National Museums of Kenya. Samples were prepared for isotope analysis (δ13C and δ18O) at the University of Utah. Archaeological enamel prep involves treatment of samples (1-0.5mg enamel) with a weak acid (0.1M buffered acetic acid) to for 30 minutes remove labile carbonates, weighed into silver capsules and dried under vacuum at 200 °C before digestion in a common phosphoric acid bath at 90 °C for 15 minutes. Samples were analyzed for δ13C and δ18O on a Finnigan MAT 252 coupled to a Carboflo dual-inlet carbonate device (common phosphoric acid bath at 90°C for 15 minutes reaction time). Stable isotope ratios are reported as delta (δ) values relative to the international carbon isotope standard, Vienna Pee Dee Belemnite (VPDB), using the standard permil (‰) notation, where δ13C = (Rsample/Rstandard -1) x 1000, and Rsample and Rstandard are the 13C/12C ratios of the sample and standard, respectively. Standard deviation of an internal standards (Carrara marble, MHS, FRS) was between +/- 0.1 to 0.2‰. Modern samples were corrected to MHS and fossil samples to FRS for the temperature effect for phosphoric acid digestion of tooth enamel (Passey et al., 2007). Dietary designations for herbivores are based on estimated consumption of C4 plants (tropical grasses) and C3 plants (trees, shrubs, herbs), which is calculated using a hypothetical 100% C4 diet and an isotope enrichment (ε*) between diet and herbivore tooth enamel of 14.1‰ (Cerling and Harris, 1999). Estimated carbon isotope composition of tooth enamel for a 100% C4 diet is based on the average isotopic value of modern C4 plants in eastern and central Africa (-12.9‰, n = 764) (Cerling, 2014). Changes in the isotopic composition of atmospheric CO2 due to the combustion of fossil fuels between 88 modern environments and pre-industrial Holocene (presented here as δ 13C1750) were accounted for by adding 1.6‰ to the modern average δ13C of C4 plants (Cerling and Harris, 1999; Francey et al., 1999; Keeling et al., 2010). 4.4.2 Sedimentary leaf wax biomarkers Early Holocene lacustrine sediments were collected from Area 103 near Lake Turkana (Figure 4.1) in 2011. Stratigraphic correlation to previous maps of these deposits (Owen and Renaut, 1986) provides ages for the sediments (11.5 - 5.8 Kya, dates from mollusk shells in situ). Compound specific isotope analysis (CSIA) was completed at the Pennsylvania State University. Eighteen leaf wax biomarker samples were extracted using an Automated Solvent Extractor in 9:1 DCM:Methanol, purified by column chromatography. The resultant polar fraction containing n-alkanoic acids were derivitized into fatty acid methyl esthers (FAMEs) for isotope analysis using a phthalic acid standard (Arndt Schimmelmann, U. Indiana) with a known δ13C value of -73.16‰. δ13C values of FAMEs were corrected to their original carbon isotopic value after methylization using a mass-balance correction of the derivitization standard. Percent C4 input was calculated using endmember values from previously published values (-22 to -34‰) (Castañeda et al., 2009; Magill et al., 2013a). 4.5. Results 4.5.1 Lake Victoria Basin tooth enamel isotopes Data from herbivores analyzed from archaeological sites in the Lake Victoria Basin are presented in Table 4.1 and graphically in Figure 4.3. Stable isotopes from modern comparative tooth enamel are presented as δ13C1750 values from SW Kenya and 89 northern Tanzania – within the Lake Victoria Basin or the greater Loita-Mara-Serengeti ecosystem, where possible. Detailed analysis of the Gogo Falls fauna is presented elsewhere (Chritz et al., 2015). 4.5.1.1. Luanda Isotopes from tooth enamel in the early Holocene site of Luanda (9.7 – 8.5 Kya) reveal that the diets of herbivores from this site indicate that the diets of these animals were predominately dominated by C4 resources. The exceptions are hippos (-4.2‰ ± 1.6) and bushpigs (Potamochoerus sp. -10.2‰ ± 2.4), which both exhibit isotope values that indicate mixed C3/C4 diets, but both of which are within the range of isotope values seen in these animals in modern environments. 4.5.1.2. Wadh Lang’o Tooth enamel isotopes from herbivores at this site exhibit dietary isotopes that are all almost entirely >80% C4. These isotope values are very similar to their modern counterparts. Interestingly, Caprini (Subfamily Caprinae; domestic sheep and goats) indicate an entirely grazing diet. This tribe includes both sheep and goats, which in modern environments are separated in dietary space into grazers (sheep) and browsers/mixed feeders (goats) (Balasse and Ambrose, 2005b; Chritz et al., 2015). 4.5.2 Lake Turkana Basin tooth enamel isotopes Data from herbivores analyzed from archaeological sites in the Lake Turkana Basin are presented in Table 4.2 and graphically in Figure 4.4. Stable isotopes from modern comparative tooth enamel are presented as δ13C1750 values from Turkana, from collections stored at the Turkana Basin Institute and the National Museums of Kenya’s 90 Koobi Fora field station, and also field collected specimens. 4.5.2.1. Early Holocene sites – GaJj11, FxJj12, “Equid Site” Isotopic data from these sites were combined due to low densities of faunal remains from these sites. All taxa analyzed for isotopes indicate diets that are primarily C4, and all species are quite similar to their modern counterparts. 4.5.2.2. Lothagam Harpoon Site Data from mammals at this site indicate an abundance of C4 resources in their diet. The exception to this is an analysis of a single baboon molar (Papio anubis), which yields an isotope value of -6.9‰. This is more enriched than modern Turkana baboons (10.9‰), indicating more C4 resources in the mid-Holocene specimen, though it is still a mixed feeder. 4.5.2.3. Dongodien, GaJj2 Stable carbon isotope analysis from the abundant faunal remains at these two later Holocene pastoralist sites reveals an interesting departure from the dietary isotopes of fauna from the previous sites. Isotopes of the majority of these taxa (6/10) indicate primarily mixed C3/C4 feeding (<80% C4). Looking to ecologically flexible taxa, such as Aepycerotini (impala, -7.16%), Hippopotamus (-1.55% ± 1.77), and Antilopini (gazelles, -3.24% ± 0.98), there appears to be more C3 resources. Likewise, domestic caprines are also within the mixed C3/C4 dietary realm (-4.01% ± 2.33). Faunal analysis of domestic taxa from this site indicates that both sheep and goats were present (Marshall 1984), which makes these mostly mixed feeding results for both species even more intriguing. 91 4.5.3 Lake Turkana Basin leaf wax biomarkers Stable carbon isotopes from leaf wax biomarkers for early Holocene deposits indicate varying inputs from C3 and C4 plants to the sedimentary column during the early Holocene (11.1 – 7.4 Kya, Figure 4.5). These data indicate %C4 input to range from 72% to 48% over this time period, with isotope values that are comparable to other Holocene lacustrine records in terms of range in isotopic values (Costa et al., 2014). These data reveal apparent shifts in vegetation and significantly more C3 representation that are not apparent from the tooth enamel records for the same time period (Figure 4.5). 4.5.4 Ele Bor tooth enamel isotopes Tooth enamel isotope data from the site of Ele Bor indicate representation of grazers (>80% C4), mixed feeders (between 80-20% C4), and browsers (>80% C3) consistently across all three horizons (Table 4.3, Figure 4.6). Isotope values for archaeological herbivores are similar to their modern counterparts (Table 4.3). The modern comparative fauna for Ele Bor herbivores is a subset of teeth sampled from Turkana, since no modern faunal collections have ever been made in the area. 4.6 Discussion 4.6.1 Comparative records of ecology in the Lake Victoria Basin Holocene records from the Lake Victoria Basin present an opportunity to understand paleoecology over the course of the Holocene using almost continuous records of pollen and ẟ13Cwax values from piston cores (Figure 4.7). Similarly, these records present a unique opportunity to compare these commonly used but rarely integrated methods of understanding paleoecology (Feakins et al., 2005; Levin, 2015). 92 Herbivore tooth enamel isotopes from archaeological sites are depicted as integrating data across a time slice of this record (Figure 4.7), and the contrasting results of this data compilation raise intriguing questions. As has been previously noted, ẟD records from leaf wax biomarkers, a proxy for rainfall amount (Kahmen et al., 2013; Magill et al., 2013b; Niedermeyer et al., 2010; Polissar and Freeman, 2010; Schefuß et al., 2005), indicate gradual rather than abrupt aridification of the basin, in accordance with insolation forcing (Berke et al., 2012a), a trend also reflected in ẟDwax from Lake Challa (Tierney et al., 2011a) and marine sediments (Niedermeyer et al., 2010). With these caveats in mind, the tooth enamel record may be useful for determining which lacustrine proxies represent terrestrial vegetation within a certain area with more fidelity. Herbivore tooth enamel isotopes are not without their own biases, such as food selection by herbivores and taphonomic (preservation) bias at archaeological sites (Gannes et al., 1997). However, dietary isotopes still provide perspective for what resources were on the landscape, and how herbivores may have changed their diets to accommodate vegetation changes (Uno et al., 2011) (see Chapter 3). Herbivore dietary isotopes across a range of taxa, including those with more flexible diets (hippos and domestic caprines), indicate an overall abundance of C4 plants in the Lake Victoria Basin throughout these timeslices, though some C3 feeders are present in the early Holocene site of Luanda (Figure 4.7). The distribution of modern ẟ13Cenamel from SW Kenya and northern Tanzania (“modern”, Figure 4.3) from the modern counterparts of archaeological fauna indicate that most of these savanna herbivores are consuming C4 foods, with a lower representation of C3 browsers and C3/C4 mixed feeders. Indeed, isotopes of herbivore tooth enamel indicate that the diets of these herbivores have only 93 recently included more C3 foods, but overall are largely C4 feeders. Dietary trends within certain lineages reflect more ancient diets rather than modern counterparts. For example, Tragelaphini and Antilopini have a greater contribution of C4 to their diet throughout the Holocene, only shifting to more C3 after 1.8 kya (later than the Gogo Falls fauna). This one trend is seen in Pleistocene-aged individuals within this lineage (Cerling et al., 2015). Similarly, Neotragini are also much more enriched in 13C relative to modern individuals, which could be explained by all of these samples representing oribi (Ourebia ourebi) rather than dik dik (Madoqua sp.). Hippos from Luanda (9.7 – 8.5 ka) are mixed-feeders, indicating some C3-component to their diet that is not reflected in other herbivores (except Potamochoerus sp.), though returning to >80% C4-feeding in the Gogo Falls samples. This result may indicate a higher proportion of C3 herbaceous vegetation near the lake in the early Holocene (see Chapter 2), or the Luanda are not properly capturing the C3 signal, as implied by the pollen record (Figure 4.7). Reconciling these three distinct proxies is a difficult task. Herbivore tooth enamel isotopes appear to agree more with ẟ13Cwax records than with pollen data (Figure 4.7). This record provides evidence that these two proxies may be similarly spatially integrated. A connection cannot yet be made between tooth enamel isotopes and paleoecology, as such a model does not yet exist, so our discussion is limited to changes in herbivore diets through time. It could also be possible that the leaf wax record is preferentially recording C4 input to the sedimentary record, as is the case with the total soil organic pool (Wynn and Bird, 2007), though herbivore diets are still largely representative of an abundance of C4 resources. It is clear, however, that changing 94 climate and insolation forcing did not have an effect in forcing vegetation, as represented by the leaf waxes, or in herbivore diets, though these changes are synchronous with pollen data. It may be possible that plants in the family Moraceae are more sensitive to changes in total rainfall amount than C4 grasses (Poaceae), or that, despite decreasing rainfall, there was no change in rainfall seasonality, thus maintaining C4 grasses on the landscape (Hély et al., 2006). 4.6.2 Comparative records of ecology in the Lake Turkana and Ele Bor Though the Turkana paleoecology record is discontinuous throughout the Holocene, there are still opportunities to compare proxies throughout certain intervals. The Ele Bor record will also be discussed along with the Turkana record, as they are both from northern Kenya, and present a rare opportunity to compare paleoecology from sites within a lake basin and an inland archaeological site. Though ẟD records do not exist for Turkana, there are direct measures of decreased rainfall via radiocarbon dated paleobeach terraces (Garcin et al., 2012). The abrupt drop in lake level in the Turkana Basin is similar to the same observed increase in dust input to the East Atlantic during the Holocene (Figure 4.8), implying that similar climate forcing mechanisms resulting in aridity are at work in both Turkana and the Horn of Africa. The abrupt drop in lake levels indicates that Turkana responded quickly to insolation forcing and the end of the African Humid Period (deMenocal et al., 2000; Garcin et al., 2012; Tierney and deMenocal, 2013). This precipitous drop in lake level was so severe that Lake Turkana was separated into two distinct lake basins (Cerling, 1996). Tex86 records from Turkana from the 2P (northern basin) and 7P (southern basin) indicate increasing lake water temperatures shortly after the drop in lake level around 5 kya, either from increasing temperatures, 95 lower water level, or both (Berke et al., 2012b). Poaceae pollen data from core 2P indicate generally high (>40%) but fluctuating levels of grass (Mohammed et al., 1995). The Turkana tooth enamel records provide important paleoecological information for this basin. Herbivore tooth enamel isotopes from early Holocene sites all indicate mostly C4 diets (Figure 4.8). At the site of Dongodien (Marshall et al., 1984) – the earliest pastoralist site in eastern Africa – there is a shift in the diets of herbivores towards more C3 feeding compared to early Holocene sites. Relative to modern herbivores of the same taxa, though, dietary isotopes still indicate more C4 biomass (Figure 4.4). These faunal assemblages are not directly comparable, as there are taxa that were not found in older assemblages at this site (Table 4.2), and likewise the presence of domestic stock (Caprini and Bos taurus) at Dongodien that were not present in earlier faunal assemblages. Faunal assemblages from sites within the Lake Victoria Basin, however, contain many of the same taxa at both pastoralist and fisher-hunter-gather sites (Table 4.1). The appearance of previously unseen fauna in early pastoralist sites in Turkana could indicate faunal turnover with vegetation changes within the basin, or simply be a reflection of changes in human behavior and hunting. Regardless, carbon isotopes in tooth enamel tell a story of changing herbivore diets over time, and more representation of C3 throughout the Holocene. The diets of tribes Antilopini and Neotragini are more C4-dominated in the Dongodien assemblage (4.5-3.8 kya) relative to modern specimens. Once again, this is also the case with Pleistocene-ages samples of the same lineage, implying that the dietary change to more C3 happened recently in Turkana. Dongodien caprines (either sheep or goat) likewise consumed more C4 than modern counterparts, and even the most depleted 96 caprines in the assemblage (which were likely goats, see Chapter 5) are not as depleted as modern Turkana goats (see Appendix D). Either early pastoralist goats were consuming less C3 resources than modern goats by choice, or a change in herding behavior led to greater dietary separation due to herd management. The early Holocene sedimentary ẟ13Cwax values add yet another layer of complexity to the Turkana Basin record. These data indicate fluctuating C4 input to the sedimentary record over this 4,000 year period, between 40 and 80%. It is difficult to say whether these biomarker data are in agreement with tooth enamel isotope values due to the small comparative dataset and large temporal integration. During the period over which these proxies overlap, there was a decline in %C4 indicated by the ẟ13Cwax values, shortly before the 8 kyr event, an abrupt arid period (Gasse, 2000; Mayewski et al., 2004). During this event, %C4 increased to ~80%. It is possible that these herbivores (most of which exhibit dietary isotopes of >80% C4, as well), are selectively sampling these periods of higher C4 in Turkana. Another possibility is that decreasing less than 40% C4 representation on the landscape is not enough of a change in terrestrial vegetation to result in a dietary shift in these mammals. Another possibility is that these leaf wax biomarkers have been transported into Turkana from outside the basin – possibly via the Omo River which drains the Ethiopian highlands and delivers up to 80% of Turkana’s water and the majority of its sedimentary load in the northern basin (Cerling, 1986). Ele Bor The Turkana paleoecolgoical record indicates that terrestrial ecology may be sensitive to climatic changes forced by insolation. The comparative tooth enamel isotope record from Ele Bor, 200 km east and 500 m higher in elevation than Lake Turkana, is 97 needed to address whether or not ecological changes occurred in other sites in northern Kenya, particularly outside of a lake basin. Ele Bor was an arcaheolgical habitation site with little supporting evidence for pastoral activity, providing an additional perspective on changing environments and the spread of herding communities (Gifford-Gonzalez, 2003). The fauna from Ele Bor is distinctive from that for Lake Turkana and Lake Victoria, and represents tribes of bovids (e.g., Tragelaphini, Table 4.3) and other herbivores (such as giraffe) that in modern settings are classified as browsers (Cerling et al., 2003; Gagnon and Chew, 2000). Faunal assemblages and isotopic indicators of diet are similar across all three archaeological horizons (Figure 4.5, Table 4.3). Unfortunately, other proxies of paleoecology and/or climate do not exist for Ele Bor and would be difficult to generate due to its remote location. However, the isotopes of Ele Bor herbivores indicate that the environment remained ecologically stable before and after the termination of the African Humid Period at about 5.5 kyr (deMenocal et al., 2000). The trends in other basins for certain taxa that were more C4 dependent on plants than modern comparative samples – Tragelaphini and Antilopini – are not present in the Ele Bor fauna, except in the oldest layer (6.5–6.2 kya). Tragelaphini are mixed feeders, then switch to diets of >80% C3 in the 5.2–3.5 kya faunal layer, and remain that way through modern fauna. Antilopini remain mixed feeders throughout the assemblages, then consume more C3 biomass in modern samples. Of course, this modern fauna is from the Lake Turkana Basin and may not be directly comparable, but it is intrguiging that only at Ele Bor do tragelaphines appear to consume a more modern (more C3) diet much earlier than in either lake basin. The fauna from Ele Bor highlights possible biases of lake basin records when 98 reconstructing ecology over a certain time period. Large African rift lakes may have distinctive climate and environmental systems that are not representative of the region as a whole, and finding more sites outside of these basins (though challenging) would provide a rewarding opportunity to understand paleoenvironments outside of this context (Levin, 2015). 4.7.3 Paleoenvironments and advent of food production in East Africa Within eastern Africa, there are variable records of the timing and social context in which herding first appears. As herding spread (Hildebrand and Grillo, 2012), cattle and caprines appeared by the eastern shore of NW Kenya’s Lake Turkana around 4.5 Kya at GaJi4 (Dongodien) and GaJi2 (Barthelme, 1985; Marshall et al., 1984), along with wild terrestrial and aquatic fauna and novel Nderit pottery. This is accompanied both by a change in climate and a drop in lake level and also a shift in herbivore faunal assemblages with a greater representation of C3 plants in their diets than at previous sites (Figure 4.8). In the Victoria Basin, early livestock is found with distinctive Kansyore pottery, a ceramic tradition whose roots precede herding in the region. Domesticates appear in Kansyore sites (e.g., Wadh Lang’o) ~4.4–3.3 Ka, but the small frequency of domestic fauna before ~1.9–1.8 Ka (Wadh Long’o and Gogo Falls) suggests that people initially supplemented a foraging diet with domesticates during the mid-Holocene but did not practice herding themselves until later (Dale and Ashley, 2010; Prendergast, 2010; Prendergast and Lane, 2010; Robertshaw, 1991). This is not accompanied by a large change in climate as indicated by ẟD or any significant shifts or trends in herbivore diets overall, or any significant change in ẟ13Cwax values. Only gradually increasing ẟD values imply a damped increase in aridity in the region, and gradually decreasing Moraceae 99 pollen and increasing Poaceae pollen indicate changing conditions throughout this time, though slowly (Figure 4.7). Currently, archaeologists have proposed several possible environmental scenarios surrounding the spread of herding in East Africa. (1) Herding spread amidst worsening environmental conditions (i.e., people were “pushed” into moving their herds south to still-moist areas; migration model) or adopted livestock because fishing/hunting/gathering strategies were breaking down as climate became more arid (diffusion model). (2) It spread amidst an economic boom in a favorable climate regime (i.e., people were “pulled” south because herding was so successful it triggered an expansion; migration model). (3) Local populations saw livestock as an attractive supplement to a fairly secure fisher-hunter-gatherer subsistence strategy (diffusion model). The paleoenvironmental research here cannot resolve migration vs. diffusion debates concerning the spread of herding, but it can refine them by clarifying the environmental context in which livestock appeared. These records indicate distinctive environmental changes associated with the archaeological records in each basin. In Turkana, around the time of the appearance of the first herders, there was a fairly abrupt (<1,000 years) change in both climate and either herbivore faunal communities and/or diet with the appearance of the first herders. At Ele Bor, there appears not to have been a distinctive change in herbivore diets over the last 6,000 years, and no evidence for any change in food producing economies. In Lake Victoria, the adoption of herding was a complicated process without a gradual and somewhat diffuse cultural transition between foragers (Kansyore) to herders (Elmenteitan) (Prendergast, 2011; 2010b), and likewise a 100 gradual change in climate and complicated and contrasting picture of ecological change. These environmental data for these basins imply that there is no “one size fits all” model for the spread of herders southward, and that distinctive environments are only associated with the appearance of the first herders only in the Lake Turkana region, but not in the Lake Victoria region. These data call for archaeologists to refine their models concerning the spread of herders southward through East Africa, and that the environmental context of these early pastoralists must be determined on a case-by-case basis. 4.7 Conclusions This comprehensive paleoecological dataset provides a needed comparative record for two Kenyan lake basins which record separate aspects of the spread of early food producers across East Africa during the Holocene – the northern Lake Turkana Basin and the south-western Lake Victoria Basin. The appearance of the first herders in the Lake Turkana Basin appears to coincide with both a climatological (deMenocal et al., 2000) and ecological (increasing C3 in herbivore diets) shift. Evaluating whether this change in conditions was favorable or unfavorable (i.e., push vs. pull scenario of early herders) for the first food producers is difficult, but certainly some environmental change occurred which may have influenced their southward spread into the Turkana Basin. Similarly, no environmental change is evident from tooth enamel isotopes at the archaeological site of Ele Bor before and after the end of the African Humid Period, and likewise, there is no evidence for the presence of early herders at the site. In contrast, paleoecolgoical proxies from the Lake Victoria Basin, with a more diffuse and complex model of the adoption of early food production at archaeological sites, do not indicate any large shifts in ecology or climate. Rather, aridity set in gradually (Berke et al., 101 2012a), with no evidence of major ecological changes from leaf wax records or tooth enamel isotopes. These data are in contrast with pollen records, however, and there are still few paleoecological studies that combine such proxies. These Holocene paleoecolgoical records highlight the contrasting information provided by these proxies and call for further evaluation of the spatio-temporal aspects of ecology that they may sample, and caution researchers against relying on any one proxy to provide a comprehensive reconstruction of ecology. 4.8 Acknowledgements I would like to acknowledge the following people, as this work could never have taken place without their intellectual guidance, mentorship, and assistance in the lab and field: Elisabeth Hildebrand, John Shea, Fiona Marshall, John Harris, Katherine Freeman, Frank Brown, Brett Tipple, Kevin Uno, Amanuel Beyin, Casey Kidney, Glynis Jelhe, Steven Foreman, Francis Ekai, Heather Graham, Clay Magill, Esperanza Zagal, Dan Davis, Scott Blumenthal, Anneke Janzen, Kate Grillo, Sarah Pilliard, Emma Mbua, Purity Kiura, F. Kyalo Manthi, and Louise Leakey. 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Global Change Biology 13(10), 2206–2217. 112 L ak e Tu r kan N NT RA LR IF T a Ele Bor KENYA ive r oR Om A NY ILEMI ET KE UG AN DA Lake Victoria CE Equator Ileret FwJj5 FxJj12,12N GaJj11,12 GaJj1 Lake Turkana GaJi2,4 Area 103 Kisumu Lake Victoria 40 miles Winam Gulf Wadh Lang’o Kanjera Luanda Kanam White Rock Point Gogo Falls HI KE OPI NY A A North Horr Lodwar Lothagam fishing site Loiyangalani 40 miles Figure 4.1: Map of the study areas. Callouts to specific sampling areas in red boxes. 113 Figure 4.2: Holocene paleoenvironment and archeology in Kenya A - Solar insolation (w/m2) at 20 N; B -elevation of paleo lake terraces as a proxy for aridity (Garcin et al., 2012); C – deuterium leaf wax isotopes from sedimentary cores from Lake Victoria as a proxy for aridity (Berke et al., 2012a); D – radiocarbon dates for archaeological sites from the Turkana Basin (blue sites indicate domestic fauna present) (Barthelme, 1985; Garcin et al., 2012); E – radiocarbon dates from archaeological layers at Ele Bor (Gifford-Gonzalez, 2003); F - radiocarbon dates from archaeological sites in the Victoria Basin (Prendergast, 2010; Prendergast and Lane, 2010; Robertshaw, 1991) 114 115 Table 4.1: δ 13C of herbivore tooth enamel from the Lake Victoria Basin. Luanda Family Bovidae Bovidae Suidae Suidae Bovidae Bovidae Tribe Bovini Reduncini Reduncini Reduncini Hippopotamidae Equidae Bovidae Bovidae Bovidae Bovidae Bovidae Wadh Lang'o Family Bovidae Bovidae Suidae Bovidae Bovidae Bovidae Bovidae Bovidae Bovidae Bovidae Bovidae Alcelaphini Species Syncerus caffer Redunca sp. Potamochoerus sp. Phacochoerus sp. Kobus kob Kobus ellipsiprymnus defassa Hippoptamus amphibius Connochaetes taurinus class size 4 class size 3 Bovini Alcelaphini Tribe Reduncini Reduncini Reduncini Caprini Alcelaphini unk. Bovini Alcelaphini Species Redunca redunca Phacochoerus sp. Kobus ellipsiprymnus defassa Connochaetes taurinus class size 2 class size 0 Bos taurus δ 13C +/- 2.1 1.6 -10.5 1.5 2.0 2.4 No. 17 2 3 Citation this work this work this work 0.7 2.0 2.3 0.6 0.9 0.8 2 3 3 this work this work this work -4.2 1.6 6 this work 1.3 1 7 this work this work 6 1 1 2 this work this work this work this work No. 1 3 1 1 Citation this work this work this work this work 0.6 2.2 1.7 1.4 3.2 2.8 1.1 δ 13C +/- 2.5 -0.5 0.4 1.4 0.5 2.9 1.2 2.8 1.5 19 1 this work this work 0.9 -0.3 -0.6 1.8 2.1 1.6 2.8 0.1 6 2 2 1 6 this work this work this work this work this work 1.1 116 Table 4.1 (cont’d) Gogo Falls (from Chritz et al., 2015) Family Tribe Species Bovidae Tragelaphini Taurotragus oryx Bovidae Cephalophini Sylvicapra sp. Bovidae Reduncini Suidae Potamochoerus sp. Suidae Phacochoerus sp. Bovidae Neotragini Ourebia ourebi Bovidae Hippotragini Hippopotamidae H. amphibius Equidae Bovidae Caprini Bovidae unk. Bovidae Bovini Bos taurus Bovidae Alcelaphini δ 13C +/- -6.6 -8.8 0.7 -3.9 2.8 -0.1 0.3 1.2 0.6 0.8 0.3 0.5 1.6 1.5 0.8 0.8 1.4 0.6 0.6 1.4 1.3 1.5 1.2 1.7 No. 5 1 5 1 4 8 5 2 13 14 2 11 15 117 Table 4.1: (cont’d) modern Family Bovidae Tribe Tragelaphini Species Taurotragus oryx Bovidae Cephalophini Sylvicapra sp.* Bovidae Reduncini δ13C1750 +/- -10.4 2.2 No. 18 -10.6 1.4 9 2.5 1.4 51 Suidae Potamochoerus sp. -6.9 4.9 15 Suidae Phacochoerus sp. 1.0 1.1 56 Ourebia ourebi 1.2 4.4 4 1.8 2.1 29 -2.3 1.7 14 2.2 0.4 9 -4.0 3.4 22 2.7 1.2 5 3.2 1.5 93 Bovidae Neotragini* Bovidae Hippotragini Hippopotamidae H. amphibius Equidae Bovidae Caprini Bovidae Bovini Bovidae Alcelaphini Bos taurus *values calculated from keratin using 𝜀keratin-enamel of 11.1‰ (Cerling et al., 2003). Citation (Cerling et al., 2003; Van Der Merwe, 2013) (Cerling et al., 2003) (Cerling et al., 2003; Van Der Merwe, 2013) (Cerling and Harris, 2002) (Bocherens et al., 1996; Cerling and Harris, 2002; Kingston, 2011) (Cerling et al., 2003) (Bocherens et al., 1996; Cerling et al., 2003; Van Der Merwe, 2013) (Cerling and Harris, 2002; Cerling et al., 2011) (Bocherens et al., 1996; Cerling et al., 2011; Kingston, 2011) (Balasse and Ambrose, 2005a) (Balasse and Ambrose, 2005a; Cerling et al., 2003) (Cerling et al., 2003; Kingston, 2011; Van Der Merwe, 2013) 118 Luanda (9.7 - 8.5 Ka) Syncerus caffer (13) Redunca sp. (2) Potamochoerus sp. (3) Phacochoerus aethiopicus (2) Kobus kob (3) Kobus defassa (3) Hippopotamus amphibius (6) Equidae (1) Connochaetes taurinus (7) Bovidae, class 4 (6) Bovidae, class 3 (1) Bovini (1) Antilopini (2) Wadh Lang’o (3.4 - 1.3 Ka) Reduncini (1) Redunca sp. (3) Phacochoerus aethiopicus (1) Kobus defassa (1) Caprinae (18) Connochaetes taurinus (1) Bovidae, class 2 (6) Bovidae, class 0 (2) Bovidae (2) Bos taurus (1) Gogo Falls (1.9 - 1.7 Ka) modern Alcelaphini (6) Tragelaphini (5) Sylvicapra sp. (1) Reduncini (5) Potamochoerus sp. (1) Phacochoerus sp. (4) Neotragini (8) Hippotragini (5) Hippopotamidae (2) Equidae (13) Caprini (14) Bovidae (2) Bos taurus (11) Alcelaphini (15) Tragelaphini (18) Sylvicapra sp. (9) Reduncini (51) Potamochoerus sp. (15) Phacochoerus sp. (56) Neotragini (4) Hippotragini (29) Hippopotamidae (14) Equidae (9) Caprini (44) Alcelaphini (93) -15 -10 -5 0 5 13 δ C (V-PDB) Figure 4.3: Boxplots of δ13C values and dietary designations for herbivores in the Lake Victoria Basin. 119 Table 4.2: δ13C of herbivore tooth enamel from the Lake Turkana Basin. GaJj11, FxJj12, Equid Site Family Tribe Suidae Bovidae Reduncini Hippopotamidae Equidae Bovidae Bovidae Dongodien, GaJj2 Family Suidae Bovidae Bovidae Hippopotamidae Equidae Bovidae Bovidae Bovidae Bovidae Bovidae Hippopotamus amphibius unk Alcelaphini Lothagam Harpoon Site Family Tribe Suidae Hippopotamidae Equidae Cercopithecidae Bovidae Species Phacochoerus sp. Species Phacochoerus sp. Hippopotamus amphibius Papio anubis unk Tribe Species Phacochoerus sp. Neotragini unk. H. amphibius Caprini Bovini Antilopini Alcelaphini Aepycerotini δ 13C +/- 0.15 2.00 0.25 No. 2 1 Citation this work this work -1.02 -0.56 0.93 1.28 1.53 1.08 2.55 0.76 2 7 2 3 this work this work this work this work δ 13C +/- 0.90 1.57 No. 5 Citation this work -0.59 0.61 -6.87 0.50 0.96 0.58 5 4 1 9 this work this work this work this work δ 13C +/- -0.27 -2.23 -1.95 -1.55 -0.24 -4.01 1.62 -3.24 -1.02 -7.16 1.64 4.09 2.61 1.77 No. 5 3 9 4 1 20 1 4 6 1 Citation this work this work this work this work this work this work this work this work this work this work 1.60 2.33 0.98 2.80 120 Table 4.2:(cont’d) Modern (data from Cerling et al., in revision) Family Tribe Species Suidae Phacochoerus sp. Bovidae Neotragini Hippopotamidae H. amphibius Equidae Cercopithecidae Papio anubis Bovidae Caprini Bovidae Bovini Bos taurus Bovidae Antilopini Bovidae Alcelaphini δ13C1750 +/- 0.13 -10.75 -0.34 -2.09 -10.90 -5.69 3.13 -8.96 3.13 2.10 1.78 2.11 3.69 4.31 0.87 4.48 0.98 No. 4 16 8 14 1 14 4 3 6 121 GaJj11, FxJj12, Equid Site (10.5 - 9.1 Ka) >80% C3 >80% C 4 Phacochoerus aethiopicus (2) Reduncini (1) Hippopotamidae (2) Equidae (8) Bovidae (2) Lothagam Harpoon Site (8.0 - 6.5 Ka) Alcelaphini (3) Phacochoerus aethiopicus (5) Hippopotamidae (5) Equidae (4) Papio anubis (4) Dongodien, GaJj2 (4.5 - 3.8 Ka) modern (<50 ya) Bovidae (9) Phacochoerus aethiopicus (5) Neotragini (3) Bovidae (9) Hippopotamidae (4) Equidae (1) Caprini (20) Bovini (1) Antilopini (4) Alcelaphini (6) Aepycerotini (1) Phacochoerus aethiopicus (4) Neotragini (16) Hippopotamidae (8) Equidae (14) Papio anubis (1) Caprini (14) Bos taurus (4) Antilopini (3) Alcelaphini (6) -15 -10 -5 0 5 13 δ C (V-PDB) Figure 4.4: Boxplots of δ13C values and dietary designations for herbivores in the Lake Turkana Basin. 122 Depth Lithology 25 %C4 20 40 60 80100 increasing sand content (regression) 5,825 +/- 272 YBP (mollusc shell in sand) 20 laminated diatomaceous silts with carbonate nodules 15 paleosol beach sands 11,455 +/- 885 YBP (Corbicula africana) 10 5 0m -32 -28 -24 -20 δ13C(C28-FAME ) Figure 4.5: Leaf wax biomarker isotope values for terrestrial lacustrine deposits in Area 103, Turkana (dates recalibrated using OxCal 4.2 using IntCal13 from Owen and Renaut, 1986). 123 Table 4.3: δ13C of herbivore tooth enamel from Ele Bor. Horizon C Family Bovidae Bovidae Bovidae Bovidae Horizon B Family Equidae Bovidae Giraffidae Bovidae Bovidae Bovidae Bovidae Tribe δ13C Antilopini Hippotragini Tragelaphini 1.5 -5.0 2.4 -3.7 Tribe δ13C +/- 0.5 -4.1 -12.4 -11.8 1.5 -2.1 -10.7 0.6 3.5 δ13C +/- -2.8 -10.7 1.3 3.8 2.4 δ13C1750 +/- -10.8 -9.0 -9.8 0.7 -2.4 -11.3 1.8 4.5 0.8 1.7 3.7 Antilopini Neotragini Hippotragini Cephalophini. Tragelaphini Horizon A1/A2 Family Tribe Bovidae Antilopini Bovidae Neotragini Bovidae Hippotragini modern Family Bovidae Bovidae Bovidae Bovidae Equidae Giraffidae Tribe Neotragini Antilopini Tragelaphini Hippotragini +/2.5 5.2 2.8 1.4 No. 1 3 1 3 Citation this work this work this work this work No. 2 13 1 1 8 1 4 Citation this work this work this work this work this work this work this work No. 7 3 1 Citation this work this work this work No. 16 3 3 9 13 1 Citation Cerling et al., in press Cerling et al., in press Cerling et al., in press Cerling et al., in press Cerling et al., in press Cerling et al., in press 124 Horizon C, (6.5 -6.2 Kya) Tragelaphini (3) Hippotragini (1) Bovidae (1) Antilopini (3) Horizon B, (5.2 - 3.5 Kya) Tragelaphini (4) Neotragini (1) Hippotragini (8) Giraffidae (1) Equidae (2) Cephalophini (1) Horizon A1/A2, (1.8 - 0.5 Kya) Antilopini (13) Neotragini (3) Hippotragini (1) Antilopini (7) modern (Turkana, <50 ya) Tragelaphini (3) Neotragini (16) Hippotragini (9) Giraffidae (1) Equidae (13) Antilopini (3) -15 -10 -5 0 5 13 δ C (V-PDB) Figure 4.6: Boxplots of δ13C values and dietary designations for herbivores in the Ele Bor. 125 Figure 4.7: Holocene paleoenvironment at Lake Victoria, Kenya. A – Solar summer insolation at 20 N; B - ẟDwax values from sedimentary n-alkanes in piston cores from Lake Victoria (Berke et al., 2012a); C - ẟ13Cwax values from sedimentary n-alkanes in piston cores from Lake Victoria (Berke et al., 2012a); D – Poaceae (grass) pollen from Pilkington Bay piston cores (Kendall, 1969); E – Moraceae (moist trees) pollen from Pilkington Bay (Kendall, 1969); F – tooth enamel isotopes from modern (see Table 4.1) archaeological sites in the Lake Victoria Basin (this study). 126 127 Figure 4.8: Holocene paleoenvironment at LakeTurkana and Ele Bor, Kenya. A – Solar summer insolation at 20 N; B – East Atlantic terrigneous dust input to sedimentary record from Core 231 in the Gulf of Aden (deMenocal et al., 2000); C – Lake Turkana paleolake terraces (Garcin et al., 2012); D – Poaceae (grass) pollen from core 2P (north basin)(Mohammed et al., 1995); E - ẟ13Cwax values from sedimentary n-alkanoic acids from terrestrial lacustrine sediments (this study); F – Lake Turkana surface temperatures from Tex86 from cores 2P and 7P (Berke et al., 2012b); G – tooth enamel isotopes from modern (see Table 4.2) archaeological sites in the Lake Turkana Basin (this study); H – tooth enamel isotopes from archaeological sites from Ele Bor, Site M (this study). 128 CHAPTER 5 ENVIRONMENTS AND TRYPANOSOMAISIS RISKS FOR EARLY HERDERS IN THE LATER HOLOCENE OF THE LAKE VICTORIA BASIN, KENYA Printed with permission from: Chritz, K.L., F.B. Marshall, M.E. Zagal, F. Kirera, T.E. Cerling. 2015. Environments and trypanosomaisis risks for early herders in the later Holocene of the Lake Victoria basin, Kenya. Proceedings of the National Academy of Sciences 112(12): 3674-3679, doi: 10.1073/pnas.1423953112. 130 Environments and trypanosomiasis risks for early herders in the later Holocene of the Lake Victoria basin, Kenya Kendra L. Chritza,1, Fiona B. Marshallb, M. Esperanza Zagalc,d, Francis Kirerae, and Thure E. Cerlinga,f a Department of Biology, University of Utah, Salt Lake City, UT 84112; bDepartment of Anthropology, Washington University in St. Louis, St. Louis, MO 63130; cDepartment of Chemistry and dDepartment of Anthropology, University of Utah, Salt Lake City, UT 84112; eDepartment of Anatomy, Ross University School of Medicine, Miramar, FL 33027; and fDepartment of Geology and Geophysics, University of Utah, Salt Lake City, UT 84112 Specialized pastoralism developed ∼3 kya among Pastoral Neolithic Elmenteitan herders in eastern Africa. During this time, a mosaic of hunters and herders using diverse economic strategies flourished in southern Kenya. It has been argued that the risk for trypanosomiasis (sleeping sickness), carried by tsetse flies in bushy environments, had a significant influence on pastoral diversification and migration out of eastern Africa toward southern Africa ∼2 kya. Elmenteitan levels at Gogo Falls (ca. 1.9–1.6 kya) preserve a unique faunal record, including wild mammalian herbivores, domestic cattle and caprines, fish, and birds. It has been suggested that a bushy/woodland habitat that harbored tsetse fly constrained production of domestic herds and resulted in subsistence diversification. Stable isotope analysis of herbivore tooth enamel (n = 86) from this site reveals, instead, extensive C4 grazing by both domesticates and the majority of wild herbivores. Integrated with other ecological proxies (pollen and leaf wax biomarkers), these data imply an abundance of C4 grasses in the Lake Victoria basin at this time, and thus little risk for tsetse-related barriers to specialized pastoralism. These data provide empirical evidence for the existence of a grassy corridor through which small groups of herders could have passed to reach southern Africa. | archaeology carbon isotopes livestock disease | food production | East Africa | H erding was the earliest form of food production in Africa, originating in the arid eastern Sahara ∼8 kya. Adoption of food production and increasing mobility in northeastern Africa allowed prehistoric people to manage resource availability amid a drastically changing climate (1) and transformed local populations of people and animals. Saharan herders and hunters spread southward to the Sahel, reaching eastern Africa by 4.5 kya, and eventually reaching southern Africa with sheep and cattle around 2 kya (1–8). The southern African data have been much debated, but recent genetic studies support at least limited movement of early herders from eastern to southern Africa (5, 7). Iron Age Bantu agriculturalist migrations and forager exchange processes also contributed to livestock spread. In contrast to Iron Age farmers, however, the sparse archaeological evidence suggests slow and small-scale southern pastoral migrations (9). Researchers have attributed the limited penetration of southern Africa by stone-using pastoralists to the prevalence of woodland habitats, the distribution of tsetse fly, and the influence of trypanosomiasis on livestock production from Lake Victoria and the Serengeti southward (10, 11). However, site-based paleoenvironmental data have proven difficult to obtain, and although often assumed, the proposition that ancient closed or bushy environments represented a barrier to herding has been seldom examined. Here we present stable isotope analysis of herbivore tooth enamel (n = 86) from Elmenteitan Pastoral Neolithic levels at Gogo Falls (ca. 1.9–1.6 kya) near Lake Victoria, one of the northernmost “woodland tsetse belts” modeled for Africa ca. 2,000 y ago (12, 13). Rather than providing evidence www.pnas.org/cgi/doi/10.1073/pnas.1423953112 for the Lake Victoria basin as a wooded tsetse fly harboring habitat, these new fine-resolution paleoenvironmental data document extensive C4 grazing by both domesticates and the majority of wild herbivores. A synthesis of lacustrine and terrestrial signals further supports the existence of grassy areas in the Lake Victoria basin at 2,000 B.P., indicating a change in ecology to bushy environments more recently than previously thought, and an ecosystem that would not have supported the tsetse fly around Lake Victoria. Early herders in northern Kenya relied on sheep, goats, fish, and diverse wild vertebrates (1, 14). Research in western Kenya, and specifically the Lake Victoria basin (Fig. 1), has revealed southwestward movement of Elmenteitan herders into an area populated by complex ceramic-using, fishing hunter-gatherers and a more varied process of adoption of food production. Domestic sheep and goats appear in low numbers starting around 3.7 kya at Wadh Lang’o, and perhaps also at Gogo Falls and Usenge 3 (15), but not at other sites such as Siror (16), suggesting patchy adoption of herding. Specialized dependence on livestock in Africa is first documented in Elmenteitan sites, which date at the earliest to 3.1 kya at Njoro River Cave, east of Lake Victoria (17, 18). Elmenteitan sites occur to the north in Laikipia, on the Mau Escarpment, on the southern end of the Loita Hills and the northern Mara Plains, and at Gogo Falls (the westernmost extent) (18). Despite opportunities for hunting, fauna from Elmenteitan sites on the Mara plains are >90% domestic (19, 20). The exception to this is the site of Gogo Falls, which has a diverse and abundant faunal assemblage, including Significance Herding was the earliest form of African food production and transformed local populations of people and animals. Herders migrated from eastern to southern Africa around 2,000 years ago, but only in small numbers. Zoonotic disease vectors, specifically the tsetse fly, which carries sleeping sickness, are thought to have impeded these movements. Archaeologists have argued that the presence of tsetse flies around Lake Victoria, Kenya, created a barrier that prevented migration and forced subsistence diversification. This study, using stable isotope analysis of animal teeth, reveals the existence of ancient grassy environments east of Lake Victoria, rather than tsetserich bushy environments. This overturns previous assumptions about environmental constraints on livestock management in a key area for southward movement of early herders. Author contributions: K.L.C. designed research; K.L.C. and M.E.Z. performed research; F.K. provided samples; T.E.C. contributed new reagents/analytic tools; K.L.C., F.B.M., M.E.Z., F.K., and T.E.C. analyzed data; and K.L.C., F.B.M., and T.E.C. wrote the paper. The authors declare no conflict of interest. This article is a PNAS Direct Submission. 1 To whom correspondence should be addressed. Email: [e-mail]. This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10. 1073/pnas.1423953112/-/DCSupplemental. PNAS Early Edition | 1 of 6 ANTHROPOLOGY Edited by Richard G. Klein, Stanford University, Stanford, CA, and approved February 4, 2015 (received for review December 14, 2014) 131 N KENYA Equator of archaeological sites, however, terrestrial paleoecological data for the Holocene in Kenya are scarce and have been based primarily on lacustrine archives, rather than sites themselves. Pollen and leaf wax biomarkers from lacustrine records from Lake Victoria yield somewhat conflicting paleoenvironmental interpretations: pollen data indicate decreasing moist tree and shrub presence (indicated by Moraceae pollen) in the Lake Victoria basin until ∼2 kya, when grass pollen (Poaceae) increased (Fig. 2) (34), contrasting with stable carbon isotopes from lacustrine leaf wax biomarkers, which indicate persistence of C4 grasses throughout the last 6 kya, with only a sharp decrease in C4 around 3 kya (Fig. 2) (35). These contrasting results could be explained by differences in spatial integration, with lipids recording a more localized signal within the catchment and pollen recording a more regional vegetation signal (36), and likely also point to varied, fluctuating input from C4 grasses during this period. Even so, pollen evidence suggests only minor (∼10%) moist forest in the region compared with ∼20–30% input from Kisumu A B 100 miles Fig. 1. Location of Gogo Falls in relation to other Holocene archaeological sites (●) and towns (○) in the Lake Victoria basin. 2 of 6 | www.pnas.org/cgi/doi/10.1073/pnas.1423953112 100 -22 -24 80 -26 60 -28 40 -30 -32 50 C 40 30 20 10 D 60 50 40 30 Moraceae pollen (%) fish and wild herbivores (21). The unique faunal assemblages from Elmenteitan levels at Gogo Falls and from Prolonged Drift (22), which has a mixed Savanna Pastoral Neolithic stone tool assemblage, have sparked considerable debate over the significance of subsistence variability for understanding environmental risks and the dynamics of frontier pastoral-hunter-gather subsistence during the spread of food production. Foragers in the process of adopting herding, loss of stock by pastoralists, and environmental and disease constraints have all been considered possible factors leading to heavy reliance on wild resources (21– 24). South of the Mara plains and Lake Victoria, ancient herders left fewer material traces, leading to arguments regarding the existence of a long-term pastoral-forager frontier, south of which herders did not thrive (25–27). The presence of tsetse flies in eastern Africa has been used as an explanation for observed patterns of subsistence diversity among early pastoralists in the Lake Victoria basin (21, 28) and for limited southward movement (15, 20, 29, 30). A range of zoonoses posed threats for early herders, but trypanosomiasis risks have an especially widespread effect on human communities, even today (10, 12, 13, 30). Mesic, bushy/wooded environments that may have harbored large numbers of tsetse fly (Glossina spp.) would have been poorly suited to extensive cattle stock-keeping. The tsetse fly carries trypanosome parasites that transfer to hosts during blood meals and eventually cause oftenfatal sleeping sickness in ungulates (predominately domesticates) and humans. In modern and precolonial times, herders have controlled tsetse fly numbers by heavy grazing, burning and destruction of woody areas, and managed livestock to avoid areas with abundant tsetse (10, 31, 32). It is not unreasonable to assume, then, that bushy, tsetse-rich environments would have impeded heavy reliance on livestock by early herders and may have even created a boundary beyond which it was difficult for large numbers of herders to settle (20, 33). Despite an abundance -20 Kansyore %C4 plants 20 miles δ13C Gogo Falls Elmenteitan Urewe 0 Gogo Falls, Trench III White Rock Point Poaceae pollen (%) Luanda Wadh Lang’o Kanjera Kanam 20 10 0 2 4 Age (ka) 6 Fig. 2. (A) Holocene pottery traditions present at archaeological sites in the Victoria basin. (B) Leaf wax δ13C and modeled %C4 from lacustrine sedimentary cores in Lake Victoria (35). (C) Moraceae (tropical mesic trees and shrubs) pollen counts from lacustrine sedimentary cores in Lake Victoria (data from ref. 34). (D) Poaceae (grass) pollen counts from lacustrine sedimentary cores in Lake Victoria (data from ref. 34). B, C, and D modified with permission from Elsevier; www.sciencedirect.com/science/journal/02773791. Chritz et al. 132 Results Isotopic data are presented in in Fig. 3 and SI Appendix, Tables S1 and S3. For comparison, a compilation of modern comparable herbivore δ13C1750 values, primarily from southern Kenya and northern Tanzania, are presented in Fig. 4 (SI Appendix, Table S2) (47–53). The average δ13C values indicate a diet with >80% C4 grass for 10 of 13 taxa from Gogo Falls. Potamochoerus sp. (δ13C = −3.86‰; n = 1), Sylvicapra sp. (δ13C = −8.81‰; n = 1), and browsers mixed feeders grazers Bos taurus (11) Alcelaphini (15) Hippotragini (5) Equidae (13) Reduncini (5) Bovidae (2) Neotragini (8) Caprini (14) Phacochoerus sp. (4) Potamochoerus sp. (1) Tragelaphini (5) Sylvicapra sp. (1) -15 -10 -5 0 5 13 δ C (V-PDB) Fig. 3. Box plots of δ13C tooth enamel values from archaeological tooth enamel, Trench III, Gogo Falls. Chritz et al. browsers mixed feeders grazers Bos taurus (5) Alcelaphini (93) Hippotragini (29) Equidae (9) Reduncini (51) Hippopotamidae (14) Bovidae (0) Neotragini (4) Caprini (37) Phacochoerus sp.(56) Potamochoerus sp. (15) Tragelaphini (18) Sylvicapra sp. (9) -15 -10 -5 0 5 13 δ C (V-PDB) Fig. 4. Box plot of δ13C1750 tooth enamel values from modern comparative fauna from Kenya and Tanzania. *Neotragini and Sylvicapra sp. values were calculated to tooth enamel values from keratin, using ekeratin-enamel values of 11.1‰ because of a lack of enamel values in the literature. Tragelaphini (δ13C = −6.6‰; n = 5) are the most 13C-depleted herbivores in the assemblage, but they are more 13C-enriched than their modern counterparts (Fig. 4; SI Appendix, Table S2). Modern hippos average −2.3 ± 1.7‰, whereas hippos at Gogo Falls average 0.6 ± 0.6‰. The presence of both large migratory and small, local bovids provides further evidence for both locally and basinally grassy environments (21). Domestic fauna are pure C4 feeders at Gogo Falls. Bos taurus is within the range of pure C4 diet at both Gogo Falls and in the modern environment. Tribe Caprini (Subfamily Caprinae), which includes both goats and sheep, is again overwhelmingly within the range of a pure C4 diet (1.6 ± 1.4‰). Modern goat δ13C1750 values from locations across Kenya range from −2.6‰ to −12.3‰, whereas modern sheep range from −0.3‰ to −1.5‰ (SI Appendix, Fig. S1 and Table S4) because of the different dietary preferences of sheep and goats. Sheep are primarily grazers, whereas goats are mixed feeders that tend to browse, as their highly efficient digestive physiology confers an advantage for consuming low-quality woody forage (50, 54–57). Criteria for distinguishing domestic sheep and goat teeth have not been considered reliable in Africa (58), and Gogo Falls samples, at the time of identification, were simply identified as sheep or goat. Future studies may distinguish teeth and postcranial bones in African samples (59). If both sheep and goats were present at Gogo Falls, then these isotopic results indicate a much higher consumption of C4 resources by modern goats than is typically observed (50, 56, 60). If the isotope data represent values exclusively from sheep, which is more likely, this level of C4 intake is not unusual, yet the presence of sheep only at this site is unusual compared with current herding practices in East Africa. Discussion In comparison with the present day, isotopic analyses of herbivores indicate that during the Elmenteitan occupation at Gogo Falls, there were few C3 browsers or mixed C3–C4 feeders, suggesting a landscape dominated by C4 grasses. This interpretation is in general agreement with δ13Cwax values (Fig. 2) from lacustrine cores in Lake Victoria (which likely indicate vegetation locally, within the lake catchment), and to a lesser extent with pollen records, which indicate mixed environments regionally. These three lines of evidence, taken together, strongly imply an abundance of C4 grasses in the basin during this period. Vegetation in PNAS Early Edition | 3 of 6 ANTHROPOLOGY grasses. These records offer the important wider ecological context of regional (pollen) and basinal (leaf waxes) ecology, yet they are too coarse to understand specialized pastoralist expansions, and especially for local evidence (site-based reconstruction) of habitats harboring tsetse. To determine whether or not shifting tsetse-rich environments were present at particular locales during specific periods, direct stable isotope-based paleoecological analysis at archaeological sites can provide a complement to faunal analysis (21, 37–39). Isotopic measurements of enamel carbon (δ13C) and oxygen (δ18O) can be used to understand diet, habitat, and climate (40, 41). Although bone and tooth collagen have been previously used to assess paleoenvironment at Holocene archaeological sites in Kenya (42), tooth enamel is preferred, as it resists digenetic alteration and can be easily compared with fauna from older periods of geologic time (43). In some instances, faunal tooth enamel isotopes may provide our best estimates of environmental change over time, because these signals are less temporally and spatially attenuated than other proxies, such as leaf wax biomarkers (44), which may be influenced by reservoir effects during catchment transport before deposition (45, 46). The environmental context of the Elmenteitan layers of Gogo Falls has been previously interpreted as a grassland/bushland mosaic, based on analysis of wild species diversity and the presence of modern browsing taxa such as roan or sable antelope (Tragelaphini) and bushpig (Potamochoerus), as well as grazers such as oribi (21). The relatively low proportions of domestic stock were interpreted as reflecting environmental constraints on pastoral productivity in the region. The presence of the tsetse fly in a presumed seasonally mesic, bushy/woody environment, it was argued, prevented herders from relying exclusively on domesticates, which resulted in seasonal fishing and hunting of large wild ungulates (21). These interpretations have not yet been ground-truthed with a local, terrestrial paleoecological indicator. 133 and around the Winam Gulf today has been characterized as “evergreen and semi-evergreen bushland and thicket,” “edaphic wooded grasslands and grasslands on drainage impeded or seasonally flooded soils,” and “moist combretum wooded grassland” (which comprises the ecology area around the Kuja River, where the Gogo Falls is located) (61). Faunally based interpretations, primarily the presence of grazers and modern browsers such as Tragelaphini and Potamochoerus sp., of the ancient environment at Gogo Falls suggested a woodland/grassland/bushland mosaic that is climatologically and ecologically similar to today (21), which would have supported tsetse flies. This bushland interpretation is not supported by the isotopic data, which are dominated by C4-grazing taxa with few C3-browsing taxa, contradicting long-held assumptions of modern climate and ecology persisting over the last 3 millennia in most regions of eastern Africa (19, 21, 24). Given the resolution of data available, it is not yet possible to quantify the relative proportions of grassy and woody vegetation; however, abundant C4 grasses must have been present. Abundant grassy vegetation may have been the result of changes in rainfall seasonality in the later Holocene (62, 63), changes in total rainfall in the Lake Victoria basin, increased burning, or ecological factors, such as heavy grazing by domesticates or wild herbivores. Controlled grazing by cattle, sheep, and goats can stimulate new grass growth where grazing herbivores thrive (64, 65). Overgrazing, however, makes environments more bushy and reduces grass height (66). Burning promotes new grass and may have maintained grasslands if herders were regularly burning savanna for livestock, as is a common practice in modern eastern Africa (67–69). The presence of megaherbivores maintains grasslands as well, and when both burning and elephant or giraffe populations are high, grasslands expand (67, 68, 70). Given such complex environmental interactions in African grasslands today, the question arises whether ancient herders in the Lake Victoria basin helped create grasslands through grazing and fire, or whether they were created and maintained by other ecological and climatological factors (71, 72). The isotopic data from Gogo Falls and leaf wax data from the Lake Victoria basin reveal an ideal environment for large grazing herds of domesticates in the immediate Lake Victoria basin catchment, one rich in C4 grasses. Such grasslands were not an ideal environment for tsetse flies, which calls into question the maintenance of long-term areas of hunter-gatherer-pastoral interaction and the inhibition of southward movements of herders by stable and extensive bands of tsetse bush (2). This study fits with the findings of recent genetic research including Y-chromosome, lactase persistence, and livestock data that point to some migration of herders from eastern to southern Africa (5– 7). Archaeologists arguing for successful early pastoral migrations from eastern to southern Africa have pointed to some similarities between Elmenteitan pottery and early herder ceramics in southern Africa (73). Our data imply that a tsetserich barrier preventing herders from moving through the Lake Victoria basin into northern Tanzania and other parts of subSaharan Africa (9, 24, 33, 74) was not widespread 2,000 y ago, bolstering the possibility of such connections. On a local level, our data show that Elmenteitan herders at Gogo Falls were not forced into hunting and fishing because of ecological constraints on stock-keeping. The Elmenteitan mammalian fauna at Gogo Falls are made up of 53.5% domestic livestock comprising cattle and sheep (based on isotopes). Fishing was also a significant activity (21). Recent research in the Lake Victoria basin has shown that the nearby site of Wadh Lang’o shares similar dates of 1,950 ± 35 y B.P. for early Elmenteitan levels (27, 75) and a number of similarities with Gogo Falls: Wadh Lang’o is also a large open-air site with a long sequence containing Kansyore, Elmenteitan, and Iron Age horizons (27, 75). It is situated in a similar environmental setting, on the banks of a river flowing into Lake Victoria. There are as yet no direct 4 of 6 | www.pnas.org/cgi/doi/10.1073/pnas.1423953112 paleoecological proxies from this site, but according to lacustrine paleoenvironmental archives (Fig. 2), it was also likely situated in a grassy environment. Fishing is a significant activity at both sites; however, the mammals in the Elmenteitan faunal horizons in Trench 1, Wadh Lang’o (below datum 140–190 cm), are predominantly (89–91%) domestic, suggesting limited hunting (75). Alcelaphines are represented, but the zebra and oribi present at Gogo Falls are absent at Wadh Lang’o. Herding strategies also differed between Wadh Lang’o and Gogo Falls, with cattle making up 1% or less of the domestic fauna at Wadh Lang’o (n = 88 below datum 140–190 cm; n = 156 below datum 140–160 cm; SI Appendix, Table S5) compared with 32% (n = 328) at Gogo Falls. In strong contrast with these sites in the Lake Victoria basin, there is no fishing and very little hunting in Elmenteitan sites on the plains of southern Kenya (19, 20). Cattle, sheep, and goats make up 97–100% of the fauna at the open air settlement sites in the Lemek-Mara, including Ngamuriak, Sugenya, and Oldorotua, as well as the Central Rift Valley rock shelter of Maasai Gorge (SI Appendix, Table S5). Reliance on cattle and some small stock, combined with fishing and significant hunting at Gogo Falls, is an unexpected contrast with nearby Elmenteitan sites and does not fit a typical Elmenteitan subsistence pattern. Nor do the fauna, material culture, and organization of this site suggest hunter-gatherers in the process of adopting herding. Gogo Falls appears to have been populated by a unique group of people, potentially a small group of herders, who took flexible approaches to subsistence opportunities and interactions with hunter-gatherer-fishers as they followed grassy corridors in eastern Africa. Conclusions Our results provide much-needed late Holocene paleoenvironmental data for the Lake Victoria basin and demonstrate that substantial ecological changes occurred over the last 2,000 y. Isotopic data reveal that the fauna were dominated by C4 grazers with very few C3 browsing taxa, suggesting a grassy paleoecological setting for the unique Elmenteitan site of Gogo Falls. This challenges interpretations based on macrofaunal remains and models of disease-depressed livestock production (21). The isotopic findings also draw attention to herder relations with hunter-gatherers, rather than ecology, in consumption of large wild ungulates in western Kenya. This suggests that social factors may have played a greater role than previously thought in subsistence diversity during the spread of pastoralism in Eastern Africa. High levels of hunting at Gogo Falls can no longer be attributed to closed or bushy habitats that inhibited livestock production and southward movement of herders (21) and contributed to long-term frontier interactions between pastoralists and hunter-gatherers (25, 27, 74). These data raise interesting questions about the interplay among herders, environmental management and adaptation, and ecological drivers for grassland maintenance. Finally, in light of the isotope data from Gogo Falls and lacustrine leaf wax data (35), the Lake Victoria basin may have been the setting for grassy corridors through which demic migration of pastoralists occurred out of eastern Africa and toward southern Africa (30). Materials and Methods Site Description. The archaeological site of Gogo Falls is located on the banks of the Kuja River by a dam that bears the same name (34°21′E, 0°34′S; Fig. 1). The area receives ∼1,060 mm/y in rainfall, with bimodal rainfall seasonality, with rains occurring in March–May and October–November (28). At this time, the area surrounding the site has been ecologically altered by farming, but natural vegetation is characterized as a wooded grassland (61). Nearby local wild fauna include roan antelope, zebra, and giraffe, and historically, a diverse mammalian population including elephant, lion, cheetah, and rhinoceros (61). Browsers are rare, and thus woodland and thicket encroachment has been occurring in the area (76). Gogo Falls was excavated by Robertshaw in 1980 and 1983 (28, 77), and teeth analyzed for this article were uncovered during the 1983 excavation. Chritz et al. Five noncontinuous trenches were dug in the study area, each yielding abundant faunal remains, artifacts, and potsherds spanning three cultural traditions: Kansyore, Elmenteitan, and Urewe (28). The stratigraphic details of Trench III is presented by Robertshaw (28), but the highlight of this excavation is a 1.5-m ash and dung midden layer containing Elmenteitan pottery exclusively, which Robertshaw interpreted as a stock-keeping area. Radiocarbon dating of charcoal near the top of Trench III gave an uncalibrated age of 1,770 ± 80 y B.P., and another date from the bottom of the trench gave an age of 1,990 ± 80 y B.P. (28), with no evidence of stratigraphic mixing. These dates calibrate to 1,646 calibrated y B.P. (calBP) ± 96 and 1,900 calBP ± 100 at 95.4% confidence using ShCal13 (78). Teeth were selected from faunal analysis by Marshall and Stewart (21). All the faunal material excavated from the 1.5-m ash midden is in excellent preservation. The fauna from this trench are diverse, containing fish, avian, and mammalian remains. Domestic caprines (goat and sheep) constitute the largest proportion of the assemblage (36.4%), followed by cattle (17.1%), topi/hartebeest (presented here at the tribal level of identification, “Alcelaphini”; 15.2%), oribi (presented as “Neotragini”; 11.8%), zebra (11.3%), reedbuck (“Reduncini”; 2.8%), warthog (1.8%), and eland and roan/sable (“T. oryx” and “Hippotragini”, respectively; 1.1%) (21). Stable isotope analysis was carried out on a subset of these teeth, including samples from rarer fauna as well (i.e., Potamochoerus sp. and Sylvicapra sp.). silver capsules and dried under vacuum at 200 °C for 2 h before analysis. Samples were analyzed for δ13C and δ18O on a Finnigan MAT 252 coupled to a Carboflo dual-inlet carbonate device (common phosphoric acid bath at 90 °C for 15 min reaction time). Stable isotope ratios are reported as delta (δ) values relative to the international carbon isotope standard, Vienna Pee Dee Belemnite, using the standard permil (‰) notation, where δ13C = (Rsample/ Rstandard − 1) × 1,000 and Rsample and Rstandard are the 13C/12C ratios of the sample and standard, respectively. SD of an internal carbonate standard (Carrara marble) was ±0.1‰. Dietary designations for herbivores are based on estimated consumption of C4 plants (tropical grasses) and C3 plants (trees, shrubs, herbs), which is calculated using a hypothetical 100% C4 diet and an isotope enrichment (e*) between diet and herbivore tooth enamel of 14.1‰ (81). Estimated carbon isotope composition of tooth enamel for a 100% C4 diet is based on the average isotopic value of modern C4 plants in eastern and central Africa (−12.9‰; n = 764) (82). Changes in the isotopic composition of atmospheric CO2 resulting from the combustion of fossil fuels between modern environments and preindustrial Holocene (presented here as δ 13C1750) were accounted for by adding 1.6‰ to the modern average δ13C of C4 plants (81, 83, 84). Laboratory Analysis. Following the sampling procedures outlined by the National Museums of Kenya, teeth were photographed and sampled along broken edges, using a Dremel tool and diamond drill bit. About 1 mg enamel powder was drilled from each sample. Enamel powder was prepared and analyzed at the University of Utah. Enamel powders were treated with 0.1 M buffered acetic acid (pH ∼5.3) for 30 min to remove labile carbonates (79, 80). After acid treatment, powders were rinsed three times with distilled water and dried at 60 °C overnight. Sample powders were weighed into ACKNOWLEDGMENTS. We thank P. Robertshaw for permission to conduct stable isotope analysis on excavated material and S. 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Cerling TE, Harris JM (1999) Carbon isotope fractionation between diet and bioapatite in ungulate mammals and implications for ecological and paleoecological studies. Oecologia 120(3):347–363. 82. Cerling TE (2014) 14.12 Stable Isotope Evidence for Hominin Environments in Africa. Treatise on Geochemistry, ed Cerling TE (Elsevier, Amsterdam), 2nd Ed, pp 157–167. 83. Francey RJ, et al. (1999) A 1000-year high precision record of δ13C in atmospheric CO2. Tellus B Chem Phys Meterol 51(2):170–193. 84. Keeling CD, Piper S, Bollenbacher AF, Walker SJ (2010) Monthly atmospheric 13C/12C isotopic ratios for 10 SIO stations. Available at cdiac.ornl.gov/trends/co2/iso-sio/iso-sio. html. Accessed October 1, 2014. Chritz et al. 136 SI Appendix Table S1: 13C tooth enamel values (average and standard deviation) from archaeological tooth enamel, Trench III, Gogo Falls. no. 13 samples Family Tribe Species C Std. dev. Bovidae Bovini Bos taurus 1.6 1.5 11 Bovidae Alcelaphini 1.5 1.2 15 Bovidae Hippotragini 1.2 1.4 5 Equidae Equus sp. 0.8 0.6 13 Bovidae Reduncini 0.7 1.7 5 Hippopotamidae Hippopotamus 0.6 0.6 2 amphibius Bovidae unk. 0.5 1.3 2 Bovidae Neotragini Ourebia ourebi 0.3 0.8 8 Bovidae Caprini 0.2 1.4 14 Suidae Phacochoerus sp. -0.1 0.9 4 Suidae Potamochoerus sp. -3.9 1 Bovidae Tragelaphini Taurotragus oryx -6.6 2.8 5 Bovidae Cephalophini Sylvicapra sp. -8.8 1 137 Table S2: 13C1750 tooth enamel values (average and standard deviation) from modern comparative fauna from Kenya and Tanzania. *Neotragini and Sylvicapra values were calculated to tooth enamel values from keratin using keratin-enamel values of 11.1‰ because of a lack of enamel values in the literature. std. dev. no. 13 (±) samples Family Tribe Species C Ref. Bovidae Bovini Bos taurus 2.7 1.2 5 (1, 2) Equus sp. 3.2 1.8 2.2 2.5 -2.3 1.5 2.1 0.4 1.4 1.7 93 29 9 51 14 (3-6) (3-5) (5, 7, 8) (3, 4) (7, 9) 1.3 -4.0 Phacochoerus sp. 1.0 Potamochoerus sp. -6.9 Tragelaphini T. oryx -10.4 Cephalophini Sylvicapra sp.* -10.6 4.4 3.4 1.1 4.9 2.2 1.4 4 22 56 15 18 9 (3) (10) (5, 6, 9) (9) (3, 4) (3) Bovidae Alcelaphini Bovidae Hippotragini Equidae Bovidae Reduncini Hippopotamidae Bovidae Bovidae Suidae Suidae Bovidae Bovidae Neotragini* Caprini Hippopotamus amphibius Ourebia ourebi 138 Table S3: 13C and 18O tooth enamel values from Gogo Falls fauna. Calculated %C4 has been rounded to the nearest 5% to account for uncertainty in the dietary model. 13 18 Common/ Family/SubTribe Specimen ID C O %C4 Species name Family KKC13-E-296 Bos taurus Bovidae Bovini -1.0 1.0 80 KKC13-E-297 Bos taurus Bovidae Bovini -0.8 -2.7 80 KKC13-E-298 Bos taurus Bovidae Bovini 1.9 1.8 100 KKC13-E-299 Bos taurus Bovidae Bovini 1.5 1.2 100 KKC13-E-300 Bos taurus Bovidae Bovini 1.8 0.3 100 KKC13-E-301 Bos taurus Bovidae Bovini 2.9 1.7 100 KKC13-E-302 Bos taurus Bovidae Bovini 2.7 2.8 100 KKC13-E-303 Bos taurus Bovidae Bovini 0.6 0.5 90 KKC13-E-304 Bos taurus Bovidae Bovini 3.4 2.3 100 KKC13-E-305 Bos taurus Bovidae Bovini 3.2 2.1 100 KKC13-E-306 Bos taurus Bovidae Bovini 1.3 2.5 95 KKC13-E-307 Bos taurus Bovidae Bovini 1.8 0.7 100 KKC13-E-342 Bovidae Alcelaphini 0.2 1.4 90 KKC13-E-343 Bovidae Alcelaphini 0.1 1.5 90 KKC13-E-345 Bovidae Alcelaphini 3.2 2.4 100 KKC13-E-350 Bovidae Alcelaphini -0.4 -0.4 85 KKC13-E-351 Bovidae Alcelaphini 1.3 0.3 95 KKC13-E-352 Bovidae Alcelaphini 3.0 1.9 100 KKC13-E-353 Bovidae Alcelaphini 0.6 0.1 90 KKC13-E-354 Damaliscus spp. Bovidae Alcelaphini 2.7 1.5 100 KKC13-E-355 Damaliscus spp. Bovidae Alcelaphini 3.0 1.9 100 KKC13-E-356 topi/hartebeest Bovidae Alcelaphini 2.1 -2.1 100 KKC13-E-357 Damaliscus spp. Bovidae Alcelaphini 0.6 -0.3 90 KKC13-E-358 Damaliscus spp. Bovidae Alcelaphini 2.2 1.3 100 Alcelaphus/ Damaliscus KKC13-E-346* Bovidae Alcelaphini 0.0 -0.3 85 KKC13-E-348* Bovidae Alcelaphini 2.2 1.3 100 KKC13-E-377 topi/hartebeest Bovidae Alcelaphini 0.3 1.1 90 KKC13-E-349 roan/sable Bovidae Hippotragini -0.6 -2.7 85 KKC13-E-373 roan/sable Bovidae Hippotragini 1.4 -2.1 100 KKC13-E-376 roan/sable Bovidae Hippotragini 2.3 0.4 100 KKC13-E-372* roan Bovidae Hippotragini 2.7 0.2 100 KKC13-E-374* roan/sable Bovidae Hippotragini 0.9 2.4 95 KKC13-E-317 Equus burchelli Equidae 1.5 1.9 100 KKC13-E-318 Equus burchelli Equidae 2.0 1.4 100 KKC13-E-319 Equus burchelli Equidae 1.3 0.0 100 KKC13-E-320 Equus burchelli Equidae 0.8 1.3 95 139 Specimen ID KKC13-E-315 KKC13-E-316 KKC13-E-321 KKC13-E-323 KKC13-E-324 KKC13-E-312* KKC13-E-314* KKC13-E-367 KKC13-E-378 KKC13-E-368* KKC13-E-370* KKC13-E-371* KKC13-E-330 KKC13-E-331 KKC13-E-309* KKC13-E-347 KKC13-E-333 KKC13-E-334 KKC13-E-335 KKC13-E-336 KKC13-E-337 KKC13-E-338 KKC13-E-339 KKC13-E-340 KKC13-E-283* KKC13-E-280 KKC13-E-281 KKC13-E-282 KKC13-E-284 KKC13-E-285 KKC13-E-288 KKC13-E-289 KKC13-E-290 KKC13-E-291 KKC13-E-292 KKC13-E-293 KKC13-E-286* KKC13-E-287* KKC13-E-326 KKC13-E-327 KKC13-E-328 Common/ Species name Equus spp. Equus spp. Equus spp. Equus spp. Equus spp. Equus spp. Equus spp. kob/waterbuck kob/waterbuck H. amphibius H. amphibius unk unk Ourebia ourebi Ourebia ourebi Ourebia ourebi Ourebia ourebi Ourebia ourebi Ourebia ourebi Ourebia ourebi Ourebia ourebi goat/sheep goat/sheep goat/sheep goat/sheep goat/sheep goat/sheep goat/sheep goat/sheep goat/sheep goat/sheep goat/sheep goat/sheep goat/sheep goat/sheep Phacochoerus sp. Phacochoerus sp. Phacochoerus sp. Family/SubFamily Equidae Equidae Equidae Equidae Equidae Equidae Equidae Bovidae Bovidae Bovidae Bovidae Bovidae Hippopotamidae Hippopotamidae Bovidae Bovidae Bovidae Bovidae Bovidae Bovidae Bovidae Bovidae Bovidae Bovidae Caprinae Caprinae Caprinae Caprinae Caprinae Caprinae Caprinae Caprinae Caprinae Caprinae Caprinae Caprinae Caprinae Caprinae Suidae Suidae Suidae Tribe Reduncini Reduncini Reduncini Reduncini Reduncini Neotragini Neotragini Neotragini Neotragini Neotragini Neotragini Neotragini Neotragini 13 C 0.1 1.6 0.6 0.5 0.0 0.1 0.0 2.8 2.3 -0.4 -1.0 1.0 0.2 -0.5 1.4 1.0 0.0 0.9 -0.6 -0.9 -0.3 1.2 0.9 0.9 -0.4 2.2 0.2 -2.0 -0.8 0.5 1.7 -0.7 0.5 -2.1 2.6 0.7 0.3 -1.2 0.7 0.4 -0.3 18 O 1.0 2.4 1.2 -0.4 -0.3 1.4 0.6 0.9 1.3 -0.3 -0.2 -2.6 -2.6 0.7 0.8 0.7 -1.1 -3.0 0.9 2.1 1.9 0.5 0.8 0.7 1.9 3.0 1.9 1.6 3.2 4.3 1.3 -2.8 3.8 -0.6 2.9 1.6 3.3 -1.9 1.4 1.7 -0.2 %C4 90 100 90 90 90 90 90 100 100 85 80 95 90 85 100 95 95 94 85 80 85 95 95 95 85 100 90 75 80 90 100 80 90 70 100 90 90 80 90 90 85 140 Specimen ID KKC13-E-332 KKC13-E-360 KKC13-E-363 KKC13-E-364 KKC13-E-361* KKC13-E-362* Common/ Species name Potamochoerus sp. Taurotragus oryx Taurotragus oryx Taurotragus oryx Taurotragus oryx Sylvicapra sp. Family/SubFamily Suidae Tribe 13 C -5.8 18 O 2.3 %C4 45 Bovidae Bovidae Bovidae Bovidae Bovidae Tragelaphini Tragelaphini Tragelaphini Tragelaphini Cephalophini -9.6 -3.9 -4.1 -9.6 -8.8 -0.8 0.0 1.2 2.0 -1.1 20 60 60 20 25 *sample was run untreated due to low sample volume 141 Table S4: Modern caprine Common/Species name sheep sheep sheep sheep sheep sheep sheep sheep goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat goat 13 C1750 tooth enamel data. Locality Naivasha Basin Naivasha Basin Naivasha Basin Naivasha Basin Naivasha Basin Naivasha Basin Central Rift valley Central Rift valley Baragoi Baringo Borana Maua Ciakariga Emuhaya Kainauk Kibwezi Kilungu Kilungu Kimende Lamu Lodwar Loyangalani Maimanti Malaral Mandera Marimanti Matiiri Maua Mericho Nanyuki Nariokotome Ndaragwa South Horr Suguta Malmar Tharaka Tunyai Naivasha Basin Naivasha Basin Naivasha Basin Naivasha Basin Naivasha Basin 13 C1750 0.1 0.3 -0.9 0.2 -0.6 -1.5 -0.7 -0.4 -7.9 -10.2 -10.7 -10.9 -10 -8.5 -6.3 -9.7 -10.1 -4.7 -12.3 -3.5 -9.5 -9.7 -4.1 -5.6 -11.7 -6 -9.5 -3.6 -6.3 -5.4 -3.5 -10.3 -4.4 -11.2 -11.6 -6.3 -7.4 -7.2 -6.1 -6.5 Citation (10) (10) (10) (10) (10) (10) (2) (2) this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study (10) (10) (10) (10) (10) 142 Common/Species name goat goat goat goat Locality Naivasha Basin Naivasha Basin Naivasha Basin Naivasha Basin 13 C1750 -5.9 -3.4 -2.6 -8.5 Citation (10) (10) (10) (10) 143 Table S5: Comparison of wild vs. domestic fauna at Neolithic sites in Kenya Site Tradition wild % domestic N (NISP) Narosura SPN* 7 93 1215 Crescent Island Main SPN 18 82 526 Prolonged Drift SPN 79.9 20.1 1491 Ngamuriak Elmenteitan 0.5 99.56 4653 Sugenya Elmenteitan 1.9 98.1 1774 Oldorotua Elmenteitan 1.6 98.4 2127 Maasai Gorge Elmenteitan 4.3 96.7 115 Gogo Falls Elmenteitan 46.5 53.5 612 Wadh Lango 140-160 Elmenteitan 10.9 89.1 156 Wadh Lango 160-190 Elmenteitan 10 90 88 *SPN = Savanna Pastoral Neolithic Ref. (11) (11) (12) (13, 14) (15) (15) (11) (16) (17) (17) 144 browsers mixed feeders grazers Gogo Falls caprines (14) Ovis aries (8) Capra hircus (37) -15 -10 13 -5 0 5 δ C (V-PDB) Figure S1: Boxplots of 13C1750 tooth enamel values of modern goat, Capra hircus (Table S4), modern sheep, Ovis aries (Table S4), and Gogo Falls caprines (Table S3). 145 SI References 1. Cerling TE, Harris JM, Leakey M, Mudida N (2003) Stable isotope ecology of northern Kenya with emphasis on the Turkana Basin. Lothagam: The Dawn of Humanity in Eastern Africa, eds Leakey MG, Harris JM (Columbia University Press, New York), pp.583–603. 2. Balasse M, Ambrose SH (2005) Mobilité altitudinale des pasteurs néolithiques dans la vallée du Rift (Kenya) : premiers indices de l’analyse du 13C de l’émail dentaire du cheptel domestique. Anthropozoologica 40(1):147–166. 3. Cerling TE, Harris JM, Passey BH (2003) Diets of East African Bovidae based on stable isotope analysis. Journal of Mammalogy 84(2):456–470. 4. Van Der Merwe NJ (2013) Isotopic ecology of fossil fauna from Olduvai Gorge at ca 1.8 Ma, compared with modern fauna. South African Journal of Science 109(11/12):1-14. 5. Bocherens H, Koch PL, Mariotti A, Geraads D, Jaeger JJ (1996) Isotopic biogeochemistry (13C, 18O) of mammalian enamel from African Pleistocene hominid sites. PALAIOS 11(4):306–318. 6. Kingston JD (2011), Stable isotopic analyses of Laetoli herbivores, Paleontology and Geology of Laetoli: Human Evolution in Context. Volume 1: Geology, Geochronology, Paleoecology and Paleoenvironment, ed Harrison T (Springer Netherlands, Dordrecht), pp 293–328. 7. Cerling TE et al. (2011) Diet of Paranthropus boisei in the Early Pleistocene of East Africa. Proceedings of the National Academy of Science 108(23):9337-9341. 8. Kingston JD, Harrison T (2007) Isotopic dietary reconstructions of Pliocene herbivores at Laetoli: implications for early hominin paleoecology. Palaeogeography, Palaeoclimatology, Palaeoecology 243(3-4):272–306. 9. Cerling TE, Harris JM (2002) Dietary adaptations of extant and Neogene African suids. Journal of Zoology 256(1):45–54. 10. Balasse M, Ambrose SH (2005) Distinguishing sheep and goats using dental morphology and stable carbon isotopes in C4 grassland environments. Journal of Archaeological Science 32(5):691-702. 11. Gifford-Gonzalez DP (1998) Early pastoralists in East Africa: Ecological and social dimensions. Journal of Anthropological Archaeology 17(2):166–200. 12. Gifford-Gonzalez DP, Isaac GL, Nelson CM (1980) Evidence for predation and pastoralism at Prolonged Drift: a Pastoral Neolithic site in Kenya. Azania 15(1):57108. 13. Marshall F (1990) Origins of specialized pastoral production in East Africa. 146 American Anthropologist 92(4):873–894. 14. Marshall F, Grillo K, Arco L (2011) Prehistoric pastoralists and social responses to climatic risk in East Africa. Sustainable Lifeways: Cultural Persistence in an Everchanging Environment, eds. Miller NF, Moore KM, Ryan K(University of Pennsylvania Press, Philadelphia), pp 74-105. 15. Simons A (2004) The development of early pastoral societies in south-western Kenya: A study of the faunal assemblages from Sugenya and Oldorotua 1 (La Trobe Univeristy, Melbourne). Ph.D. dissertation. 16. Marshall F, Stewart K (1994) Hunting, fishing and herding pastoralists of western Kenya: the fauna from Gogo Falls. Archaeozoologia 7(1):7-27. 17. Prendergast ME (2010) Kansyore fisher-foragers and transitions to food production in East Africa: The view from Wadh Lang'o, Nyanza Province, Western Kenya. Azania 45(1):83–111. APPENDIX A RAW HIPPO CANINE ISOTOPE DATA AND FIGURES 148 Aberdares NP δ18O δ13C 0 −1 −1 −2 −2 −3 −3 −4 −4 −5 −5 0 50 100 150 200 mm from proximal end 250 δ18Oenamel δ13Cenamel 0 300 Figure A.1: δ13C and δ18O (V-PDB) canine enamel profile plot of Aberdares NP hippo (0mm = death). 149 Amboseli #1 (Amboseli H1/H2) 0 0 −2 −2 −4 −4 −6 −6 −8 −8 0 50 100 150 200 250 300 350 400 δ18Oenamel δ13Cenamel δ18O δ13C 450 mm from proximal end Figure A.2: δ13C and δ18O (V-PDB) canine enamel profile plot of Amboseli #1 hippo (0mm = death). 150 Amboseli #2 (KWS-AMB-0401) −2 δ18O δ13C −1 −3 −2 −4 −3 −5 −4 −6 0 50 100 150 200 250 300 350 δ18Oenamel δ13Cenamel 0 400 mm from proximal end Figure A.3: δ13C and δ18O (V-PDB) canine enamel profile plot of Amboseli #2 hippo (0mm = death). 151 Arabuko Sokoke #1 (AS168) 2 2 0 0 −2 −2 −4 −4 −6 −6 0 50 100 150 200 250 300 δ18Oenamel δ13Cenamel δ18O δ13C 350 mm from proximal end Figure A.4: δ13C and δ18O (V-PDB) canine enamel profile plot of Arabuko Sokoke #1 hippo (0mm = death). 152 Arabuko Sokoke #2 (K00-AS-167) 0 δ18O δ13C −1 −1 −2 −2 −3 −3 −4 −4 −5 −5 50 100 150 200 250 300 350 δ18Oenamel δ13Cenamel 0 400 mm from proximal end Figure A.5: δ13C and δ18O (V-PDB) canine enamel profile plot of Arabuko Sokoke #2 hippo (0mm = death). 153 Arabuko Sokoke #3 (Arabuko Sokoke - 2008) 0 δ18O δ13C −1 −1 −2 −2 −3 −3 −4 −4 −5 −5 −6 −6 0 50 100 150 200 250 δ18Oenamel δ13Cenamel 0 300 mm from proximal end Figure A.6: δ13C and δ18O (V-PDB) canine enamel profile plot of Arabuko Sokoke #3 hippo (0mm = death). 154 Arabuko Sokoke #4 (AS166) 0 δ18O δ13C −1 −1 −2 −2 −3 −3 −4 −4 −5 −5 −6 −6 −7 −7 50 100 150 200 δ18Oenamel δ13Cenamel 0 250 mm from proximal end Figure A.7: δ13C and δ18O (V-PDB) canine enamel profile plot of Arabuko Sokoke #4 hippo (0mm = death). 155 Buffalo Springs −1 δ18O δ13C −2 −2 −3 −3 −4 −4 −5 −5 0 50 100 150 200 250 300 350 400 δ18Oenamel δ13Cenamel −1 450 mm from proximal end Figure A.8: δ13C and δ18O (V-PDB) canine enamel profile plot of Buffalo Springs hippo (0mm = death). 156 Chyulu −4 1 δ18O δ13C −6 −1 −7 −2 −8 −3 −9 −4 −10 −5 δ13Cenamel 0 0 50 100 150 200 250 δ18Oenamel −5 300 mm from proximal end Figure A.9: δ13C and δ18O (V-PDB) canine enamel profile plot of Chyulu hippo (0mm = death). 157 Kisumu (Lake Victoria - Winam Gulf) 0 0 δ18O δ13C −4 −4 −6 −6 −8 −8 −10 −10 δ13Cenamel −2 0 50 100 150 200 250 300 350 δ18Oenamel −2 400 mm from proximal end Figure A.10: δ13C and δ18O (V-PDB) canine enamel profile plot of Kisumu hippo (0mm = death). 158 Laikipia 1 δ18O δ13C 0 0 −1 −1 −2 −2 −3 −3 −4 −4 −5 −5 250 300 350 400 mm from proximal end δ18Oenamel δ13Cenamel 1 450 Figure A.11: δ13C and δ18O (V-PDB) canine enamel profile plot of Laikipia hippo (0mm = death). 159 Mara (Maasai Mara NR) 1 δ18O δ13C 0 0 −1 −1 −2 −2 −3 −3 −4 −4 −5 −5 −6 −6 0 50 100 150 200 250 300 δ18Oenamel δ13Cenamel 1 350 mm from proximal end Figure A.12: δ13C and δ18O (V-PDB) canine enamel profile plot of Mara hippo (0mm = death). 160 Meru NP −2 −2 −4 −4 −6 −6 −8 −8 0 50 100 150 200 δ18Oenamel δ13Cenamel δ18O δ13C 250 mm from proximal end Figure A.13: δ13C and δ18O (V-PDB) canine enamel profile plot of Meru NP hippo (0mm = death). 161 Minjila (KWS Minjila - Tana River) 1 δ18O δ13C 0 0 −1 −1 −2 −2 −3 −3 −4 −4 0 50 100 150 200 250 300 350 400 450 δ18Oenamel δ13Cenamel 1 500 mm from proximal end Appendix Figure 14: δ13C and δ18O (V-PDB) canine enamel profile plot of Minjila hippo (0mm = death). 162 Mokowe δ18O δ13C 3 0 1 −2 0 −3 −1 −4 −2 −5 δ13Cenamel −1 0 50 100 150 200 250 300 350 400 450 δ18Oenamel 2 500 mm from proximal end 13 18 Appendix Figure 15: δ C and δ O (V-PDB) canine enamel profile plot of Mokowe hippo (0mm = death). 163 Mpeketoni δ18O δ13C 1 0 0 −1 −1 −2 −2 −3 −3 −4 −4 0 50 100 150 200 250 300 350 400 450 500 δ18Oenamel δ13Cenamel 1 550 mm from proximal end Appendix Figure 16: δ13C and δ18O (V-PDB) canine enamel profile plot of Mpeketoni hippo (0mm = death). 164 Mwea NR − Gitaru Dam 0 0 −2 −2 −4 −4 −6 −6 −8 −8 0 50 100 150 200 250 mm from proximal end 300 350 δ18Oenamel δ13Cenamel δ18O δ13C 400 Appendix Figure 17: δ13C and δ18O (V-PDB) canine enamel profile plot of Mwea – Gitaru Dam hippo (0mm = death). 165 Mwea NR - 2009 δ18O δ13C −2 −2 −6 −6 −8 −8 −10 −10 δ13Cenamel −4 0 50 100 150 200 250 300 350 400 450 500 δ18Oenamel −4 550 mm from proximal end Appendix Figure 18: δ13C and δ18O (V-PDB) canine enamel profile plot of Mwea #1 hippo (0mm = death). 166 Mwea #2 (Mwea NR - 2005) 0 0 δ18O δ13C −4 −4 −6 −6 −8 −8 −10 −10 δ13Cenamel −2 0 50 100 150 200 250 300 350 400 450 500 δ18Oenamel −2 550 mm from proximal end Appendix Figure 19: δ13C and δ18O (V-PDB) canine enamel profile plot of Mwea #2 hippo (0mm = death). 167 Mwea #3 (Mwea NP 2007) δ18O δ13C −2 −2 −6 −6 −8 −8 −10 −10 δ13Cenamel −4 0 50 100 150 200 250 300 350 400 mm from proximal end 450 500 550 δ18Oenamel −4 600 Appendix Figure 20: δ13C and δ18O (V-PDB) canine enamel profile plot of Mwea #3 hippo (0mm = death). 168 Naivasha - Crater Lake (KWS-Naivasha-H Crater Lake) 8 δ18O δ13C 0 6 −2 4 −4 2 −6 0 −8 −2 0 50 100 150 200 250 300 350 400 mm from proximal end 450 500 550 δ18Oenamel δ13Cenamel 2 600 Appendix Figure 21: δ13C and δ18O (V-PDB) canine enamel profile plot of Naivasha hippo (0mm = death). 169 Naivasha - Crayfish Camp (KWS-NAIV-CF-1108) 5 δ18O δ13C 1 4 0 3 −1 2 −2 1 −3 0 −4 −1 −5 −2 0 50 100 150 200 250 300 350 400 mm from proximal end 450 500 550 δ18Oenamel δ13Cenamel 2 600 Appendix Figure 22: δ13C and δ18O (V-PDB) canine enamel profile plot of Naivasha hippo (0mm = death). 170 Lake Nakuru (KWS-LNakuru-Kabutini-5) δ18O δ13C 2 1 1 0 0 −1 −1 −2 −2 −3 −3 0 50 100 150 200 250 δ18Oenamel δ13Cenamel 2 300 mm from proximal end Appendix Figure 23: δ13C and δ18O (V-PDB) canine enamel profile plot of Lake Nakuru hippo (0mm = death). 171 Lake Ol Bolossat 2 δ18O δ13C −2 1 −3 0 −4 −1 −5 −2 −6 −3 −7 −4 −8 −5 0 50 100 150 200 mm from proximal end 250 δ18Oenamel δ13Cenamel −1 300 Appendix Figure 24: δ13C and δ18O (V-PDB) canine enamel profile plot of Lake Ol Bolossat hippo (0mm = death). 172 Turkana - Koobi Fora #2 (KEN-09-115) 4 δ18O δ13C 3 3 2 2 1 1 0 0 −1 −1 0 50 100 150 200 250 300 350 400 δ18Oenamel δ13Cenamel 4 450 mm from proximal end 13 18 Appendix Figure 25: δ C and δ O (V-PDB) canine enamel profile plot of Turkana – Koobi Fora 2 hippo (0mm = death). 173 Witu 1 δ18O δ13C 0 0 −1 −1 −2 −2 −3 −3 −4 −4 −5 −5 0 50 100 150 200 250 300 350 400 450 500 δ18Oenamel δ13Cenamel 1 550 mm from proximal end Appendix Figure 26: δ13C and δ18O (V-PDB) canine enamel profile plot of Witu hippo (0mm = death). 174 Table A.1: Aberdares NP hippo isotope values. Specimen Locality Position Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP Aberdares NP 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 265 275 285 295 305 315 325 δ13C (V-PDB) -1.4 -1.5 -1.0 -1.3 -1.4 -1.7 -2.1 -2.3 -2.5 -2.2 -2.2 -1.8 -1.9 -1.7 -1.9 -2.1 -2.4 -2.3 -2.5 -3.4 -3.4 -3.3 -3.0 -3.2 -3.4 -3.5 -3.5 -3.5 -4.0 -4.2 -4.4 -4.6 -4.9 δ 18O (V-PDB) -1.8 -2.3 -1.8 -1.8 -0.9 -1.6 -1.6 -1.5 -2.1 -2.6 -2.8 -3.0 -3.2 -3.3 -2.9 -3.3 -2.7 -2.7 -2.5 -2.5 -2.7 -2.7 -2.6 -2.8 -2.8 -2.8 -2.7 -3.0 -3.1 -3.0 -3.0 -3.2 -3.3 175 Table A.2: Adhi Dam (Boni NR) hippo isotope values. Specimen KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam Locality Position Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 245 255 265 275 285 295 305 315 325 335 345 355 365 375 385 395 405 δ13C δ 18O (V-PDB) (V-PDB) -7.9 -1.6 -7.1 0.3 -5.7 -0.2 0.1 -5.5 -5.7 -0.7 -6.8 -1.2 -7.2 -1.7 -7.0 -1.6 -7.0 -1.4 -6.2 -1.2 -5.7 -1.3 -5.4 -1.1 -5.3 -2.0 -5.5 -1.9 -5.5 -2.2 -5.5 -1.7 -5.2 -1.9 -6.2 -1.6 -5.0 -1.4 -6.2 -1.6 -6.6 -1.6 -7.2 -1.7 -7.0 -1.8 -7.7 -1.7 -7.0 -2.0 -6.7 -1.6 -6.3 -1.5 -6.4 -1.6 -6.4 -1.6 -7.0 -1.8 -6.7 -2.2 -7.0 -2.1 -7.0 -3.1 -6.8 -2.9 -6.5 -3.4 -6.5 -3.1 -5.6 -3.7 -5.1 -3.5 -3.9 -2.7 -3.0 -2.5 176 Table A.2: (cont’d) Specimen Locality Position KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam KWS-Adhidam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam Adhi Dam 415 425 455 465 475 485 495 δ13C δ 18O (V-PDB) (V-PDB) -2.3 -1.3 -2.4 -1.7 -2.3 -1.1 -2.3 -1.4 -2.0 -1.9 -2.5 -1.4 -2.9 -1.8 177 Table A.3: Amboseli #1 hippo isotope values. Specimen Locality Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Position 5 15 25 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 265 275 285 295 305 315 325 335 345 355 365 375 385 395 405 δ13C (V-PDB) -0.9 -1.3 -1.0 -1.9 -1.6 -1.2 -1.1 -2.5 -2.3 -1.8 -1.1 -1.3 -1.4 -2.4 -2.8 -2.2 -1.7 -1.6 -3.5 -2.3 -2.0 -1.7 -1.8 -1.8 -0.9 -1.1 -0.5 -0.8 -1.4 -1.2 -1.2 -0.9 -1.5 -1.7 -1.3 -1.2 -0.9 -0.8 -1.2 -1.1 δ 18O (V-PDB) -5.9 -6.1 -6.0 -5.7 -6.1 -6.2 -6.1 -6.0 -6.2 -6.1 -5.9 -5.9 -6.0 -5.6 -6.2 -6.5 -5.9 -5.8 -5.9 -5.3 -5.6 -5.5 -5.2 -5.5 -5.6 -5.1 -5.3 -5.1 -5.4 -5.3 -5.2 -5.7 -5.5 -5.3 -5.8 -5.9 -5.4 -5.6 -5.5 -5.7 178 Table A.3: (cont’d) Specimen Locality Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli-H1-H2 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Amboseli #1 Position 415 425 435 445 455 465 δ13C (V-PDB) -0.7 -0.3 -0.3 -0.9 -0.9 -1.4 δ 18O (V-PDB) -5.8 -5.4 -5.6 -5.7 -5.5 -6.0 179 Table A.4: Amboseli #2 hippo isotope values. Specimen KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 Locality Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Amboseli #2 Position 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 265 275 285 295 305 315 325 335 345 355 365 375 385 395 δ13C (V-PDB) -1.8 -3.3 -2.4 -1.1 -1.2 -1.1 -1.6 -1.3 -1.2 -1.1 -2.0 -1.7 -1.6 -1.7 -1.8 -2.3 -2.2 -1.9 -2.0 -2.2 -3.5 -2.8 -3.0 -2.6 -3.2 -2.8 -2.4 -2.4 -2.3 -3.3 -2.9 -2.6 -2.9 -2.6 -2.4 -2.0 -2.4 -1.8 -1.6 -1.7 δ 18O (V-PDB) -4.8 -4.1 -4.7 -5.2 -4.6 -5.2 -5.2 -5.3 -5.4 -5.5 -5.6 -5.7 -5.6 -5.5 -4.8 -5.6 -5.5 -5.4 -5.2 -5.1 -4.7 -4.9 -4.9 -4.9 -5.1 -5.2 -5.3 -5.5 -4.5 -4.5 -4.9 -4.8 -4.9 -5.0 -5.5 -6.1 -6.1 -5.3 -5.3 -5.1 180 Table A.4: (cont’d) Specimen Locality KWS-Amb-0401 KWS-Amb-0401 KWS-Amb-0401 Amboseli #2 Amboseli #2 Amboseli #2 Position 405 415 425 δ13C (V-PDB) -1.6 -1.3 -1.8 δ 18O (V-PDB) -5.7 -5.1 -4.8 181 Table A.5: Arabuko Sokoke #1 hippo isotope values. Specimen Locality Position Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko-Sokoke-AS-168 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 Arabuko Sokoke #1 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 264 271.5 280 288 296 307 315 320 327 333 340 δ13C (V-PDB) -5.9 -6.1 -6.3 -4.7 -3.5 -2.6 -2.3 -1.9 -1.7 -1.5 -0.7 -0.7 -1.4 -1.3 -1.0 -0.8 -0.6 -0.4 -0.3 -0.5 -0.5 -0.5 -0.5 -0.6 -0.6 -0.6 -0.2 -0.1 0.0 -0.4 -1.4 -1.9 -2.3 -2.6 -2.7 -2.4 -2.5 δ 18O (V-PDB) -4.3 -5.1 -5.4 -6.0 -5.1 -4.5 -4.7 -4.9 -4.4 -4.0 -3.8 -3.4 -4.6 -4.4 -4.5 -3.9 -4.8 -5.0 -4.2 -4.5 -5.3 -5.0 -5.2 -5.4 -5.3 -4.6 -4.4 -4.0 -3.7 -4.0 -3.9 -4.1 -4.5 -3.8 -3.6 -4.5 -4.5 182 Table A.6: Arabuko Sokoke #2 hippo isotope values. Specimen Locality Position K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 K00-AS-167 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 Arabuko Sokoke #2 24 34 46 56 69 79 91 101 112 123 135 146 157 167 179 192 206 216 228 240 263 275 288 298 308 319 332 342 353 364 375 385 399 415 432 δ13C (V-PDB) -1.9 -1.2 -1.8 -1.5 -1.6 -1.6 -1.4 -1.2 -0.9 -0.7 -0.7 -0.6 -0.4 -0.5 -1.1 -1.0 -1.5 -2.5 -3.9 -3.9 -4.3 -4.5 -4.5 -4.1 -3.9 -4.1 -4.0 -3.4 -2.8 -2.4 -2.3 -2.0 -2.1 -2.7 -4.1 δ 18O (V-PDB) -2.2 -2.3 -2.1 -2.1 -2.2 -2.3 -2.5 -2.7 -3.2 -3.0 -3.4 -3.1 -3.1 -3.5 -4.1 -4.4 -4.2 -4.4 -4.0 -4.1 -4.2 -3.9 -3.8 -3.4 -3.7 -3.8 -3.3 -3.4 -3.8 -3.5 -3.4 -3.6 -3.6 -3.7 -4.0 183 Table A.7: Arabuko Sokoke #3 hippo isotope values. Specimen Locality Position Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko-Sokoke-08 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 Arabuko Sokoke #3 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 265 275 285 295 δ13C (V-PDB) -3.8 -3.5 -3.5 -3.6 -3.4 -3.2 -3.2 -2.9 -3.0 -3.2 -3.6 -4.4 -4.4 -4.8 -4.6 -4.2 -3.9 -4.0 -3.4 -3.6 -3.6 -3.7 -3.5 -3.3 -3.6 -3.4 -3.0 -3.3 -3.1 -3.5 δ 18O (V-PDB) -3.8 -3.3 -4.4 -3.5 -3.2 -3.3 -3.6 -2.8 -2.5 -3.2 -3.1 -3.6 -3.0 -4.1 -3.6 -3.7 -3.6 -3.9 -3.0 -2.7 -2.6 -2.6 -2.7 -2.5 -2.8 -2.9 -3.1 -2.9 -3.1 -3.3 184 Table A.8: Buffalo Springs hippo isotope values. Specimen Locality Position Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 265 275 285 295 305 315 325 335 345 355 365 375 385 395 δ13C (V-PDB) -4.3 -3.8 -3.3 -3.2 -3.6 -4.1 -3.7 -3.1 -2.5 -2.2 -2.4 -2.4 -2.3 -2.3 -3.2 -3.6 -3.8 -3.8 -3.7 -3.4 -3.4 -3.3 -3.3 -3.3 -2.5 -2.3 -2.8 -2.9 -3.0 -3.0 -3.0 -3.3 -3.3 -3.6 -3.6 -3.4 -3.2 -3.3 -3.3 -3.3 δ 18O (V-PDB) -3.1 -2.5 -2.8 -2.9 -3.1 -2.9 -3.3 -3.3 -3.4 -3.5 -3.6 -3.7 -3.9 -3.5 -3.7 -3.8 -3.5 -3.2 -3.4 -3.4 -3.5 -4.0 -3.9 -4.2 -3.9 -3.7 -3.5 -3.8 -3.6 -3.7 -3.4 -3.2 -3.3 -3.4 -3.5 -3.3 -3.5 -3.6 -3.8 -3.5 185 Table A.8: (cont’d) Specimen Locality Position Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo-Springs Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR Buffalo Springs NR 405 415 425 435 445 455 465 δ13C (V-PDB) -3.3 -3.5 -3.6 -3.8 -3.4 -3.5 -3.3 δ 18O (V-PDB) -3.9 -3.7 -3.7 -3.7 -3.7 -3.7 -3.7 186 Table A.9: Chyulu hippo isotope values Specimen Locality Position Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu Chyulu 5 15 25 35 45 55 65 75 85 95 105 125 135 145 155 165 175 185 195 205 215 225 235 245 255 265 275 285 295 305 315 δ13C (V-PDB) -5.1 -6.5 -7.7 -8.1 -8.8 -8.1 -6.8 -7.8 -7.5 -8.0 -7.8 -7.5 -6.9 -6.5 -6.6 -6.8 -6.6 -5.8 -6.2 -6.0 -6.4 -8.5 -8.1 -7.4 -6.5 -6.2 -6.3 -6.3 -6.5 -6.4 -6.5 δ 18O (V-PDB) -0.6 -0.9 -0.3 -0.4 -1.4 -2.3 -2.8 -3.6 -4.2 -4.2 -4.1 -4.0 -5.0 -4.5 -4.3 -4.3 -4.5 -4.2 -4.4 -4.4 -3.0 -5.0 -4.6 -4.4 -4.5 -4.4 -4.1 -4.6 -4.9 -4.7 -4.9 187 Table A.10: Kisumu (Lake Victoria) hippo isotope values δ13C δ 18O Specimen Locality Position (V-PDB) (V-PDB) Kisumu Kisumu 5 -6.9 -4.7 Kisumu Kisumu 15 -7.6 -4.4 Kisumu Kisumu 25 -7.8 -4.2 Kisumu Kisumu 35 -8.9 -4.1 Kisumu Kisumu 45 -8.1 -3.9 Kisumu Kisumu 55 -7.5 -3.9 Kisumu Kisumu 65 -7.8 -4.3 Kisumu Kisumu 75 -7.4 -3.7 Kisumu Kisumu 95 -7.7 -4.3 Kisumu Kisumu 105 -7.2 -4.1 Kisumu Kisumu 115 -7.1 -4.2 Kisumu Kisumu 125 -7.7 -4.3 Kisumu Kisumu 135 -7.6 -4.0 Kisumu Kisumu 145 -6.7 -4.4 Kisumu Kisumu 155 -6.0 -3.9 Kisumu Kisumu 165 -5.7 -4.3 Kisumu Kisumu 175 -6.3 -3.8 Kisumu Kisumu 185 -6.2 -4.2 Kisumu Kisumu 195 -6.7 -4.1 Kisumu Kisumu 205 -5.9 -4.0 Kisumu Kisumu 215 -5.4 -4.5 Kisumu Kisumu 225 -5.6 -4.2 Kisumu Kisumu 235 -5.5 -3.7 Kisumu Kisumu 245 -6.0 -3.7 Kisumu Kisumu 255 -5.9 -3.8 Kisumu Kisumu 265 -5.7 -3.8 Kisumu Kisumu 275 -6.5 -3.6 Kisumu Kisumu 285 -6.3 -3.4 Kisumu Kisumu 295 -5.8 -3.4 Kisumu Kisumu 305 -5.4 -3.7 Kisumu Kisumu 315 -3.9 -3.1 Kisumu Kisumu 325 -5.1 -3.6 Kisumu Kisumu 355 -5.9 -3.4 Kisumu Kisumu 365 -6.0 -4.0 Kisumu Kisumu 375 -5.4 -3.8 Kisumu Kisumu 385 -5.1 -3.3 Kisumu Kisumu 395 -5.8 -3.3 Kisumu Kisumu 405 -6.3 -3.4 188 Table A.11: Laikipia hippo isotope values (tusk still in skull – estimated 240mm missing). δ13C δ 18O Specimen Locality Position (V-PDB) (V-PDB) K01-LAI-191 Laikipia 245 -2.8 -4.0 K01-LAI-191 Laikipia 255 -2.5 -4.3 K01-LAI-191 Laikipia 265 -2.9 -3.5 K01-LAI-191 Laikipia 275 -2.9 -3.8 K01-LAI-191 Laikipia 285 -3.0 -3.2 K01-LAI-191 Laikipia 295 -2.8 -3.3 K01-LAI-191 Laikipia 305 -2.5 -3.1 K01-LAI-191 Laikipia 315 -2.7 -2.4 K01-LAI-191 Laikipia 325 -2.6 -2.0 K01-LAI-191 Laikipia 335 -2.5 -2.1 K01-LAI-191 Laikipia 345 -2.4 -3.0 K01-LAI-191 Laikipia 355 -2.5 -2.5 K01-LAI-191 Laikipia 365 -2.3 -2.7 K01-LAI-191 Laikipia 375 -2.1 -2.1 K01-LAI-191 Laikipia 385 -2.2 -1.9 K01-LAI-191 Laikipia 395 -2.0 -2.1 K01-LAI-191 Laikipia 405 -2.9 -1.2 K01-LAI-191 Laikipia 415 -3.2 -1.1 K01-LAI-191 Laikipia 425 -3.2 -1.1 K01-LAI-191 Laikipia 435 -2.9 -1.8 K01-LAI-191 Laikipia 445 -2.8 -1.5 K01-LAI-191 Laikipia 455 -2.8 -1.7 K01-LAI-191 Laikipia 465 -2.9 -1.1 K01-LAI-191 Laikipia 475 -3.3 -1.6 189 Table A.12: Maasai Mara hippo isotope values. δ13C Specimen Locality Position (V-PDB) Mara Maasai Mara 5 -3.3 Mara Maasai Mara 15 -3.0 Mara Maasai Mara 25 -2.6 Mara Maasai Mara 35 -3.3 Mara Maasai Mara 45 -3.5 Mara Maasai Mara 55 -3.5 Mara Maasai Mara 65 -3.4 Mara Maasai Mara 75 -3.2 Mara Maasai Mara 85 -3.7 Mara Maasai Mara 95 -3.9 Mara Maasai Mara 105 -3.7 Mara Maasai Mara 115 -3.3 Mara Maasai Mara 125 -3.6 Mara Maasai Mara 135 -3.3 Mara Maasai Mara 155 -3.1 Mara Maasai Mara 165 -3.0 Mara Maasai Mara 175 -2.5 Mara Maasai Mara 185 -3.1 Mara Maasai Mara 195 -3.2 Mara Maasai Mara 205 -3.1 Mara Maasai Mara 215 -3.2 Mara Maasai Mara 225 -2.8 Mara Maasai Mara 235 -3.1 Mara Maasai Mara 245 -3.5 Mara Maasai Mara 255 -3.0 Mara Maasai Mara 265 -3.0 Mara Maasai Mara 285 -3.7 Mara Maasai Mara 295 -2.8 Mara Maasai Mara 305 -3.2 Mara Maasai Mara 315 -2.5 Mara Maasai Mara 325 -1.8 Mara Maasai Mara 335 -1.9 Mara Maasai Mara 345 -3.0 δ 18O (V-PDB) -3.3 -4.0 -3.6 -4.0 -4.6 -4.0 -4.2 -4.2 -4.4 -4.5 -4.3 -4.1 -3.8 -4.0 -4.1 -5.1 -5.5 -4.4 -4.0 -4.2 -4.2 -4.2 -4.0 -4.3 -4.2 -4.0 -4.2 -4.4 -4.2 -4.1 -3.3 -3.7 -3.7 190 Table A.13: Meru NP hippo isotope values. Specimen Locality Position Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP Meru NP 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 255 265 275 δ13C (V-PDB) -2.5 -1.7 -2.2 -1.8 -2.4 -2.9 -3.2 -3.6 -3.6 -3.6 -3.4 -3.6 -3.9 -4.4 -4.9 -5.5 -5.9 -6.9 -7.1 -8.4 -7.8 -8.3 -8.1 -8.1 -8.1 -8.5 -8.6 δ 18O (V-PDB) -4.1 -4.0 -4.3 -3.5 -2.9 -3.2 -3.9 -3.9 -3.7 -3.9 -4.2 -4.4 -4.5 -4.4 -5.0 -5.1 -5.1 -5.1 -5.2 -7.1 -5.6 -5.7 -5.8 -6.0 -5.8 -5.8 -5.6 191 Table A.14: Minjila hippo isotope values. Specimen Locality Position KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 265 275 285 295 305 315 325 335 345 355 365 375 385 395 δ13C (V-PDB) -1.0 -1.4 -1.5 -1.8 -1.9 -1.9 -1.6 -1.2 -1.3 -0.9 -1.2 -1.2 -1.3 -1.3 -1.6 -1.8 -1.7 -2.0 -2.1 -2.3 -2.4 -2.3 -2.1 -2.0 -1.9 -1.6 -1.5 -1.5 -1.5 -1.5 -1.4 -1.6 -1.8 -1.8 -1.7 -1.8 -2.2 -2.2 -2.5 -2.3 δ 18O (V-PDB) 0.0 -0.7 -0.9 -0.8 -1.4 -2.3 -2.4 -2.3 -2.8 -2.5 -3.0 -2.3 -2.6 -2.8 -2.9 -2.7 -2.2 -2.6 -2.6 -2.6 -2.7 -2.4 -2.7 -3.1 -2.9 -3.0 -2.6 -2.4 -2.1 -1.7 -1.5 -1.8 -0.9 -0.9 -0.9 -0.9 -2.0 -1.2 -1.6 -1.9 192 Table A.14: (cont’d) Specimen Locality Position KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila KWS-Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila Minjila 405 415 425 435 445 455 465 475 485 495 505 525 δ13C (V-PDB) -2.5 -3.1 -2.7 -2.9 -3.0 -3.1 -3.4 -3.6 -3.7 -3.6 -4.0 -1.8 δ 18O (V-PDB) -2.1 -1.9 -1.6 -2.6 -2.7 -3.0 -3.2 -3.5 -3.1 -3.2 -3.6 -1.2 193 Table A.15: Mokowe hippo isotope values. Specimen Locality Position Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 265 275 285 295 305 315 325 335 345 355 365 375 385 395 δ13C (V-PDB) 0.5 0.0 -0.1 -0.2 -0.2 -0.5 -0.5 -0.7 -0.6 -0.5 -0.2 -0.1 -0.2 0.1 0.2 0.1 -0.5 -0.4 -0.3 0.0 -0.3 -0.3 -0.5 -0.3 -0.3 -0.4 -0.2 0.1 0.3 0.1 -0.1 -0.1 0.1 -0.3 0.1 0.0 0.1 0.1 -0.1 0.0 δ 18O (V-PDB) -4.2 -3.8 -3.7 -3.6 -3.7 -3.4 -3.1 -2.9 -3.1 -3.1 -3.3 -3.4 -3.2 -3.2 -3.6 -3.6 -4.0 -4.0 -3.9 -3.7 -3.7 -3.7 -3.6 -3.5 -3.3 -3.4 -3.4 -3.5 -3.6 -3.7 -3.8 -3.9 -3.7 -4.0 -3.7 -3.6 -3.5 -3.1 -3.1 -3.4 194 Table A.15: (cont’d) Specimen Locality Position Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe Mokowe 405 415 425 435 445 455 465 475 485 δ13C (V-PDB) 0.0 -0.2 -0.3 1.2 0.0 0.1 -0.3 -0.3 1.2 δ 18O (V-PDB) -3.4 -3.3 -3.4 -3.6 -3.5 -3.4 -3.5 -3.4 -3.5 195 Table A.16: Mpeketoni hippo isotope values. Specimen Locality Position Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 275 285 295 305 315 325 335 345 355 365 375 385 395 405 δ13C (V-PDB) -2.6 -2.7 -2.4 -2.7 -2.5 -2.1 -2.1 -1.1 -2.3 -2.5 -2.0 -2.1 -2.2 -2.0 -1.9 -2.1 -2.2 -2.1 -2.1 -2.0 -2.0 -1.7 -1.6 -2.0 -1.5 -1.7 -1.7 -1.7 -1.6 -1.5 -1.4 -1.6 -1.6 -1.9 -2.0 -2.1 -2.1 -2.4 -2.3 -2.6 δ 18O (V-PDB) -0.8 0.0 0.1 -0.1 -0.2 0.8 0.1 0.0 0.2 -1.2 0.0 -0.8 -0.5 0.0 0.8 -0.8 -0.8 -0.7 -0.8 -2.6 -1.4 -0.4 -0.6 -1.2 -1.2 -1.3 -2.0 -2.1 -2.4 -2.8 -2.7 -2.4 -2.1 -1.9 -2.0 -3.0 -2.5 -3.1 -3.3 -1.8 196 Table A.16: (cont’d) Specimen Locality Position Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni Mpeketoni 415 425 435 455 465 475 485 495 505 515 525 535 δ13C (V-PDB) -2.5 -2.6 -2.5 -2.5 -2.4 -2.3 -2.1 -1.8 -1.7 -2.3 -2.0 -2.3 δ 18O (V-PDB) -2.5 -1.4 -2.9 -2.8 -1.5 -1.8 -2.1 -1.9 -2.0 -2.2 -2.9 -3.1 197 Table A.17: Mwea – Gitaru Dam hippo isotope values Specimen Locality Position Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea-Gitaru Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam Mwea - Gitaru Dam 5 15 35 45 55 75 85 95 105 115 125 135 145 155 165 175 185 205 215 235 245 255 265 285 295 305 315 325 335 345 355 365 375 385 395 405 δ13C (V-PDB) -7.6 -6.6 -6.3 -5.8 -5.1 -4.6 -5.1 -5.0 -5.1 -5.4 -5.4 -5.5 -5.1 -5.4 -5.5 -5.6 -5.3 -5.3 -5.4 -5.7 -5.7 -5.9 -6.4 -5.8 -5.9 -6.0 -5.8 -5.4 -5.3 -5.7 -6.0 -6.0 -6.2 -5.8 -5.6 -5.3 δ 18O (V-PDB) -2.7 -2.0 -2.4 -3.3 -2.9 -2.7 -3.0 -3.2 -3.8 -3.5 -3.4 -3.2 -3.0 -3.1 -3.3 -3.2 -2.9 -3.5 -3.4 -3.7 -3.9 -4.1 -4.3 -4.2 -3.9 -4.1 -3.9 -3.6 -3.4 -3.6 -3.3 -3.7 -3.3 -3.2 -3.3 -3.2 198 Table A.18: Mwea #1 isotope values. Specimen Locality Position Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 265 275 285 295 305 315 325 335 345 355 365 375 385 395 δ13C (V-PDB) -9.4 -8.4 -9.1 -8.4 -7.5 -7.5 -7.3 -7.7 -7.1 -6.9 -6.2 -6.0 -5.9 -6.5 -6.6 -6.5 -6.4 -6.0 -5.4 -5.2 -4.8 -4.5 -4.6 -4.6 -5.3 -5.6 -6.5 -6.0 -6.9 -6.7 -6.2 -6.6 -6.5 -6.6 -6.5 -6.2 -5.8 -5.7 -5.2 -5.1 δ 18O (V-PDB) -3.3 -2.8 -2.2 -2.7 -3.3 -3.3 -3.3 -2.6 -3.9 -3.5 -3.5 -3.2 -3.4 -3.1 -3.3 -3.2 -3.7 -3.6 -3.9 -4.0 -3.9 -4.9 -5.0 -5.3 -5.3 -5.1 -5.7 -5.5 -5.4 -5.3 -4.2 -3.8 -4.0 -4.1 -3.7 -3.9 -3.9 -4.4 -4.0 -4.4 199 Table A.18: (cont’d) Specimen Locality Position Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea-09 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 Mwea #1 405 415 425 435 445 455 465 475 485 495 505 525 535 545 555 565 575 δ13C (V-PDB) -4.8 -4.7 -5.0 -5.0 -5.2 -5.0 -4.7 -4.5 -4.7 -4.9 -4.6 -4.9 -5.3 -5.4 -4.9 -4.6 -5.0 δ 18O (V-PDB) -3.6 -3.8 -3.7 -4.1 -3.9 -4.4 -4.2 -4.3 -4.3 -4.4 -4.4 -4.7 -4.7 -4.6 -4.9 -4.3 -4.8 200 Table A.19: Mwea #2 hippo isotope values. Specimen Locality Position Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 liMwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 265 275 285 295 305 315 325 335 345 355 365 375 385 395 δ13C (V-PDB) -9.2 -10.0 -9.6 -8.9 -8.0 -7.2 -7.5 -7.3 -7.1 -7.2 -6.7 -6.1 -6.0 -5.7 -6.0 -6.8 -7.5 -7.3 -6.7 -6.3 -6.5 -5.6 -5.5 -5.7 -5.5 -5.9 -5.7 -4.6 -5.9 -6.3 -6.8 -6.7 -6.5 -6.5 -5.2 -6.6 -6.8 -6.8 -6.7 -6.2 δ 18O (V-PDB) -3.7 -3.4 -1.9 -1.6 -1.8 -5.9 -3.6 -3.0 -4.1 -3.7 -4.3 -4.4 -4.4 -4.2 -4.3 -4.7 -4.9 -5.5 -5.5 -5.9 -6.4 -5.6 -5.3 -5.8 -5.7 -5.9 -6.3 -4.5 -6.2 -6.1 -6.1 -6.0 -5.4 -5.3 -4.4 -5.2 -5.1 -4.7 -4.7 -4.4 201 Table A.19: (cont’d) Specimen Locality Position Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea-NR-MAR-2005 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 Mwea #2 405 415 425 435 445 455 465 475 485 495 505 515 525 535 545 555 565 575 δ13C (V-PDB) -5.8 -4.3 -4.5 -5.2 -4.4 -4.5 -5.0 -4.6 -4.3 -4.2 -4.6 -4.6 -4.4 -4.4 -4.2 -4.2 -4.7 -4.3 δ 18O (V-PDB) -4.6 -4.8 -2.3 -2.9 -3.0 -3.0 -3.1 -3.4 -3.6 -3.4 -3.4 -4.4 -3.7 -3.0 -2.7 -3.8 -3.7 -3.7 202 Table A.20: Mwea #3 hippo isotope values. Specimen Locality Position Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 265 275 285 295 305 315 325 335 345 355 365 375 385 395 δ13C (V-PDB) -6.4 -6.2 -6.3 -6.3 -6.7 -7.5 -7.5 -7.3 -7.5 -7.8 -8.1 -9.4 -8.8 -7.6 -6.6 -6.3 -5.8 -5.1 -4.6 -5.0 -6.9 -5.1 -5.4 -5.4 -5.5 -5.1 -5.4 -5.5 -5.6 -5.3 -7.3 -7.0 -7.3 -7.3 -6.6 -6.0 -6.1 -7.0 -7.3 -7.4 δ 18O (V-PDB) -2.7 -2.3 -2.9 -3.0 -3.0 -3.1 -3.4 -3.6 -3.4 -3.4 -3.1 -3.7 -3.0 -2.7 -3.8 -3.7 -3.7 -4.2 -4.5 -5.0 -4.8 -4.3 -4.3 -4.4 -4.0 -3.9 -3.6 -3.4 -3.3 -3.1 -3.4 -2.8 -3.0 -3.1 -2.6 -3.3 -3.2 -4.0 -3.3 -3.6 203 Table A.20: (cont’d) Specimen Locality Position Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea-NP-2007 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 Mwea #3 405 415 425 435 445 455 465 475 485 495 505 515 525 535 545 555 565 575 585 595 605 δ13C (V-PDB) -7.3 -7.2 -7.3 -7.3 -6.6 -6.4 -6.5 -6.7 -6.3 -6.1 -6.1 -6.2 -5.9 -5.6 -5.9 -6.2 -6.5 -6.5 -7.2 -7.6 -7.9 δ 18O (V-PDB) -3.1 -3.0 -3.0 -3.4 -3.1 -4.0 -3.1 -3.5 -3.4 -3.5 -3.4 -3.3 -3.1 -3.4 -3.1 -3.1 -3.1 -3.2 -3.8 -4.1 -4.6 204 Table A.21: Naivasha – Crater Lake hippo isotope values. Specimen Locality Pos. KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 265 275 285 295 305 315 325 335 345 355 365 375 385 395 405 δ13C (V-PDB) 0.0 -0.5 -0.5 -1.0 -1.1 -0.8 -0.6 -0.6 -0.2 0.2 0.1 0.1 -0.3 -0.7 -1.1 -1.0 -1.1 -1.0 -1.7 -2.1 -2.6 -2.3 -2.3 -2.5 -2.9 -3.2 -3.5 -3.8 -3.5 -2.8 -2.6 -2.3 -2.7 -3.7 -4.3 -4.1 -3.6 -3.6 -5.1 -5.3 δ 18O (V-PDB) -1.7 -2.2 -2.2 -2.7 -2.9 -3.5 -4.0 -4.3 -4.2 -4.8 -5.1 -5.5 -5.1 -5.2 -5.2 -4.7 -5.0 -5.1 -4.8 -4.7 -4.6 -4.5 -4.4 -4.5 -4.7 -4.4 -4.5 -3.9 -4.0 -3.9 -3.8 -3.3 -3.9 -3.0 -2.9 -2.8 -3.4 -3.1 -2.8 -2.4 205 Table A.21: (cont’d) Specimen Locality Pos. KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL KWS-Naivasha-CraterL Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake Naivasha - Crater Lake 415 425 435 445 455 465 475 485 495 505 515 525 δ13C (V-PDB) -6.7 -6.4 -7.4 -7.4 -7.1 -6.5 -4.2 -1.4 -5.6 -4.9 -4.8 -4.5 δ 18O (V-PDB) -2.1 -2.4 -2.1 -2.3 -2.3 -3.0 -3.9 -3.6 -3.0 -3.3 -3.4 -3.8 206 Table A.22: Naivasha – Crayfish hippo isotope values. Specimen Locality Position KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 265 275 285 295 305 315 325 335 345 355 365 375 385 395 δ13C (V-PDB) -2.0 -1.7 -2.2 -3.0 -3.8 -4.3 -4.3 -4.4 -4.2 -4.3 -4.2 -3.9 -3.7 -3.5 -3.1 -3.0 -3.0 -2.7 -2.5 -2.7 -2.9 -2.3 -1.9 -2.5 -2.1 -2.1 -2.7 -2.0 -2.0 -1.6 -1.1 -0.6 -0.5 -0.8 -0.8 -0.7 -0.3 -0.7 -1.2 -0.8 δ 18O (V-PDB) -0.6 0.0 1.6 2.0 1.0 0.5 0.3 -1.1 -0.5 -0.8 -1.1 -0.5 0.1 -0.7 0.4 0.0 0.0 -0.1 -0.1 0.5 0.6 0.7 1.3 0.6 1.0 0.7 0.2 0.6 0.3 0.8 0.2 1.4 1.7 1.3 0.9 1.5 1.4 0.5 0.3 0.7 207 Table A.22: (cont’d) Specimen Locality Position KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 KWS-NAIV-1108 Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish Naivasha - Crayfish 405 415 425 435 445 455 465 475 485 495 505 515 535 545 555 565 575 585 δ13C (V-PDB) -1.9 -1.1 -0.8 -0.7 -1.2 -0.9 -0.8 -0.6 -0.7 -1.1 -1.1 -0.9 -0.6 -0.8 -0.3 -0.8 -0.6 -1.0 δ 18O (V-PDB) 0.5 0.4 0.5 -0.1 -0.3 0.2 0.5 0.4 0.2 0.3 0.3 0.5 0.9 0.5 1.4 0.8 1.1 0.5 208 Table A.23: Lake Nakuru hippo isotope values. Specimen Locality Position KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini KWS-LNakuru-Kabutini Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru Nakuru 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 185 195 205 215 225 235 245 255 265 275 285 295 305 315 δ13C (V-PDB) -2.0 -0.6 -0.6 -0.6 -1.2 -2.2 -1.5 -1.4 -1.6 -2.0 -0.1 -0.1 -0.1 0.2 0.1 0.2 0.7 -0.2 0.0 0.2 -0.7 -0.7 -1.1 -1.0 -0.9 -0.9 -1.5 -1.6 -1.7 -2.2 -2.0 δ 18O (V-PDB) -0.5 0.5 1.7 1.8 1.1 0.5 0.7 0.0 -0.7 -2.9 -1.9 -1.1 -1.6 -1.5 -1.6 -1.7 -1.2 -2.5 -2.4 -1.5 -1.6 -2.0 -1.9 -1.9 -1.7 -2.1 -2.0 -2.2 -1.9 -2.3 -2.3 209 Table A.24: Lake Ol Bolossat hippo isotope values. Specimen Locality Position Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat Olbolossat 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 265 275 285 295 305 δ13C (V-PDB) -5.0 -5.7 -4.6 -4.9 -5.8 -6.3 -6.7 -5.0 -4.3 -2.0 -4.5 -5.6 -5.0 -4.8 -7.1 -7.3 -7.7 -6.5 -7.2 -7.9 -5.5 -6.2 -5.5 -6.0 -4.7 -3.8 -3.6 -4.1 -5.1 -3.7 -5.6 δ 18O (V-PDB) -2.1 -1.7 -2.4 -2.5 -0.8 -1.5 -3.2 -1.4 -0.8 -3.1 -0.7 -1.3 -0.8 -0.9 -1.1 -1.2 -2.0 -3.1 -3.5 -2.9 -3.6 -2.7 -2.5 -1.6 -2.4 -3.5 -4.5 -3.7 -4.0 -2.6 -2.3 210 Table A.25: Tsavo #1 hippo isotope values. Specimen Locality Position K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 275 285 295 305 315 325 335 345 345 355 355 365 365 375 δ13C (V-PDB) -7.4 -6.8 -5.6 -5.6 -6.3 -7.9 -8.5 -5.7 -4.5 -4.4 -3.7 -3.1 -3.0 -2.6 -3.1 -2.6 -2.3 -2.0 -1.7 -1.9 -2.1 -2.7 -3.4 -2.8 -2.1 -2.3 -2.8 -2.9 -2.5 -2.5 -2.5 -2.5 -2.5 -2.6 -2.5 -2.4 -2.3 -2.0 -1.9 -2.1 δ 18O (V-PDB) -0.9 -0.4 -1.9 -3.2 -3.8 -3.8 -3.9 -4.5 -4.4 -4.4 -4.4 -4.7 -4.2 -4.5 -4.8 -4.6 -4.1 -4.4 -4.6 -4.6 -4.7 -4.5 -4.0 -4.2 -4.0 -4.1 -4.5 -4.3 -4.4 -3.8 -4.0 -4.5 -4.9 -4.7 -4.3 -4.8 -4.7 -4.6 -4.4 -4.6 211 Table A.25: (cont’d) Specimen Locality Position K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 K01-TSW-291 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 Tsavo #1 375 385 385 395 405 415 425 435 445 455 465 475 485 495 505 515 525 535 545 555 565 575 585 595 605 δ13C (V-PDB) -1.8 -1.5 -1.5 -1.3 -1.6 -1.5 -1.8 -2.0 -2.2 -2.3 -1.9 -2.3 -2.3 -2.7 -3.2 -3.6 -3.7 -3.4 -3.1 -3.1 -3.4 -3.3 -2.8 -2.8 -2.3 δ 18O (V-PDB) -4.3 -4.3 -4.5 -4.2 -4.4 -4.0 -3.8 -4.1 -3.7 -4.0 -4.6 -4.4 -4.7 -4.9 -5.2 -4.9 -4.9 -5.0 -4.7 -5.0 -5.1 -4.7 -4.6 -4.7 -4.5 212 Table A.26: Tsavo #2 hippo isotope values. Specimen Locality Position Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 175 185 195 205 215 225 235 245 255 265 275 285 295 305 315 325 335 345 355 365 375 385 395 405 δ13C (V-PDB) -2.4 -3.5 -4.2 -4.2 -3.8 -3.3 -3.1 -3.3 -3.5 -3.3 -3.2 -3.4 -3.0 -2.8 -3.6 -3.6 -3.1 -3.3 -3.1 -3.5 -3.9 -4.5 -4.6 -4.6 -4.6 -3.3 -2.6 -3.7 -3.6 -3.1 -2.7 -2.5 -2.2 -2.2 -2.7 -2.8 -3.2 -3.5 -2.7 -2.6 δ 18O (V-PDB) -3.6 -3.3 -3.3 -3.5 -3.1 -4.1 -4.1 -4.5 -4.6 -4.6 -4.3 -4.7 -4.8 -4.3 -4.2 -4.6 -4.5 -4.5 -4.3 -4.7 -4.8 -4.3 -4.5 -5.0 -5.3 -5.3 -5.6 -5.6 -5.1 -4.8 -4.6 -4.6 -4.7 -4.7 -4.7 -4.9 -3.9 -4.2 -4.4 -4.2 213 Table A.26: (cont’d) Specimen Locality Position Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo-09 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 Tsavo #2 415 425 435 445 455 465 475 485 495 505 515 525 535 545 555 565 575 585 595 δ13C (V-PDB) -2.4 -2.6 -2.6 -2.1 -2.5 -3.3 -3.5 -3.5 -3.3 -3.5 -2.9 -2.3 -2.6 -2.5 -3.1 -3.5 -3.2 -3.2 -2.3 δ 18O (V-PDB) -4.1 -4.6 -4.2 -4.0 -4.1 -4.6 -4.5 -4.3 -4.1 -5.1 -4.3 -3.9 -3.5 -4.2 -4.5 -4.3 -3.9 -4.2 -3.4 214 Table A.27: Tsavo #3 hippo isotope values. Specimen Locality Position K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 265 275 285 295 305 315 325 335 345 355 365 375 385 395 δ13C (V-PDB) -2.8 -2.9 -2.9 -3.3 -3.7 -5.6 -5.5 -5.4 -4.7 -5.3 -5.7 -4.5 -4.5 -3.9 -3.6 -3.3 -2.7 -2.5 -2.9 -2.7 -2.3 -2.1 -2.1 -2.3 -2.6 -2.2 -2.2 -2.0 -1.7 -2.0 -2.4 -2.4 -2.4 -2.8 -4.4 -5.8 -7.3 -7.5 -7.8 -9.4 δ 18O (V-PDB) -3.3 -3.1 -1.4 -0.3 0.8 1.8 1.0 0.7 0.3 -1.6 -1.5 -1.2 -1.2 -1.6 -1.5 -1.7 -2.1 -1.9 -1.4 -1.3 -1.9 -2.2 -1.7 -0.8 -0.9 -1.2 -1.2 -1.1 -2.3 -2.4 -2.1 -1.5 -1.2 -1.6 -3.0 -2.8 -2.9 -3.0 -3.3 -2.9 215 Table A.27: (cont’d) Specimen Locality Position K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 K08-201 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 Tsavo #3 405 415 425 435 445 455 465 475 485 495 505 515 525 535 δ13C (V-PDB) -10.0 -9.7 -8.5 -8.9 -9.4 -9.4 -7.7 -7.2 -6.6 -6.3 -5.4 -4.8 -4.7 -6.2 δ 18O (V-PDB) -4.3 -3.3 -3.4 -4.4 -3.9 -4.3 -4.1 -4.2 -4.4 -4.6 -3.9 -3.8 -4.0 -4.2 216 Table A.28: Turkana – Koobi Fora 1 hippo isotope values. Specimen Locality Position K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 15 25 35 45 52 54 56 58 60 62 64 66 68 70 72 74 76 78 80 82 84 86 88 90 92 94 96 98 100 102 104 106 108 110 112 114 116 118 120 122 δ13C (V-PDB) 1.0 1.2 0.6 0.7 0.4 0.5 0.4 0.3 0.4 0.4 0.6 0.6 0.8 1.1 1.0 1.1 0.9 1.0 0.8 0.3 0.6 0.4 0.4 -0.1 0.3 0.1 0.2 0.1 0.1 0.3 0.3 0.4 0.4 0.4 0.5 0.6 0.6 0.7 0.7 1.0 δ 18O (V-PDB) 0.9 1.5 1.2 1.3 2.9 2.9 2.2 2.3 2.7 2.8 2.7 2.6 2.2 2.8 2.1 2.1 2.2 2.3 2.3 1.2 2.0 1.4 1.5 1.1 1.5 1.5 1.5 1.0 1.7 2.5 2.2 2.5 2.5 2.4 2.2 2.5 2.2 2.0 1.6 1.9 217 Table A.28: (cont’d) Specimen Locality Position K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 124 126 128 130 132 134 136 138 140 142 144 146 148 150 151 153 155 157 159 161 163 165 167 169 171 173 175 177 179 181 183 185 187 189 191 193 195 197 199 201 δ13C (V-PDB) 1.1 1.2 1.3 1.4 0.7 1.2 1.4 1.1 1.2 1.4 1.5 1.5 1.2 1.2 0.2 0.0 0.1 0.4 0.3 0.5 0.5 0.7 0.7 0.7 0.9 1.2 0.9 1.1 0.9 1.0 0.8 1.3 0.9 0.7 0.8 0.4 0.5 0.3 0.3 0.0 δ 18O (V-PDB) 2.0 1.8 1.7 1.4 2.2 2.3 2.2 1.8 1.8 2.0 2.0 2.1 2.4 2.1 0.6 0.9 0.7 1.1 1.3 1.0 1.2 1.3 1.4 1.7 1.5 1.2 0.9 1.1 0.6 0.7 0.0 0.4 -0.2 0.3 0.7 0.5 0.9 0.6 0.5 0.4 218 Table A.28: (cont’d) Specimen Locality Position K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 203 205 207 209 211 213 215 217 219 221 223 225 227 229 231 233 235 237 239 241 243 245 247 249 250 252 254 256 258 260 262 264 266 268 270 272 274 276 278 280 δ13C (V-PDB) 0.0 -0.1 0.2 0.0 0.1 0.3 0.0 -0.1 0.0 -0.1 -0.2 -0.4 -0.5 -0.7 -0.3 -0.1 0.1 0.1 0.2 0.1 0.2 0.3 0.3 0.2 1.1 1.0 1.1 1.0 1.0 0.9 0.7 0.9 0.9 0.8 0.9 0.6 0.9 1.0 1.1 1.0 δ 18O (V-PDB) 0.8 0.6 1.0 0.6 0.3 0.6 0.9 0.8 0.9 0.9 1.0 0.5 0.6 1.1 1.1 0.9 0.3 0.5 1.1 0.7 1.1 1.1 0.9 0.9 2.3 2.2 2.2 2.2 1.7 1.7 1.7 1.7 1.2 1.6 1.0 1.3 1.3 1.4 1.5 1.7 219 Table A.28: (cont’d) Specimen Locality Position K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 282 284 286 288 290 292 294 296 298 300 302 304 306 308 310 312 314 316 318 320 322 324 326 328 330 332 334 336 338 340 342 344 346 348 350 355 365 375 385 395 δ13C (V-PDB) 1.1 1.0 1.2 1.3 1.2 1.1 1.3 1.4 1.0 1.2 1.2 1.3 1.3 1.5 1.6 1.4 1.2 1.3 1.2 1.4 1.6 1.4 1.5 0.7 1.3 1.4 1.3 1.2 1.3 1.3 1.7 1.4 1.3 1.1 1.6 1.3 1.3 1.4 1.3 1.1 δ 18O (V-PDB) 1.5 1.4 1.8 1.9 2.0 2.0 2.2 2.7 1.0 1.7 1.4 1.3 1.4 1.8 1.9 1.6 1.4 1.3 0.9 1.1 1.1 0.8 0.7 0.3 0.7 0.9 0.8 1.3 1.1 0.9 1.5 1.7 1.0 1.8 1.6 0.0 1.1 1.0 1.2 1.1 220 Table A.28: (cont’d) Specimen Locality Position K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF K11-KF Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 Turkana - Koobi Fora 1 405 415 425 435 445 455 465 475 485 δ13C (V-PDB) 1.2 1.4 1.4 1.1 1.4 1.0 0.7 0.7 0.5 δ 18O (V-PDB) 0.8 2.0 1.2 0.6 0.9 1.2 0.9 2.2 2.4 221 Table A.29: Turkana – Koobi Fora 2 hippo isotope values. Specimen Locality Position KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 KEN-09-115 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 14 20 30 60 71 82 93 103 113 123 132 142 152 162 172 182 193 203 213 222 233 244 253 262 272 281 292 302 313 323 333 343 353 364 374 383 393 403 413 422 δ13C (V-PDB) 1.9 1.5 1.3 1.7 1.3 1.2 1.1 0.7 0.7 1.1 0.4 0.4 0.9 0.9 1.1 1.4 1.4 1.0 0.2 0.4 0.9 1.4 0.6 0.3 0.8 0.6 -0.4 -0.1 0.3 0.2 0.4 0.2 0.5 0.3 0.4 0.6 0.2 -0.3 -0.1 0.1 δ 18O (V-PDB) 0.5 1.4 0.5 0.7 1.4 1.2 1.0 1.4 1.2 1.0 1.3 1.4 1.1 1.7 1.5 2.0 1.1 1.5 0.9 1.3 1.0 1.1 1.3 1.2 1.5 1.9 1.4 1.4 1.7 1.8 1.7 1.1 1.3 1.2 1.1 1.1 1.1 1.2 0.8 1.0 222 Table A.29: (cont’d) Specimen Locality Position KEN-09-115 KEN-09-115 KEN-09-115 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 Turkana - Koobi Fora 2 432 443 453 δ13C (V-PDB) 0.2 0.2 -0.2 δ 18O (V-PDB) 0.6 0.6 1.1 223 Table A.30: Witu hippo isotope values. Specimen Locality Position Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 195 205 215 225 235 245 255 265 275 295 305 315 325 335 345 355 365 375 385 395 405 δ13C (V-PDB) -1.3 -0.9 -1.2 -1.5 -2.0 -1.8 -2.1 -2.0 -2.0 -2.1 -2.3 -2.1 -2.0 -2.2 -2.0 -2.1 -2.1 -2.4 -2.2 -2.2 -2.2 -2.1 -2.0 -2.1 -2.0 -2.1 -2.0 -2.1 -1.9 -2.2 -2.1 -2.2 -2.3 -2.5 -2.3 -2.2 -2.5 -1.3 -1.3 -2.2 δ 18O (V-PDB) -1.7 -1.1 -0.8 -0.6 -0.6 -0.9 -1.2 -1.5 -1.1 -1.2 -1.8 -1.3 -1.4 -1.7 -1.5 -2.0 -2.4 -2.5 -2.4 -2.4 -2.5 -2.7 -2.8 -2.8 -2.8 -2.8 -2.7 -2.5 -2.5 -3.0 -2.9 -3.2 -3.1 -3.3 -3.1 -3.1 -3.5 -3.0 -3.0 -3.3 224 Table A.30: (cont’d) Specimen Locality Position Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu Witu 415 425 435 445 455 465 475 485 495 505 515 525 535 545 555 565 δ13C (V-PDB) -0.8 -1.4 -1.7 -1.4 -1.0 -2.0 -1.7 -1.3 -1.6 -1.1 -0.9 -1.4 -1.7 -1.7 -2.0 -2.1 δ 18O (V-PDB) -3.0 -3.2 -2.7 -2.4 -2.8 -2.7 -2.5 -2.7 -2.8 -3.0 -3.2 -3.1 -3.5 -3.5 -3.6 -3.8 225 Table A.31: Bonferroni corrected P-values, Wilcoxan ranksum test (bold = significant). Locality, Isotope Bonferroni corrected P-values Naivasha, C 1.000 Naivasha, O 1.000 AdhiDam, C 1.000 AdhiDam, O 0.162 Turkana09, C 0.122 Turkana09, O 1.000 Amboseli0401,C 1.000 Amboseli0401,O 1.000 Tsavo07, C 1.000 Tsavo07, O 1.000 Tsavo96, C 0.002 Tsavo96, O 0.002 AS167, C 1.000 AS167, O 0.139 Minjila, C 1.000 Minjila, O 0.169 Nakuru, C 1.000 Nakuru, O 0.006 Mwea09, C 0.002 Mwea09, O 0.009 MweaNR, C 0.002 MweaNR, O 0.950 Mwea05, C 1.000 Mwea05, O 0.169 MweaGitaru, C 1.000 MweaGitaru, O 0.182 NaivashaH, C 0.317 NaivashaH, O 0.193 AmboseliH, C 1.000 AmboseliH, O 0.577 AS08, C 1.000 AS08, O 1.000 Chyulu, C 1.000 Chyulu, O 0.005 Kisumu, C 0.014 Kisumu, O 1.000 Mara, C 1.000 Mara, O 1.000 Meru, C 0.000 Meru, O 0.032 226 Table A.31: (cont’d) Locality Mokowe, C Mokowe, O Mpeketoni, C Mpeketoni, O Olbolossat, C Olbolossat, O Witu, C Witu, O AS168, C AS168, O AS166, C AS166, O Koobi Fora, C Koobi Fora, O Buffalo Springs, C Buffalo Springs, O Aberdares, C Aberdares, O Bonferroni corrected P-values 1.000 1.000 0.162 0.007 1.000 1.000 1.000 0.003 0.008 1.000 0.154 1.000 1.000 0.376 0.517 0.002 0.010 0.014 APPENDIX B RAW ISOTOPE DATA FROM QUEEN ELIZABETH NATIONAL PARK HIPPOS 228 Table B.1: Raw isotope values of QEP hippos through time. Length (mm from δ13C Specimen Cal. Year prox. end) (V-PDB) KL 1970.59 5 -0.2 KL 1970.29 45 0.0 KL 1969.99 75 0.6 KL 1969.69 95 0.7 KL 1969.39 105 0.9 KL 1969.09 115 0.7 KL 1968.79 125 0.7 KL 1968.49 135 0.8 KL 1968.19 145 -0.4 KL 1967.89 155 0.3 KL 1967.59 165 -0.5 KL 1967.29 175 -0.8 KL 1966.99 185 -1.1 KL 1966.69 195 0.5 KL 1966.39 205 -0.7 KL 1966.09 215 -1.0 KL 1965.79 225 -0.2 KL 1965.49 235 -0.2 KL 1965.19 245 -0.1 KL 1964.89 255 -0.1 KL 1964.59 265 -0.7 KL 1964.29 275 -0.1 KL 1963.99 285 0.5 KL 1963.69 295 -0.1 KL 1963.39 305 0.0 KL 1963.09 315 -0.2 KL 1962.79 325 0.0 KL 1962.49 335 0.2 KL 1962.19 345 0.3 KL 1961.89 355 -0.2 KL 1961.59 365 -0.3 KL 1961.29 375 -0.1 KL 1960.99 385 -0.2 Queen VIC 1982.87 15 -3.5 Queen VIC 1983.10 25 -3.0 Queen VIC 1983.32 45 -3.0 Queen VIC 1983.55 65 -3.8 Queen VIC 1983.77 75 -3.2 Queen VIC 1984.00 85 -3.3 Queen VIC 1984.23 95 -4.0 δ 18O (V-PDB) 0.6 -0.5 0.5 -0.8 -0.8 -2.6 -1.1 -0.7 -0.4 -0.6 0.3 -0.3 -0.6 0.3 -0.1 -1.0 0.5 0.4 0.5 0.9 0.3 0.6 1.4 0.4 0.3 0.4 0.3 0.4 -0.5 -0.4 -0.3 1.1 1.3 -1.9 0.6 -0.5 -1.7 -2.2 -2.1 -1.0 %C4 85 85 90 90 95 90 90 95 85 90 85 80 80 90 82 80 85 85 85 85 80 85 90 85 85 85 85 90 90 85 85 85 85 60 65 65 60 65 65 60 229 Table B.1: (cont’d) Specimen Cal. Year Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Queen VIC Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL 1984.45 1984.68 1985.13 1985.58 1985.81 1986.03 1986.26 1986.71 1986.94 1987.39 1987.61 1987.84 1988.06 1988.29 1988.52 1988.74 1988.97 1989.19 1989.42 1989.65 1989.87 2000.00 1999.77 1999.55 1999.32 1999.10 1998.87 1998.65 1998.42 1998.19 1997.97 1997.74 1997.52 1997.29 1997.06 1996.84 1996.61 1996.39 1996.16 1995.94 Length (mm from prox. end) 105 115 135 155 165 175 185 205 225 245 255 265 275 285 295 305 315 325 335 345 355 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185 δ13C (V-PDB) -3.2 -1.6 -3.6 -3.0 -2.8 -3.3 -3.2 -3.5 -3.0 -3.3 -3.0 -3.3 -3.3 -3.5 -4.0 -4.0 -3.5 -3.4 -4.1 -3.6 -3.6 -2.7 -2.3 -3.0 -3.4 -3.2 -3.1 -3.6 -3.4 -3.2 -3.3 -3.1 -3.0 -2.7 -2.8 -2.9 -2.7 -2.5 -2.4 -2.3 δ 18O (V-PDB) -1.2 -0.9 -0.4 -1.7 -1.2 -0.8 -0.9 -0.8 -0.6 -0.1 0.1 -0.7 -1.0 -1.7 -1.9 -1.6 -0.8 -0.7 -1.5 0.2 -0.9 1.2 1.0 0.4 -0.5 -0.4 -0.1 -0.3 -1.8 -1.5 -2.5 -2.4 -3.5 -4.1 -4.2 -4.9 -5.6 -4.3 -2.8 -3.3 %C4 65 75 60 65 65 65 65 65 65 65 65 65 56 60 60 60 65 65 60 60 60 70 70 65 65 65 65 60 65 65 65 65 65 70 65 65 70 70 70 70 230 Table B.1: (cont’d) Specimen Cal. Year Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL 1995.71 1995.48 1995.26 1995.03 1994.81 1994.58 1994.36 1994.13 1993.9 1993.68 1993.45 1993.23 1993.00 1992.77 1992.55 1992.32 1992.1 1991.87 1991.65 1991.42 1991.19 1990.97 1990.74 1990.52 1990.29 1990.06 1989.84 1989.61 1989.39 1989.16 1988.94 1988.71 1988.48 1988.26 1988.03 1987.81 1987.58 1987.36 1987.13 1986.9 Length (mm from prox. end) 195 205 215 225 235 245 255 265 275 285 295 305 315 325 335 345 355 365 375 385 395 405 415 425 435 445 455 465 475 485 495 505 515 525 535 545 555 565 575 585 δ13C (V-PDB) -2.6 -2.5 -2.5 -2.4 -2.4 -2.4 -1.9 -2.9 -2.8 -2.1 -2.5 -2.5 -2.6 -2.7 -2.6 -2.5 -2.3 -2.6 -2.5 -2.6 -2.8 -2.8 -2.8 -3.2 -3.4 -3.4 -4.1 -4.0 -4.4 -4.3 -3.9 -3.9 -4.0 -3.4 -3.2 -3.2 -3.0 -3.0 -2.8 -2.6 δ 18O (V-PDB) -2.8 -2.2 -2.1 -1.8 -2.3 -2.0 -2.9 -1.3 -2.5 -2.1 -1.7 -2.5 -2.3 -2.5 -2.4 -2.0 -1.0 -2.2 -1.7 -0.9 -1.5 -2.0 -1.4 -1.9 -1.7 -1.4 -1.9 -1.7 -1.7 -1.8 -1.2 -1.4 -1.3 -1.7 -2.2 -2.5 -1.2 -1.7 -2.4 -1.7 %C4 70 70 70 70 70 70 75 65 65 70 70 70 70 70 70 70 70 70 70 70 65 65 70 65 65 65 60 60 55 55 60 60 60 65 65 65 65 65 65 70 231 Table B.1: (cont’d) Specimen Cal. Year Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL Q-09-KL 1986.68 1986.45 1986.23 1986.00 1985.77 1985.55 1985.32 1985.10 Length (mm from prox. end) 595 605 615 625 635 645 655 665 δ13C (V-PDB) -2.9 -3.0 -3.0 -3.1 -3.1 -3.0 -2.9 -2.9 δ 18O (V-PDB) -1.5 -1.6 -1.7 -2.3 -1.8 -1.5 -1.5 -2.0 %C4 65 65 65 65 65 65 65 65 APPENDIX C RAW ISOTOPE VALUES OF ARCHAEOLOGICAL TOOTH ENAMEL AND LEAF WAX BIOMARKERS 233 Table C.1: Isotope values of Luanda (9.7 – 8.5 Kya) fauna. δ13C δ 18O Specimen KLC ID (V-PDB) (V-PDB) Alcelaphini Alcelaphini Bovini BVD 3 BVD 4 BVD 4 BVD 4 BVD 4 BVD 4 BVD 4 Connochaetes taurinus Connochaetes taurinus Connochaetes taurinus Connochaetes taurinus Connochaetes taurinus Connochaetes taurinus Connochaetes taurinus Equus spp. H. amphibius H. amphibius H. amphibius H. amphibius H. amphibius H. amphibius Kobus defassa Kobus defassa Kobus defassa Kobus kob Kobus kob Kobus kob Potamochoerus spp. P. aethiopicus P. aethiopicus Potamochoerus spp. Potamochoerus spp. Redunca spp. Redunca spp. Syncerus caffer Syncerus caffer Syncerus caffer 3.1 2.4 3.2 1.4 1.7 1.2 0.5 1.7 1.3 3.6 -0.1 2.6 1.6 3.4 1.6 2.6 3.6 0.6 -1.9 -3.4 -5.5 -5.7 -3.2 -5.5 2.4 1.5 3.0 1.8 3.0 1.3 -12.0 1.1 0.2 -7.8 -11.8 0.3 3.0 0.7 -1.4 0.1 0.3 -2.8 -1.2 -0.2 -2.0 -3.4 2.3 -1.0 -1.5 -0.8 0.4 -0.3 -0.7 0.2 0.0 0.6 -0.5 -0.6 -3.1 -3.0 -2.3 -4.1 -2.8 -4.6 -4.7 -1.2 -0.1 -1.0 0.0 -1.4 -5.2 1.4 -0.6 -4.2 -5.5 0.0 -1.5 0.5 0.3 -0.6 KKC13-E-191 KKC13-E-178 KKC13-E-190 KKC13-E-189 KKC13-E-151 KKC13-E-154 KKC13-E-157 KKC13-E-166 KKC13-E-174 KKC13-E-205 KKC13-E-158 KKC13-E-164 KKC13-E-171 KKC13-E-193 KKC13-E-196 KKC13-E-200 KKC13-E-201 KKC13-E-152 KKC13-E-155 KKC13-E-167 KKC13-E-172 KKC13-E-173 KKC13-E-185 KKC13-E-188 KKC13-E-176 KKC13-E-202 KKC13-E-203 KKC13-E-148 KKC13-E-149 KKC13-E-150 KKC13-E-181 KKC13-E-198 KKC13-E-206 KKC13-E-179 KKC13-E-180 KKC13-E-147 KKC13-E-192 KKC13-E-153 KKC13-E-194 KKC13-E-159 NMK ID Tooth L/691/BS/3 L/583/FS/7 L/691/OB/0 L/691/OB/0 L/414/BR/6 L/414/OB/1 L369/BS/2 L/256/OB/1 L/828/BH/7 L/828/BH/8 L/369/BS/3 L/256/OB/1 L/828/OB/1 L/533/OB/2 L/607/BS2/4 L/639/BS/3 L/639/GM L/414/BS/5 L/369/BS/3 L/369/OB/0 L/828/BH/7 L/828/BH/7 L/583/GM/8 L/583/GM/8 L/533/BS/5 L/828/BH/8 L/828/BH/8 L/414/OB/1 L/583/GM/8 L/828/BH/8 L/583/FS/7 L/639/OB/0 L/828/BH/8 L/583/FS/7 L/583/FS/7 L/414/OB/1 L/756/OB/0 L/414/OB/1 L/756/BS/3 L/369/OB/1 LI2 frag LI M0 frag frag P4 frag frag frag M1 I1 M2 P3 LM0 LM1 UM3 LP3/4 UM3 Um2 frag tusk frag frag M0 LM1 LM0 M3 M2 M1,2? frag LM3 LM3 LM3 M2 UM3 UM3 M0 LM0 M2 234 Table C.1: (cont’d) Specimen δ13C (V-PDB) δ 18O (V-PDB) KLC ID NMK ID Tooth Syncerus caffer Syncerus caffer Syncerus caffer Syncerus caffer Syncerus caffer Syncerus caffer Syncerus caffer Syncerus caffer Syncerus caffer Syncerus caffer Syncerus caffer Syncerus caffer Syncerus caffer Syncerus caffer 0.5 1.5 1.9 1.9 2.1 2.3 2.9 3.0 3.1 3.2 3.4 3.7 3.7 3.8 -0.7 -3.3 -1.7 -0.1 -0.9 -2.1 -1.3 -2.5 -2.3 -2.8 -1.8 -1.6 -3.2 -1.9 KKC13-E-170 KKC13-E-177 KKC13-E-163 KKC13-E-175 KKC13-E-187 KKC13-E-160 KKC13-E-168 KKC13-E-197 KKC13-E-161 KKC13-E-169 KKC13-E-199 KKC13-E-184 KKC13-E-162 KKC13-E-204 L/828/OB/0 L/533/BS/5 L/256/OB/1 L/533/OB/3 L/583/OB/2 L/369/BS/3 L/828/OB/0 L/607/GS/7 L/256/OB/1 L/828/OB/0 L/639/OB/0 L/583/FS/7 L/256/OB/1 L/828/BH/8 M0 I1 M3 M2 M3 M3 M0 M3 M0 M3 frag LP2 M0 P0 235 Table C.2: Isotope values of Wadh Lang’o (~3 Kya) fauna. δ13C δ 18O Specimen (V-PDB) (V-PDB) KLC ID BVD 2 2.2 -2.4 KKC13-E-240 Caprini 2.7 1.1 KKC13-E-245 Caprini -0.1 0.1 KKC13-E-254 Connochaetes taurinus 2.8 1.9 KKC13-E-244 Kobus defassa 1.4 2.6 KKC13-E-243 Phacochoerus 0.4 -0.1 KKC13-E-257 aethiopicus NMK ID 208 280 2945 281 181 2944 Tooth UM1 LM3 LM1/2 UM0 LM0 frags 236 Table C.3: Isotope values of Wadh Lang’o (~2 Kya) fauna. δ13C δ 18O Specimen KLC ID (V-PDB) (V-PDB) Alcelaphini 3.0 5.2 KKC13-E-215 Alcelaphini 0.0 4.0 KKC13-E-213 Alcelaphini 2.7 4.1 KKC13-E-214 Alcelaphini 2.2 2.8 KKC13-E-216 Alcelaphini 3.0 3.6 KKC13-E-217 Alcelaphini 1.8 1.3 KKC13-E-218 Bos taurus 1.8 3.6 KKC13-E-238 BVD 0 1.7 -0.5 KKC13-E-227 BVD 0 -2.3 2.1 KKC13-E-228 BVD 2 2.2 5.7 KKC13-E-251 BVD 2 -1.8 0.2 KKC13-E-255 BVD 2 0.6 1.2 KKC13-E-256 BVD 2 2.0 2.5 KKC13-E-262 BVD 2 0.3 3.1 KKC13-E-263 BVD 2-3 -0.6 4.7 KKC13-E-253 BVD 3-4 -0.7 2.2 KKC13-E-242 Caprini 0.1 1.6 KKC13-E-237 Caprini 3.1 4.7 KKC13-E-247 Caprini 1.6 4.1 KKC13-E-252 Caprini 2.8 5.4 KKC13-E-208 Caprini 4.0 5.0 KKC13-E-209 Caprini 1.1 2.6 KKC13-E-210 Caprini 2.2 -2.4 KKC13-E-219 Caprini -0.3 1.7 KKC13-E-220 Caprini 2.2 -0.3 KKC13-E-222 Caprini -0.2 2.0 KKC13-E-223 Caprini -1.4 0.1 KKC13-E-230 Caprini 1.7 3.9 KKC13-E-241 Caprini -0.5 4.3 KKC13-E-249 Caprini 2.1 4.1 KKC13-E-250 Caprini 2.0 3.7 KKC13-E-258 Caprini -0.7 2.1 KKC13-E-260 Caprini -2.2 4.2 KKC13-E-235 Redunca -0.7 1.7 KKC13-E-221 redunca Redunca -3.3 -1.6 KKC13-E-224 redunca Redunca 2.6 0.7 KKC13-E-225 redunca Reduncini 2.5 0.4 KKC13-E-226 NMK ID Tooth 2916 2929 2915 2921 2909a 2909b 4448 2914 16626-28 2870 2894 5246 n/a n/a 2939 1612 2886 2878 1702 2926 2928 2927 2908 2911 2910 2924 4537 1615a 2871 2876 5096 2888 2865 UM3 UP4 UP4 frag M3? M2 M2 frag frag Lm3 ind UM3 frag frag frag M3 M UM3 LM3 LM2 M3 DUP4 UM2 UM2 LM2 UM3 UM3 M3 M2 M0 M1 LM12 P4 2912 UM2 2918 UM0 2923 UM0 2913 UM0 237 Table C.4: Lothagam faunal isotopes (8.3 Kya). δ13C δ 18O Specimen Site (V-PDB) (V-PDB) "Equid Alcelaphini 0.5 3.0 site" Alcelaphini 2.0 3.2 FxJj12 Alcelaphini 1.4 -1.8 FxJj12 Bovidae -0.9 2.1 "Equid site" Bovidae 2.7 2.5 GaJj11 Equidae -1.1 1.7 Equidae -2.0 1.1 Equidae -1.9 -2.5 Equidae 0.6 1.2 Equidae 0.6 5.8 Equidae -0.3 2.3 Equidae 0.0 2.4 H. amphibius H. amphibius -2.1 -2.0 GaJj11 0.1 -2.3 GaJj11 Reduncini 2.0 2.8 Suidae 0.3 3.1 Suidae 0.0 3.0 "Equid site" "Equid site" "Equid site" "Equid site" "Equid site" "Equid site" "Equid site" "Equid site" "Equid site" "Equid site" Age KLC ID 10.5 KC12-9AA 9.58.3 9.58.3 KKC1016F KKC1011 10.5 KC12-2T 10.29.9 KKC13E-1 KC1211AX KC1213Y 10.5 10.5 NMK ID Tooth M "area P1/P2? 116" FxJj12M2 46 M 73 M3 M M 10.5 KC12-14AE M 10.5 KC12-2S M 10.5 KC12-7N M 10.5 KC12-7O M 10.5 KC12-7P M 10.29.9 10.29.9 KKC1141 KKC13E-2 KC1210AZ KC1211AW 10.5 10.5 10.5 KC12-6AB 188 C 199 frag frag M M 238 Table C.5: Later Holocene faunal isotopes (8.3 Kya); isotopes relative to V-PDB. ẟ13C ẟ18O Aepycerotini Alcelaphini Alcelaphini Alcelaphini Alcelaphini -7.2 -5.4 -1.6 -1.4 -1.3 Alcelaphini Alcelaphini Antilopini Antilopini Antilopini Specimen Site Age KLC ID 2.3 3.9 3.4 2.4 4.5 GaJj2 GaJj2 Dongodien GaJj2 Dongodien 4.9-4.4 4.9-4.4 4.5-3.4 4.9-4.4 4.5-3.4 KKC13-E-23 KKC13-E-31 KKC13-E-6 KKC13-E-28 KKC11-6 0.3 3.2 -4.5 -3.3 -3.0 2.8 0.5 0.6 3.8 4.1 Dongodien GaJj2 Dongodien Dongodien Dongodien 4.5-3.4 4.9-4.4 4.5-3.4 4.5-3.4 4.5-3.4 KKC13-E-8 KKC13-E-29 KKC13-E-48 KKC13-E-35 KKC11-6F Antilopini Bovini Caprini Caprini Caprini Caprini Caprini -2.2 1.6 -7.9 -7.5 -6.8 -6.0 -5.9 4.0 3.6 3.2 1.5 5.2 1.7 4.4 Dongodien Dongodien Dongodien Dongodien Dongodien GaJj2 Dongodien 4.5-3.4 4.5-3.4 4.5-3.4 4.5-3.4 4.5-3.4 4.9-4.4 4.5-3.4 KKC13-E-5 KKC13-E-52 KKC13-E-4 KKC13-E-39 KKC13-E-40 KKC13-E-33 KKC10-22F Caprini Caprini Caprini Caprini Caprini -5.5 -5.4 -5.3 -4.8 -4.5 4.7 2.4 4.8 5.1 6.0 Dongodien Dongodien Dongodien Dongodien Dongodien 4.5-3.4 4.5-3.4 4.5-3.4 4.5-3.4 4.5-3.4 KKC13-E-42 KKC13-E-51 KKC13-E-36 KKC13-E-41 KKC10-20F Caprini Caprini Caprini Caprini -3.7 -3.4 -3.1 -2.9 2.1 4.9 4.4 6.7 Dongodien Dongodien Dongodien Dongodien 4.5-3.4 4.5-3.4 4.5-3.4 4.5-3.4 KKC13-E-10 KKC13-E-37 KKC13-E-45 KKC10-18F Caprini Caprini Caprini -2.3 -2.1 -1.6 4.6 5.1 4.6 Dongodien Dongodien Dongodien 4.5-3.4 4.5-3.4 4.5-3.4 KKC13-E-38 KKC13-E-49 KKC11-2 Caprini Caprini Caprini Equidae -1.5 0.0 0.1 -0.2 5.8 4.1 3.6 0.6 Dongodien Dongodien Dongodien Dongodien 4.5-3.4 4.5-3.4 4.5-3.4 4.5-3.4 KKC13-E-46 KKC13-E-43 KKC13-E-47 KKC13-E-55 NMK ID 187 110 357 122 GaJi427 1089 114 214 103 GaJi4430 306 1013 329 951 601 112 GaJi4212 822 240 822 864 GaJi41013 61 836 478 GaJi4573 1029 1193 GaJi4378 614 1029 272 104149 Tooth M M M M M2 M frag M M3 frag M3 M2 M P2 M M2 I M M M3 frag M2 M I M2 M3 M3 M2 M I M2 M2 M M jaw P2 239 Table C.5: (cont’d) Specimen ẟ13C ẟ18O Site Age KLC ID NMK ID Tooth H. amphibius H. amphibius H. amphibius H. amphibius indet indet indet indet indet indet indet indet indet Neotragini Neotragini Neotragini Suidae Suidae Suidae Suidae Suidae -4.1 -0.6 GaJj2 4.9-4.4 KKC13-E-19 80 M frag -1.2 0.5 GaJj2 4.9-4.4 KKC13-E-34 176 M frag -0.9 -1.4 GaJj2 4.9-4.4 KKC13-E-27 62 frag 0 0.4 Dongodien 4.5-3.4 KKC11-7 GaJi4-422 C serial -6 -4.4 -4.2 -2.3 -1.9 -1.4 -0.5 1.3 1.9 -6.5 -1.9 1.7 -2.4 -1.2 -0.2 0.4 2 5.3 2.2 3.5 5 1.4 1.8 1.9 4.1 3.8 2.6 -0.3 2 -1.4 0.6 1.7 1 1.2 Dongodien GaJj2 Dongodien Dongodien GaJj2 Dongodien Dongodien GaJj2 GaJj2 GaJj2 GaJj2 GaJj2 GaJj2 GaJj2 GaJj2 FxJj12N GaJj2 4.5-3.4 4.9-4.4 4.5-3.4 4.5-3.4 4.9-4.4 4.5-3.4 4.5-3.4 4.9-4.4 4.9-4.4 4.9-4.4 4.9-4.4 4.9-4.4 4.9-4.4 4.9-4.4 4.9-4.4 3.8-3.0 4.9-4.4 KKC10-19F KKC13-E-24 KKC11-5 KKC13-E-50 KKC13-E-18 KKC11-1 KKC13-E-9 KKC13-E-16 KKC13-E-22 KKC13-E-21 KKC13-E-30 KKC13-E-25 KKC11-38 KKC13-E-32 KKC11-37 KKC11-40 KKC13-E-26 GaJi4-1029 238 GaJi4-891 914 109 GaJi4-763 338 132 8 188 111 25 GaJj2-110 186 GaJj2-35 FxJj12-152 175 M M M PM4 M M M frag M frag M frag M frag M3 M3 C ? M M tusk 240 Table C.6: EBA, Horizon C faunal isotope values. δ13C δ 18O Specimen KLC ID (V-PDB) (V-PDB) Antilopini -7.8 1.4 KKC13-E-113 Antilopini -3.8 2.7 KKC13-E-112 Antilopini -3.3 3.7 KKC13-E-115 Bovidae 1.5 2.7 KKC13-E-109 Hippotragini 2.4 4.1 KKC13-E-108 Tragelaphini -8.9 5.6 KKC13-E-104 Tragelaphini -3.7 1.5 KKC13-E-105 Tragelaphini 1.5 1.1 KKC13-E-107 NMK ID Tooth 100367 100803 100622 101012 102010 100897 100375 102014 frag M2 M1 M frag M0 M0 M0 M0 241 Table C.7: EBA, Horizon B faunal isotopes. δ13C δ 18O Specimen KLC ID NMK ID (V-PDB) (V-PDB) Antilopini 0.2 1.6 KKC13-E-63 104107 Antilopini -0.2 2.3 KKC13-E-76 102873 Antilopini -0.8 3.6 KKC13-E-64 103339 Antilopini -0.8 5.3 KKC13-E-65 102974 Antilopini -1.4 4.2 KKC13-E-66 104070 Antilopini -1.7 2.8 KKC13-E-70 104177 Antilopini -3.7 1.8 KKC13-E-68 103256 Antilopini -5.8 -2.8 KKC13-E-71 200001 Antilopini -6.5 2.5 KKC13-E-72 104121 Antilopini -7.3 5.0 KKC13-E-69 103372 Antilopini -7.6 1.5 KKC13-E-75 104068 Antilopini -9.1 0.7 KKC13-E-74 103301 Antilopini -9.3 5.5 KKC13-E-67 104062 Cephalophini -2.1 1.8 KKC13-E-83 104172 Equidae 0.9 1.1 KKC13-E-56 104106 Equidae 0.1 1.3 KKC13-E-60 104093 Giraffidae -12.4 5.4 KKC13-E-61 101919 Hippotragini 4.0 2.9 KKC13-E-96 104160 Hippotragini 3.4 -0.8 KKC13-E-92 102887 Hippotragini 2.7 0.6 KKC13-E-94 101650 Hippotragini 2.6 0.2 KKC13-E-93 104148 Hippotragini 2.2 2.2 KKC13-E-90 104263 Hippotragini 2.1 1.6 KKC13-E-88 104363 Hippotragini -0.7 3.2 KKC13-E-95 103300 Hippotragini -4.7 2.8 KKC13-E-89 104072 Neotragini -11.8 1.6 KKC13-E-86 103252 Tragelaphini -9.0 5.1 KKC13-E-77 103254 Tragelaphini -10.1 6.3 KKC13-E-78 104264 Tragelaphini -11.5 3.2 KKC13-E-80 102100 Tragelaphini -12.2 7.0 KKC13-E-79 104079 Tooth M3 M3 M2 M1 M3 M2 M1 M2 M3 M3 M3 M3 M3 M0 M3 frag frag M0 frag P2 P2 M3 frag M0 M0 M1 M0 M0 M0 P0 242 Table C.8: EBA Horizons A1 and A2 isotope values. Specimen δ13C δ 18O KLC ID (V-PDB) (V-PDB) 3.3 2.7 Antilopini KKC13-E-101 1.3 -0.1 Hippotragini KKC13-E-91 -1.2 1.0 Antilopini KKC13-E-98 -1.5 3.3 Antilopini KKC13-E-100 -2.5 -0.3 Antilopini KKC13-E-103 -3.8 -4.3 Antilopini KKC13-E-102 -5.5 3.5 Antilopini KKC13-E-97 -7.9 1.4 Neotragini KKC13-E-84 -8.7 1.8 Antilopini KKC13-E-99 -11.5 3.6 Neotragini KKC13-E-87 -12.6 0.1 Neotragini KKC13-E-85 NMK ID 100198 102763 101530 100142 101014 100850 104203 102879 101506 102479 104162 Tooth M2 M1 M0 M3 P4 P3 P4 P2 M0 P4 M2 243 Table C.9: C28 FAME leaf wax biomarker values. %C4 calcuated using endmember values from -22 to -34‰, rounded to the nearest 5% to account for uncertatinty (Castañeda et al., 2009; Magill et al., 2013a). ẟ13C - C28 Depth in %C4 Section (V-PDB) -28.4 7.0 0.50 -27.7 7.5 0.55 -26.3 7.9 0.65 -27.1 8.3 0.55 -25.4 8.5 0.70 -28.1 8.9 0.50 -26.2 10.3 0.65 -25.8 10.7 0.65 -26.1 14.0 0.65 -28.5 14.4 0.45 -28.5 14.9 0.45 -24.9 15.9 0.75 -25.1 17.0 0.70 -27.5 17.5 0.55 -28.5 18.2 0.45 -24.9 18.8 0.75