Evolution of the Cetratiodactyla: Phylogeny, Life History and Conservation

I - II II. Dedication III - IV III. Introduction V- XV IV. Chapter 1 A complete phylogeny of the whales, dolphins and even-toed hoofed mammals (Cetartiodactyla). (1) Title page 1 (2) Abstract 2 (3) Table of contents 3 (4) Introduction 4 - 8 (5) Methodology 8 - 15 (6) Results and Discussion 15 - 24 (7) Conclusion 24 - 26 (8) Acknowledgements 26 - 27 (9) References 27 – 52 (10) Figure text 53 - 54 (11) Figures i. Figure 1 55 ii. Figure 2 56 iii. Figure 3 57 iv. Figure 4 58 v. Figure 5 59 vi. Figure 6 60 vii. Figure 7 61 (12) Appendix 62 – 73 V. Chapter 2 The roles of phylogeny and allometry in cetacean life histories (1) Title page 74 (2) Abstract 75 (3) Introduction 76 - 80 (4) Methods 80 - 82 (5) Results 83 - 84 (6) Discussion 85 - 90 (7) Conclusion 90 - 92 (8) Acknowledgements 92 (9) References 92 - 99 (10) Tables i. Table 1 100 - 101 ii. Table 2 102 - 103 iii. Table 3 104 - 105 (11) Figure text 106 (12) Figures i. Figure 1 107 ii. Figure 2 108 iii. Figure 3 109 (12) Appendix 110– 114 VI. Chapter 3 Bushmeat hunting, habitat loss and global extinction of the Artiodactyla (1) Title page 115 (2) Abstract 116 (3) Introduction 116 - 120 (4) Methods 120 - 124 (5) Results 124 - 126 (6) Discussion and Conclusion 126 - 131 (7) Acknowledgements 131 (8) References 131 - 139 (9) Tables i. Table 1 140 ii. Table 2 141 iii. Table 3 142 iv. Table 4 143 (10) Appendices i. Appendix 1 144- 157 ii. Appendix 2 158 - 160 Evolution of the Cetartiodactyla; Phylogeny, Life history and Conservation I Abstract Using the supertree approach of matrix representation with parsimony I present the first phylogeny to include all 290 extant species of the Cetartiodactyla (even-toed hoofed mammals and whales). At the family-level the supertree is fully resolved and supports the hypothesis that the whales and dolphins are nested within the even-toed hoofed mammals as sister-taxon to the hippopotamidae. I use the phylogeny as a statistical framework to control for the effects of shared evolutionary history on cross-species analyses of cetacean life history strategies and artiodactyl extinction risk. Cetacea are a clade of charastimatic mammals but their biology is poorly known; cetaceans are difficult to study due to the inaccessible nature of their habitats, their large size and long lives. Understanding the interrelationships among life history traits is not only important for the better understanding of cetacean biology and ecology but it is also the first step to understanding the action of natural selection upon those traits. Cetaceans follow general mammalian life history strategies with some interesting deviations; none of the cetacean reproductive traits are phylogenetically patterned and the amount of variance explained by body size allometry in those traits is exceedingly low. With over half of all artiodactyl species are threatened with extinction it is important to understand the biological processes underlying species extinction. I present a II multivariate model to assess for the first time which intrinsic (biological) and extrinsic (anthropogenic and environmental) factors influence variation in extinction risk. Species hunted for bushmeat in Africa are more susceptible to extinction via hunting if they have slower reproductive rates. In contrast, species primarily affected by habitat loss are vulnerable to extinction if they are rare as reflected by small geographic ranges or low population densities. III In dedication to my grandmother Florence May This dissertation is dedicated to the many people that have helped me throughout my four years at UVA. I would like to thank my advisor John Gittleman for all he has done, and especially for allowing me to develop my scientific method through making my own mistakes. At one time or other the Gittleman lab has included Kate Jones, Kim Dodd, Wes Sechrest, Mike Habib, Angie Skeeles, Rich Grenyer and Jonathan Davies, all of whom have been of great help both academically and as good friends. I don't think I could have got through my PhD without the friends that I have made here, especially Jessica who has been the most wonderful housemate; always willing to listen to my rants and to make me cookies whenever needed. The Sunday night dinner crew and the cabin trip/kayaking people have added many great memories and some extraordinary photos to my collection, thanks to Hugh, Jess, Dexter, Fred, Anne, Jackie, Steven, Camille, Stacey, Brian, Rich and Louise. Amy and Kim as part of the 'dissertation/thesis lunch' I have you to thank for keeping my sanity during the last 6 months. I would also like to thank Linda who helped me to transition into American life when I first arrived from England and for her continued friendship. Finally I could not have done anything without the continued love and support of my parents and my grandmother. My parents have always encouraged me in my interest in natural history and biology and what's more they believed me when at the age of 12 IIV announced I was going to get a PhD one day, since then they have tirelessly supported me in my effort to achieve that goal.

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