A Comparison of the Subspecific Divergence
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TITLE PAGE A COMPARISON OF THE SUBSPECIFIC DIVERGENCE OF TWO SYMPATRIC AFRICAN MONKEYS, PAPIO HAMADRYAS & CHLOROCEBUS AETHIOPS: MORPHOLOGY, ENVIRONMENT, DIET & PHYLOGENY BY JASON DUNN BSC HONS (ST AND.) PGCERT (HYMS) A THESIS SUBMITTED IN SATISFACTION OF THE REQUIREMENTS FOR INCEPTION TO THE DEGREE OF DOCTOR OF PHILOSOPHY IN THE HULL YORK MEDICAL SCHOOL SEPTEMBER 2011 THE UNIVERSITY OF HULL THE UNIVERSITY OF YORK THE HULL YORK MEDICAL SCHOOL 2 of 301 ABSTRACT The baboon and vervet monkey exhibit numerous similarities in geographic range, ecology and social structure, and both exhibit extensive subspecific variation corresponding to geotypic forms. This thesis compares these two subspecific radiations, using skull morphology to characterise the two taxa, and attempts to determine if the two have been shaped by similar selective forces. The baboon exhibits clinal variation corresponding to decreasing size from Central to East Africa, like the vervet. However West African baboons are small, unlike the vervet. Much of the shape variation in baboons is size-related. Controlling for this reveals a north-south pattern of shape change corresponding to phylogenetic history. There are significant differences between the chacma and olive baboon subspecies in the proportion of subterranean foods in the diet. No dietary differences were detected between vervet subspecies. Baboon dietary variation was found to covary significantly with skull variation. However, no biomechanical adaptation was detected, suggesting morphological constraint owing to the recent divergence between subspecies. Phylogeny correlates with morphology to reveal an axis between northern and southern taxa in baboons. In vervets C. a. sabaeus is the most morphologically divergent, which with other evidence, suggests a West African origin and radiation east and south, in contrast with a baboon origin in southern Africa. Path analyses of all the factors discussed revealed markedly different relationships between the two taxa, with baboons responding to the environment via diet rather than directly. The two subspecific radiations have different relationships with diet, environment and phylogeny. In spite of superficial similarities, the study taxa are sufficiently different that similar ecological and environmental selective forces have not produced convergent patterns of radiations. The baboon exhibits greater flexibility and larger size freeing it from limiting constraints faced by vervets. Additionally the two have distinct sites of origins and patterns of dispersals adding a stochastic element to the differences between radiations. 3 of 301 TABLE OF CONTENTS Title Page ..................................................................................................................... 1 Abstract ........................................................................................................................ 2 Table of Contents ......................................................................................................... 3 List of Figures ............................................................................................................ 10 List of Tables.............................................................................................................. 17 Acknowledgements .................................................................................................... 21 Author’s Declaration .................................................................................................. 23 CHAPTER 1. Introduction to the Study Taxa and Concepts of Subspecific Variation .................................................................................................................................... 24 1.1 Question and Aims ........................................................................................... 24 1.2 Taxonomy & Evolutionary history ................................................................... 25 1.3 Geographic Distribution ................................................................................... 29 1.4 Environment & Habitat .................................................................................... 32 1.5 Feeding & Diet ................................................................................................. 32 1.6 Appearance & Morphology .............................................................................. 34 1.7 Species Concepts and the Evolutionary Significance of Subspecies ............... 37 1.8 Social System & Behaviour.............................................................................. 44 1.9 Behavioural Flexibility ..................................................................................... 46 1.10 Goals of the Study .......................................................................................... 48 CHAPTER 2. Patterns of Clinal Variation in the Baboon ......................................... 50 2.1 Introduction ...................................................................................................... 50 2.1.0 Overview ................................................................................................... 50 2.1.1 Patterns of Clinal Variation ....................................................................... 51 2.1.2 Possible Underpinnings of Clinal Variation ............................................. 52 2.1.3 Resources & Growth ................................................................................. 55 2.1.4 Covariance of Size and Shape ................................................................... 56 4 of 301 2.1.5 Clinal Variation in Primates ...................................................................... 56 2.1.6 Clinal Variation in the Baboon ................................................................. 59 2.1.7 Aims, Hypotheses and Assumptions ......................................................... 60 2.2 Method .............................................................................................................. 62 2.2.1 Sample ....................................................................................................... 62 2.2.2 Capturing Shape ........................................................................................ 66 2.2.3 Sex-Correction .......................................................................................... 67 2.2.4 Geographical Averaging ........................................................................... 67 2.2.5 Environmental Variable Selection ............................................................ 68 2.2.6 Regressing Size and Shape on Geography ................................................ 68 2.2.7 Partitioning Spatial and Environmental Terms ......................................... 70 2.2.8 Size Correction .......................................................................................... 71 2.2.9 Subspecific Differences ............................................................................ 71 2.3 Results .............................................................................................................. 72 2.3.1 Baboon Size .............................................................................................. 72 2.3.2 Vervet Size ................................................................................................ 75 2.3.3 Baboon Shape ........................................................................................... 77 2.3.4 Size-Controlled Shape ............................................................................... 80 2.3.5 Subspecific Differences ............................................................................ 81 2.3.6 Environmental and Spatial Correlates ....................................................... 83 2.4 Discussion ........................................................................................................ 85 2.4.1 Clinal Patterns of Size ............................................................................... 85 2.4.2 Clinal Patterns of Shape ............................................................................ 88 2.4.3 The Kinda Baboon .................................................................................... 89 2.4.4 Morphological Variation and the Effects of Evolutionary History ........... 91 2.4.5 Comparison with the Vervet and Other Taxa ........................................... 92 2.4.6 Conclusion ................................................................................................ 94 5 of 301 CHAPTER 3. Dietary Specialisation of the Baboon and Vervet Subspecies ............ 96 chapter 3. ................................................................................................................ 96 3.1 Introduction ...................................................................................................... 96 3.1.0 Overview ................................................................................................... 96 3.1.1 Foraging Ecology ...................................................................................... 96 3.1.2 Baboon Diet .............................................................................................. 98 3.1.3 Vervet Diet .............................................................................................. 100 3.1.4 Dietary Comparison of the Study Taxa ................................................... 102 3.1.4 Aim .........................................................................................................