The Red Colobus Monkeys: Variation in Demography, Behavior, and Ecology of Endangered Species
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Description
Based on field studies spanning nearly 40 years, this reference book summarizes and integrates past research with new and previously unpublished information on the behavioral ecology of Africa's red colobus monkeys from study sites as diverse as Senegal, Uganda, and Zanzibar. It provides an unparalleled compilation of information on taxonomy, genetics, vocalizations, demography, social organization, dispersal, social behavior, reproduction, mortality factors, diet, ranging patterns, interspecific relations, and conservation. Social relationships in red colobus are less rigidly structured than in other African monkeys, resulting in considerable variation in social organization and group composition, both within and between taxa. This provides a unique opportunity to examine the extent to which social variables correlate with differences in habitat quality, demography, and predation by chimpanzees, and humans. Unfortunately, at least half of the 18 taxa of red colobus are now threatened with extinction. Conservation problems are described, causal factors identified, and solutions proposed. This volume is intended not only to serve as a reference book, but to stimulate and guide future long-term research and to encourage effective conservation action. Contents List of Figures List of Tables Preface Acknowledgments 1 General biology of red colobus 1.1 General description 1.2 Paleontology 1.3 Intrataxon variation in color 1.3.1 Summary comments and speculation on coat color 1.4 Facial color: pink noses and mouths 1.5 Taxonomy 1.6 Summary points 2 Vocalizations 2.1 Introduction 2.2 Methods and localities 2.3 Common vocalizations 2.4 Vocalizations unique to specific taxa 2.5 Intertaxa and geographical comparisons: implications for evolution and phylogeny 2.6 Long and loud call bouts: contrasts in form and function 2.7 Alarm calls and semanticity 2.7.1 Avian predators 2.7.2 Poisonous snakes 2.8 Semanticity of copulation and estrous calls 2.9 Summary points 3 Demography: social group size and composition and population density 3.1 Introduction 3.2 Methodological caveats 3.3 Variation in group size and composition 3.4 Differences between taxa in group size 3.5 Differences between populations of the same taxon in group size 3.6 Differences within populations in group size over space and time 3.7 Summary of probable determinants of group size 3.8 Solitary red colobus 3.9 Differences between taxa in adult sex ratios 3.10 Differences between populations of the same taxon in adult sex ratio 3.11 Differences within populations in adult sex ratios over space and time 3.12 Differences between taxa in ratios of immatures to adult females 3.12.1 SAJ per adult female ratios 3.12.2 Infants per adult female 3.12.3 Demographic correlates of infant per adult female ratios 3.13 Population density 3.13.1 Methodological issues 3.14 Population density estimates based on studies of focal groups: an example from Kibale 3.15 Differences in population densities between taxa 3.16 Differences in population density within taxa between and within populations 3.17 Differences within taxa over time 3.18 Summary points 4 Social organization: intergroup relations, tenure, longevity, and dispersal 4.1 Introduction 4.2 Intergroup relations 4.2.1 CW group of tephrosceles, Kanyawara, Kibale 4.2.2 RUL group of tephrosceles, Ngogo, Kibale 4.2.3 Intergroup relations of other populations and taxa 4.2.4 Summary of intergroup relations 4.3 Longevity and tenure length within social groups 4.3.1 CW group, Kanyawara, Kibale 4.3.2 RUL group, Ngogo, Kibale 4.3.3 HTL Group, Ngogo, Kibale 4.4 Immigration and tenure length 4.4.1 Female immigrants 4.4.2 Male immigrants 4.4.3 Natal females 4.4.4 Natal males 4.5 General remarks on longevity, tenure, and dispersal in Kibale 4.6 Comparison with other red colobus taxa 4.7 Summary of longevity, tenure, and dispersal 5 Social behavior and reproduction 5.1 Introduction 5.2 Grooming 5.2.1 CW group of tephrosceles, Kanyawara, Kibale 5.2.2 RUL group of tephrosceles, Ngogo, Kibale 5.2.3 Summary comparison of grooming in CW and RUL groups 5.2.4 Grooming in other taxa of red colobus 5.3 Sexual behavior and reproduction 5.3.1 General background information 5.3.2 Male copulatory/reproductive success 5.3.3 Female reproductive success and interbirth intervals (IBI) 5.3.4 Timing of births 5.3.5 Female perineal swellings 5.3.6 Summary of sexual behavior and reproduction 5.4 Aggression 5.4.1 General background information 5.4.2 Supplantations: CW group tephrosceles, Kanyawara, Kibale 5.4.3 Supplantations: RUL group of tephrosceles, Ngogo, Kibale 5.4.4 Intense aggression: CW group of tephrosceles, Kanyawara, Kibale 5.4.5 Intense aggression: RUL group of tephrosceles, Ngogo, Kibale 5.4.6 General comment on the relation between dominance and aggression among tephrosceles 5.4.7 Harassment of adults by immatures in nonsexual contexts among tephrosceles, Kibale, Uganda 5.4.8 Present type I: an appeasement gesture 5.4.9 Present type II: a dominance gesture 5.4.10 Harassment during copulation 5.4.11 Aggression in other taxa of red colobus 5.4.12 Summary of aggression 5.5 Interindividual distance 5.6 Social relations of infants and small to medium juveniles 5.6.1 Neonates 5.6.2 Older infants and small juveniles 5.6.3 Twins 5.6.4 Differential investment by mothers in sons vs. daughters 5.6.5 Summary of social relations of infants and small to medium juveniles 5.7 Greeting behavior in tephrosceles 5.7.1 Summary of greeting behavior 6 Ecology 6.1 Introduction 6.2 Diet 6.2.1 Methods used in Kibale 6.2.2 Diets of the CW and RUL groups of tephrosceles, Kibale, Uganda 6.2.3 Comparison with other studies of tephrosceles in Kibale, Uganda 6.2.4 Intertaxa comparison of plant part diet 6.2.5 Phytochemical basis of diet 6.2.6 Miscellaneous information on dietary habits 6.2.7 Summary points on diet 6.3 Activity budgets 6.3.1 Methods 6.3.2 Comparison of the RUL and CW group activity budgets 6.3.3 Comparison of activity budgets between studies and taxa 6.3.4 Summary of activity budgets 6.4 Ranging behavior 6.4.1 Home range 6.4.2 Daily Travel Distance 6.5 Mortality 6.5.1 Diseases in Kibale red colobus 6.5.2 Fatal falls 6.5.3 Predation 6.6 Non-predator interspecific relations of red colobus 6.6.1 Defense against predation 6.6.2 Food competition 6.6.3 Social 7 Conservation 7.1 Introduction 7.2 Conservation status of the 18 red colobus taxa 7.2.1 Temminckii 7.2.2 Badius 7.2.3 Waldroni 7.2.4 Epieni 7.2.5 Pennantii 7.2.6 Preussi 7.2.7 Oustaleti 7.2.8 Tholloni 7.2.9 Tephrosceles 7.2.10 Rufomitratus 7.2.11 Gordonorum 7.2.12 Kirkii 7.3 Case studies of threats 7.3.1 Hunting: the case of Miss Waldron’s red colobus (waldroni) 7.3.2 Agricultural expansion: the case of the Udzungwa red colobus (gordonorum) 7.3.3 Tourism and deforestation: the case of the Zanzibar red colobus (kirkii) 7.3.4 Selective logging: the case of the Ugandan red colobus (tephrosceles) in Kibale 7.3.5 Interagency conflicts of interest: the case of the Udzungwa red colobus (gordonorum) 7.4 Extrinsic versus intrinsic threats 7.5 Problems in protected areas 7.6 Proximate variables affecting conservation and possible solutions 7.7 Ultimate variables affecting conservation and possible solutions 7.7.1 Human population growth 7.7.2 Overconsumption of resources 7.7.3 Changing values and behavior 7.7.4 Funding for conservation 7.8 Conclusions 7.9 Summary Appendix 1 Annotated list and measurements of red colobus taxa Appendix 3.1 Age and size classes of red colobus Appendix 3.2 Red colobus group composition Appendix 3.3 Statistical comparison of adult sex ratios between and within taxa Appendix 3.4 Statistical comparison of subadult and juvenile to adult female ratios between and within taxa Appendix 3.5 Statistical comparison of infant to adult female ratios between and within taxa Appendix 4.1 Log of individuals in CW group tephrosceles Appendix 4.2 Log of individuals in RUL group tephrosceles Appendix 4.3 Log of individuals in HTL group tephrosceles Appendix 5.1 Grooming of adult males in RUL group tephrosceles Appendix 5.2 Aggression by adult males in CW group tephrosceles Appendix 6.1 Total diet CW group tephrosceles Appendix 6.2 Total diet RUL group tephrosceles Appendix 6.3 Notes on disease in tephrosceles of Kibale References Index A B C D E F G H I J K L M N O P R S T U V
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