The Genetics of Altruism covers the primary findings on social evolution, social trait, and altruism from a population genetics standpoint to establish a system of genetic boxes. It presents an evolutionary question with two faces: Why are there so many social species? Why, in all the diversity of the animal kingdom, are the social species so few? To address the evolutionary question, this book focuses on recognition of the fact that on an evolutionary time, scale genetics must underlie all changes in the capacity for social structure and other aspects of organic evolution. It presents comparative analyses framed in mathematical terms; mathematical concepts as a means of getting outside human, perhaps more generally primate and carnivore; frames of reference; and alternative network combinatorics as a natural basis for comparing social structures that are phylogenetically remote. It also discusses the comparative biology of social behavior on a purely descriptive basis through the social and evolutionary structures emergent. The book concludes by discussing major evolutionary pathways, various kinds of preadaptedness for sociality, and the use of cascade principle to suggest ways in which human evolution may have been a special case. This book is a valuable resource for biologists, social scientists, researchers, students, and all those who want to broaden their knowledge in the field of social behavior and altruism.

PrefaceAcknowledgementsList of FiguresList of TablesChapter 1 The Evolutionary Roots of Sociality 1.1. Statement of the Problem 1.2. Varieties of Selection for Social Behavior 1.3. Characteristics of Social Behavior as an Object of Selection 1.4. Genetic Models 1.5. The Evolutionary Setting 1.6. Plan of the Book NotesPart I The Theory of Reciprocity Selection Chapter 2 Mathematical Models for a Simple Cooperative Trait 2.1. A Minimal Model and Its Threshold ßerit 2.2. Behavioral Interpretation of the Parameters 2.3. Generalizations of the Minimal Model: Robustness and the Effects of Cheating 2.4. Effects of Varying the Mendelian Dominance of the Social Trait Appendix. Interactions with the Mating System Notes Chapter 3 Cascade t o Takeover by the Social Trait 3.1. Extension of the Minimal Model to an Island-Structured Metapopulation 3.2. Cascade Behavior: Numerical Examples 3.3. The Two-Island Approximation and Its Uses Appendix. Notes on a Demographic Accounting Problem Notes Chapter 4 Dynamics of the Cascade Using the Two-Island Approximation 4.1. Analysis of a True Two-Island Model 4.2. Analysis of the Two-Island Approximation 4.3. The Case Where the Source Island for Socials Remains at Fixation Appendix. Perturbation Solutions for the Fixed Points in the Small m Case and Determination of Stability Character Notes Chapter 5 The Cascade Continued-Initial Conditions and Global Dynamics 5.1. Setting the Initial Conditions by Genetic Drift 5.2. The Cascade: Basic Mechanisms of Propagation 5.3. Comparative Statics: Effect of Mother Site Centrality on Viability and Speed of Cascade 5.4. Self-Erasing Cascade Histories and the Reversibility of Social Evolution 5.5. Generalizations Appendix. Numerical Studies of Two-Dimensional Regular Stepping-Stone Models NotesPart II The Theory of Kin Selection Chapter 6 General Models for Sib and Half-Sib Selection 6.1. Précis of the Hamilton Theory 6.2. Outline of the New Models 6.3. Formalism and Derivation of Random Mating Recursions 6.4. Stability Analysis: Conditions for Stability at Fixation and Justification of Hardy-Weinberg Analysis near Dominant Fixation Appendix 6.1. The Concept of Genes Identical by Descent and the Calculation of Wright's Coefficient of Relationship Appendix 6.2. A General Haplodiploid Model Notes Chapter 7 Axiomatization of Sib Selection Theories 7.1. Axiomatic Comparisons of Fixation Stability Conditions 7.2. Analysis of Polymorphism Notes Chapter 8 Alternative Combinatorial Models and the Status of the Hamilton Theory 8.1. The Appropriateness of Sib Selection as a Model of Social Evolution in Hymenoptera 8.2. A First Combinatorial Model and Its Hamilton Limit 8.3. Alternative Combinatorics for Fitness Assignment 8.4. Concurrent Individual and Sib Selection 8.5. The Present Status of the Hamilton Conjecture: Information from the Axioms 247 8.6. Models for the Evolutionary Differentiation of Caste in Social Insects Appendix 8.1. Details of ó Graphs 261 Appendix 8.2. Stability Conditions for Sib Selection Models with Combinatorial Coefficients from Section 8.2 Notes Chapter 9 Models of Intergenerational Altruism 9.1. Child-Parent Altruism in a Haplodiploid Species 9.2. Parental Investment in a Haplodiploid Species Notes Part III The Theory of Group Selection Chapter 10 Analysis of Group Selection in the Levins E = E( x ) Formalism 10.1. Overtime Solution of the Basic Levins Equation 10.2. Biological Discussion of the Results 10.3. Comparison with the Levins Analysis Appendix. Behavior of ¿(x, 1) near x = 0 and x = 1 for h ¿ (0,1) Notes Chapter 11 Group Selection of Founder Populations 11.1. Derivation of a New Model 11.2. Analysis: Fixed Points and Stability 11.3. Interpretation Notes Chapter 12 Conclusions 12.1. Pathways to Sociality 12.2. Preadaptations for Social Evolution 12.3. Reflections on Data 12.4. Evolutionary Limits on Sociality 12.5. Notes on Hominid ApplicationsTechnical Appendix: Genetics Background Glossary ReferencesAuthor IndexSubject Index