Evolution of the Family Evolution by Kin Selection Genetic Trait Expressed in Actor (Ego) Must Affect Genotypic Fitness of Individual Related to Actor at Locus Determining Trait Altruism, Selfishness, Mutualism

Evolution of the Family Evolution by Kin Selection Kin-Based Altruism Helpers at the Nest (Diploids) Sterile Worker Caste, Eusociality (Haplo-Diploids)

Evolution of the Family Proximity of Relatives Competition Among Kin Dominant, Subordinate Offspring Siblicide, Fratricide Brood Reduction (By Offspring Sex) “Weaning” Conflict

Gauthier & Barrette Behaviour 94:128.

Feeding of Fawns in 2 Deer Spp As Fawns Age Fewer Bouts Decreasing Proportion Initiated by Mother More Frequent Rejection by Mother Fawn Independence Increases; Mother Future Reproduction

Packard et al Canadian J. Zool. 70:1269. Arctic Wolves Dominant Female Suckling Less Frequent & Bouts End Sooner as Cubs Age

Packard et al Canadian J. Zool. 70:1269.

Feeding of Cubs in Arctic Wolf Standard Pattern in Frequency, Initiation & Termination As Offspring Age Predict Why Close Kin “Fight?”

Parenting Parental Care Per Se Traditional Darwinian Explanation Adequate, But Can Justify via Hamilton’s Rule Parent-Offspring Conflict: Parent Gives Care to Offspring Offspring Demands Parental Resource

Parent-Offspring Conflict Parent-Offspring Conflict (Diploidy) Parent Altruism Offspring Selfishness Close Relatives: Hamilton’s Rule (General)

Parent-Offspring Conflict If Parent Allocates Resources to Offspring: B Benefit to Offspring C Cost to Parent, Loss of Future Reproductive Success r (Parent to Offspring) = 1/2

Kin Selection Parent Allocate Resources to Offspring? Parent Allocate Resources to Offspring? Hamilton’s Rule: Increase Parent’s Inclusive Fitness r B – C > 0, Implying C/B < (1/2) Parental Altruism Favored

Kin Selection Offspring Demand Parental Resources? Offspring Demand Parental Resources? Hamilton’s Rule Generalized: Increase Offspring Inclusive Fitness E S Effect on Self = B E R Effect on Relative (Parent) = - C r (Offspring to Parent) = 1/2

Kin Selection Offspring Demand Parental Resources? Offspring Demand Parental Resources? Increase Offspring Inclusive Fitness E S + r E R > 0 B – (1/2) C > 0; C/B < 2 Offspring Selfishness Favored

Kin Selection C/B < ½ Parental Altruism Favored C/B < 2 Offspring Selfishness Favored ½ < C/B < 2 Parent Should Not Be Altruistic Offspring Should Be Selfish Kin Conflict

Kin Selection Selfish Genes Can Lead, Evolutionarily, To Cooperation and Altruism Gene Spreads by Helping Other Individuals Carry Same Gene Hamilton’s Rule; Hamilton’s Conjecture

Kin Selection Limited Dispersal (of Newborns) Makes Neighbors Close Relatives Enough Local Relatedness Might Lead to Cooperation/Altruism Toward Any Neighbor

Kin Selection Viscous Population, Low Dispersal, Relatives = Neighbors Limited Dispersal Also Can Make Relatives Close Competitors Competitive Local Conflict Over Resources Negate Effect of Kin?

Limited Dispersal and Kin Evolution of Cooperation Relatedness Promotes (Hamilton) Local Competition Impedes Interaction

Local & Global Scales Behavioral Interactions Local, Ephemeral Drive Ecological, Evolutionary Dynamics Ecology, Evolution Global (Population), Extended Timescale