1. What are the four questions Tinbergen asked about animal behavior, and explain the gist of each question.

C – Causation (0.5 pts) refers to the proximate causes of behavior such as the hormones, genes, nerve impulses that control the expression of behaviors. (0.75 pts)

D -- Development (0.5 pts) refers to the ontogeny of behaviors such as imprinting, or in the case of cognition, issues of learning. (0.75 pts)

E -- Evolution (0.5 pts) refers to the phylogenetic context in which the behaviors are found. For example, the prevalence of parental care in birds, but not reptiles (with some exceptions) is an example of the taxonomic affiliations of some behaviors. (0.75 pts)

F -- Function (0.5 pts) refers to the adaptive value or contribution that the behavior makes to fitness. (0.75 pts)

2. Under what conditions will group selection overwhelm the force of individual selection? Discuss this in terms of Wynne-Edwards’ original proposal for explaining how self-limiting reproductive behaviors might evolve in a population.

"Classic" group selection stems from arguments that Wynne-Edwards made in the 1960's. Wynne-Edwards was arguing for the evolution of a form of population regulation by the mechanism of group selection in which

            (0.5 pts) individual behaviors evolve to be self-limiting (e.g. avoid stripping resource base)

            (0.5 pts) because they benefit the group’s existence by lowering the chance of extinction of the group.

For group selection to overwhelm individual selection:

            groups would have to be very small (1 pt), so that group survival and individual survival are contingent upon factors occurring at approximately the same time scale. (1 pt)

            and the selective advantage among groups is enormous to overwhelm the ever-present force of individual selection (i.e. entire groups have a higher relative likelihood of extinction than individuals within groups have) (1 pt), so that pro-group altruism becomes more beneficial to the individual as a member of the group than does a pro-individual cheater strategy. (1 pt)

    (0.5 pts) for mentioning altruism and/or cheater strategies.

            (0.5 pts) for mentioning altruism and/or cheater strategies.