Family Planning in the Kestrel (Falco Tinnunculus): the Ultimate Control of Covariation of Laying Date and Clutch Size

in Behaviour
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Abstract

The theory that individual birds maximize their fitness by the two major decisions in reproduction concerning date (when to start laying eggs) and clutch size (when to stop laying eggs) is empirically approached in the Kestrel by quantifying Fisher's Reproductive Value for both the clutch (Vc = c. Vo/2) and the parents (Vp). The reproductive value of an egg (Vo) was found to decrease monotonically with laying date (d) due to significant associations with d of the components So (probability for an egg to survive till fledging), S1 (probability to survive from fledging till age 1), S2 (survival age 1 till age 2), and P1 (probability of breeding at age 1). Vp declined negligibly with laying date, although there were significant associations between d (laying date) and N (probability of the nest to produce at least one fledging), Pr (probability of a repeat clutch following nest failure), and La (probability of local survival of the parents following breeding). In experiments where brood size at day 10 after hatching was increased or reduced, Vc increasing experimental brood size, while Vp simultaneously decreased. Total reproductive value (V = Vc + Vp) remained unaffected by the experiments. This result suggests that a rather broad range of clutch sizes maximizes total reproductive value, as far as detectable by the data. While the yield of kestrel hunting, and hence the number of young raisable with constant parental effort (and constant Vp), increased with the spring increase in vole population density, reproductive value of the clutches decreased. For any particular food situation (hunting yield) this leads to a unique combination of clutch size and laying date maximizing V. This could be worked out by calculating fitness contours for all combinations and for different yields (Fig. 12). The optimal solutions are on a declining slope, with smaller clutches associated with later dates. 59.4% of all clutches observed obeyed the maximization criteria. Furthermore, there was a reasonable, unbiased association between predicted laying dates and clutch sizes based on individual male hunting yields and observed dates and clutches as laid by the females. Qualitatively, any method predicts a seasonal decrease in the optimal clutch size when the environment improves while reproductive value declines with progressive date. Preliminary results from an experimental approach to test the assumption of a causal effect of date on Vo, using the release of juvenile kestrels reared in captivity under artificial light schedules, are presented.

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