OPTIMAL OUTCROSSING IN IPOMOPSIS AGGREGAT A: SEED SET AND OFFSPRING FITNEs.

Restricted gene flow and localized selection should establish a correlation between physical proximity and genetic similarity in many plant populations. Given this situation, fitness may decline in crosses between nearby plants (inbreeding depression), and in crosses between more widely separated plants ("outbreeding depression") mostly as a result of disruption of local adaptation. It follows that seed set and offspring fitness may be greatest in crosses over an intermediate "optimal outcrossing distance." This prediction was supported for Ipomopsis aggregata, a long-lived herbaceous plant pollinated by hummingbirds. In six replicate pollination experiments, mean seed set per flower was higher with an outcrossing distance of 1-10 m than with selfing or outcrossing over 100 m. A similar pattern appeared in the performance of offspring from experimental crosses grown under natural conditions and censused for a seven-year period. Offspring from 10-m crosses had higher survival, greater chance of flowering, and earlier flowering than those from 1-m or 100-m crosses. As a result, 1-m and 100-m offspring achieved only 47% and 68%, respectively, of the lifetime fitness of 10-m offspring. Offspring fitness also declined with planting distance from the seed parent over a range of 1-30 m, so that adaptation to the maternal environment is a plausible mechanism for outbreeding depression. Censuses in a representative I. aggregata population indicated that the herbaceous vegetation changes over a range of 2-150 m, suggesting that there is spatial variation in selection regimes on a scale commensurate with the observed effects of outbreeding depression and planting distance. We discuss the possibility that differences in seed set might in part reflect maternal mate discrimination and emphasize the desirability of measuring offspring fitness under natural conditions in assessing outcrossing effects.