Indirect effects of territorial neighbors on the timing of spring breeding may counteract changes in selection in North American red squirrels

Abstract

Organisms can affect one anothers phenotypes when they socially interact. Indirect genetic effects occur when an individuals phenotype is affected by genes expressed in another individual. These heritable effects can enhance or reduce adaptive potential, thereby accelerating or reversing evolutionary change. Quantifying these social effects is therefore crucial for our understanding of evolution, yet estimates of indirect genetic effects in wild animals are limited to dyadic interactions. We estimated indirect phenotypic and genetic effects, and their covariance with direct effects, for the date of spring breeding in North American red squirrels (Tamiasciurus hudsonicus) living in an array of territories of varying spatial proximity. Additionally, we estimated variance parameters and the strength of selection at low and high population densities. Social effects of neighbours on the date of spring breeding were weak at low, but stronger at high population densities. Indirect phenotypic effects accounted for a larger amount of variation in the date of breeding than direct differences among-individuals, although the genetic component to these indirect effects was not statistically significant. Nevertheless, the estimated effect size was large enough to suggest that indirect genetic effects could alter evolutionary change, resulting in less change at high densities despite stronger selection. Despite the difficulty in estimating them precisely, indirect genetic effects have clear potential to alter evolutionary trajectories in any natural systems where organisms interact.