The roles of competition and environmental heterogeneity in the maintenance of behavioral variation and covariation.

Many models of selection predict that populations will lose variation in traits that affect fitness. Nonetheless, phenotypic variation is commonly observed in natural populations. We tested the influences of competition and spatial heterogeneity on behavioral variation within and among populations of Merriam's kangaroo rats (Dipodomys merriami) and tested for the differential expression of trait correlations. We found that populations of D. merriami exhibited more aggression at sites with more competition. Contrary to theoretical predictions and empirical results in other systems, the sites with the greatest spatial heterogeneity and highest levels of competition did not exhibit the most behavioral variation among individuals. However, the greatest within-individual behavioral variability in boldness (response to cues of predator presence) was exhibited where spatial heterogeneity was highest. Aggression and boldness of D. merriami were highly repeatable, that is, individuals behaved in a consistent manner over time, and the two behaviors were also highly correlated. Interestingly, the strength of this correlation was greatest where the competitive community was least diverse. These findings add to increasing evidence that natural populations of animals exhibit patterns of behavioral covariance, or personality structure, and suggest that competitive variation may act to erode personality structure.