Within‐Individual Correlations Reveal Link Between a Behavioral Syndrome, Condition, and Cortisol in Free‐Ranging Belding's Ground Squirrels

Animals often exhibit consistent individual differences in behavior (i.e., animal personality) and correlations between behaviors (i.e., behavioral syndromes), yet the causes of those patterns of behavioral variation remain insufficiently understood. Many authors hypothesize that state‐dependent behavior produces animal personality and behavioral syndromes. However, empirical studies assessing patterns of covariation among behavioral traits and state variables have produced mixed results. New statistical methods that partition correlations into between‐individual and residual within‐individual correlations offer an opportunity to more sufficiently quantify relationships among behaviors and state variables to assess hypotheses of animal personality and behavioral syndromes. In a population of wild B elding's ground squirrels ( U rocitellus beldingi ), we repeatedly measured activity, exploration, and response to restraint behaviors alongside glucocorticoids and nutritional condition. We used multivariate mixed models to determine whether between‐individual or within‐individual correlations drive phenotypic relationships among traits. Squirrels had consistent individual differences for all five traits. At the between‐individual level, activity and exploration were positively correlated whereas both traits negatively correlated with response to restraint, demonstrating a behavioral syndrome. At the within‐individual level, condition negatively correlated with cortisol, activity, and exploration. Importantly, this indicates that although behavior is state‐dependent, which may play a role in animal personality and behavioral syndromes, feedback mechanisms between condition and behavior appear not to produce consistent individual differences in behavior and correlations between them.