Determinants of spring molt in bighorn sheep: life-history, plasticity and phenology

Anthropogenic climate change and habitat alterations increase the importance of understanding the causes and consequences of variation in phenological traits. Although the timing of phenological events may vary in response to both direct and mediated effects, methods to measure and distinguish direct and mediated effects have seldom been used. We used a Bayesian structural equation model (SEM) to evaluate potential direct and mediated effects of intrinsic individual and environmental factors on the timing and progression of spring molt in bighorn sheep. The SEM showed that molt phenology varied across years, was earlier in prime-aged and in heavier individuals, slower in males, and later in lactating ewes, especially if they were light. These results highlight how individual variation in intrinsic traits and life-history leads to substantial variation in a phenological trait. Indirect effects in the SEM predicted a delay in sheep molt phenology at high population density mediated through negative density effects on body mass and lactation probability. Cooler temperatures in late spring were also predicted to delay molt phenology via a negative effect on body mass. Finally, lactation reduced ewe mass which was predicted to delay molt phenology. This mediated effect thus increased the total delay (sum of direct and mediated effects) in molt phenology experience by lactating ewes. Our results underline the importance of estimating direct and indirect effects when modeling phenological traits. Because indirect effects could substantially affect estimates of total plasticity, they should be critically important to accurately predict phenological mismatches and demographic consequences of environmental change.