Integrating physiological stress into the movement ecology of migratory ungulates: a spatial analysis with mule deer.

Rapid climate and human land-use change may limit the ability of long-distance migratory herbivores to optimally track or 'surf' high-quality forage during spring green-up. Understanding how anthropogenic and environmental stressors influence migratory movements is of critical importance because of their potential to cause a mismatch between the timing of animal movements and the emergence of high-quality forage. We measured stress hormones (fecal glucocorticoid metabolites; FGMs) to test hypotheses about the effects of high-quality forage tracking, human land-use and use of stopover sites on the physiological state of individuals along a migratory route. We collected and analysed FGM concentrations from 399 mule deer () samples obtained along a 241-km migratory route in western Wyoming, USA, during spring 2015 and 2016. In support of a fitness benefit hypothesis, individuals occupying areas closer to peak forage quality had decreased FGM levels. Specifically, for every 10-day interval closer to peak forage quality, we observed a 7% decrease in FGMs. Additionally, we observed support for both an additive anthropogenic stress hypothesis and a hypothesis that stopovers act as physiological refugia, wherein individuals sampled far from stopover sites exhibited 341% higher FGM levels if in areas of low landscape integrity compared to areas of high landscape integrity. Overall, our findings indicate that the physiological state of mule deer during migration is influenced by both anthropogenic disturbances and their ability to track high-quality forage. The availability of stopovers, however, modulates physiological responses to those stressors. Thus, our results support a recent call for the prioritization of stopover locations and connectivity between those locations in conservation planning for migratory large herbivores.