Abstract
Coevolution between plants and their animal predators has led to diverse defensive adaptations. Multiple theories of defense propose that there are resource allocation costs associated with producing chemical defenses. One leading hypothesis, optimal defense theory (ODT), suggests that natural selection will result in the allocation of resources to defenses that optimize the cost-to-benefit ratio between defense and other functional processes. The population decline of the rare subalpine wetland species, Arizona willow (Salix arizonica), has been attributed to various biotic and abiotic factors, with browsing from wild and domestic ungulates as a significant concern for at least three decades. In a field experiment using natural populations, we compare the relationship between phytochemical defense and height in Arizona willows with and without long-term protection from browsing via browse exclosures. Consistent with the predictions of ODT, individuals with physical protection from ungulate browsing for multiple years had significantly lower phenolic glycoside (PG) concentrations and increased plant height compared to unprotected individuals. A similar pattern was found across all individuals, whereby total PG concentration and height were negatively correlated. In a short-term experiment in natural populations, changes in levels of defense were not observed when plants received protection for only one growing season. The contrasting pattern of defense plasticity in response to long-term versus short-term physical protection suggests a differential plastic response in this long-lived species. Delayed reduction in PG concentration may serve as a benefit to avoid mismatches between environmental cues and responses. Our research sheds light on the intricate dynamics between plant-defense strategies, environmental pressures, and evolutionary adaptations in shaping plant-browser interactions.