Plant defensive traits drive patterns of herbivory and herbivore diversity among plant species. Over the past 30 years, several prominent hypotheses have predicted the association of plant defenses with particular abiotic environments or geographic regions.We used a strongly supported phylogeny of oaks to test whether defensive traits of 56 oak species are associated with particular components of their climatic niche. Climate predicted both the chemical leaf defenses and the physical leaf defenses of oaks, whether analyzed separately or in combination. Oak leaf defenses were higher at lower latitudes, and this latitudinal gradient could be explained entirely by climate. Using phylogenetic regression methods, we found that plant defenses tended to be greater in oak species that occur in regions with low temperature seasonality, mild winters, and low minimum precipitation, and that plant defenses may track the abiotic environment slowly over macroevolutionary time. The pattern of association we observed between oak leaf traits and abiotic environments was consistent with a combination of a seasonality gradient, which may relate to different herbivore pressures, and the resource availability hypothesis, which posits that herbivores exert greater selection on plants in resource-limited abiotic environments.
K E Y W O R D S : Biogeography, latitude, macroevolution, Ornstein-Uhlenbeck model, plant-insect interaction, tannins.Plant defenses drive plants' associations with herbivores. Plants with certain life-history traits (such as trees) or habitat affiliations (such as desert plants) often invest more in defenses than other plants, and it has been the goal of several major theories of plant defense to understand these correlations (described in Stamp 2003). The defensive traits that a plant possesses today are a result of both its evolutionary heritage-the traits and constraints it inherited-and more recent adaptation of that plant species to its biotic and abiotic environment (Gould and Lewontin 1979;Agrawal 2007). Some comparative studies between plant species have stressed the importance of deep evolutionary history in driving plant-herbivore interactions (e.g., Mitter et al. 1991;Weiblen et al. 2006). Others have stressed local adaptation (which implies rapid evolution of defenses in environments where they convey a fitness benefit to the plant) (e.g., Fine et al. 2004;Kursar et al. 2009). Of course, the deep and recent impacts of history on any adaptive trait are not mutually exclusive (Futuyma and Agrawal 2009 and papers therein). Modern phylogenetic comparative approaches, such as phylogenetic least-squares methods and Ornstein-Uhlenbeck (O-U) modeling, allow us to begin to tease apart the effects of evolutionary history and natural selection in the evolution of plant defenses.Selection pressures imposed by different habitat types or climatic associations are thought to be a major driver of a plant's defensive investment (Stamp 2003). For example, studies have found substantial variation in plant defenses betw...