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Abstract Plasticity in plant traits, including secondary metabolites, is critical to plant survival and competitiveness under stressful conditions. The ability of a plant to respond effectively to combined stressors can be impacted by crosstalk in biochemical pathways, resource availability and evolutionary history, but such responses remain underexplored. In particular, we know little about intraspecific variation in response to combined stressors or whether such variation is associated with the stress history of a given population.Here, we investigated the consequences of combined water and herbivory stress for plant traits, including relative growth rate, leaf morphology and various measures of phytochemistry, using a common garden ofAsclepias fascicularismilkweeds. To examine how plant trait means and plasticities depend on the history of environmental stress, seeds for the experiment were collected from across a gradient of aridity in the Great Basin, United States. We then conducted a factorial experiment crossing water limitation with herbivory.Plants responded to water limitation alone by increasing the evenness of UV‐absorbent secondary metabolites and to herbivory alone by increasing the richness of metabolites. However, plants that experienced combined water and herbivory stress exhibited similar phytochemical diversity to well‐watered control plants. This lack of plasticity in phytochemical diversity in plants experiencing combined stressors was associated with a reduction in relative growth rates.Leaf chemistry means and plasticities exhibited clinal variation corresponding to seed source water deficits. The total concentration of UV‐absorbent metabolites decreased with increasing water availability among seed sources, driven by higher concentrations of flavonol glycosides, which are hypothesized to act as antioxidants, among plants from drier sites. Plants sourced from drier sites exhibited higher plasticity in flavonol glycoside concentrations in response to water limitation, which increased phytochemical evenness, but simultaneous herbivory dampened plant responses to water limitation irrespective of seed source.Synthesis. These results suggest that climatic history can affect intraspecific phytochemical plasticity, which may confer tolerance to water limitation, but that co‐occurring herbivory disrupts such patterns. Global change is increasing the frequency and intensity of stress combinations, such that understanding intraspecific responses to combined stressors is critical for predicting the persistence of plant populations. 

Abstract Intensifying drought conditions across the western United States due to global climate change are altering plant–insect interactions. Specialist herbivores must find their host plants within a matrix of nonhosts, and thus often rely upon specific plant secondary chemistry for host location and oviposition cues. Climate-induced alterations to plant chemistry could thus affect female selection of larval food plants. Here, we investigated whether host-plant water limitation influenced oviposition preference in a threatened invertebrate: the monarch butterfly (Danaus plexippus). We found that females deposited more eggs on reduced-water than on well-watered narrowleaf milkweed plants (Asclepias fascicularis), but we could not attribute this change to any specific change in plant chemistry. Specialist herbivores, such as the monarch butterfly, which are tightly linked to specific plant cues, may experience shift in preferences under global-change conditions. Understanding oviposition preferences will be important to directing ongoing habitat restoration activities for this declining insect. 

Virtually all plants employ direct and indirect defenses against herbivores. While it is known that plant defenses can be affected by belowground symbiotic microbes under controlled conditions, studies showing these multitrophic interactions in nature are surprisingly scarce. Here we tested for effects of rhizobia on insect attraction and direct defense (cyanogenesis) in wild lima bean (Phaseolus lunatus) plants in Costa Rica. We performed bioassays with rhizobia-inoculated (R+) and rhizobiafree (R-) potted plants distributed among native lima bean communities at two spatially separated field sites (450 km apart) and in two field seasons. Without affecting overall plant size, rhizobia altered leaf chemistry (cyanogenesis and soluble leaf nitrogen) and ultimately insect communities visiting the plants. Natural herbivorous chrysomelid beetles were strongly attracted to R+ plants, while natural enemies, ants and parasitoid wasps, preferred R- plants resulting in a particularly high herbivore:carnivore ratio on R + plants. This suggests that symbiotic microbes mediate trophic interactions by influencing both direct and indirect plant defenses against herbivores. Our results show that rhizobia affect the plant defensive phenotype and have cascading effects on plant-insect interactions in nature.