INCREASED MOTH HERBIVORY ASSOCIATED WITH ENVIRONMENTAL-STRESS OF PINYON PINE AT LOCAL AND REGIONAL LEVELS

Using 6 years of observational and experimental data, we examined the hypothesis that water and nutrient stress increase the susceptibility of pinyon pine (Pinus edulis) to the stem- and cone-boring moth (Diory ctria albovittella). At two geographic levels, a local scale of 550 km (2) and a regional scale of 10,000 km(2), moth herbivory was strongly correlated with an edaphic stress gradient. At a local scale, from the cinder soils of Sunset Crater to nearby sandy-loam soils, nine of ten soil macro- and micronutrients, and soil water content were lowest in cinder-dominated soils. Herbivore damage was six times greater on tre es growing in the most water and nutrient deficient site at Sunset Cra ter compared to sites with well-developed soils. Percentage silt-clay content of soil, which was highly positively correlated with soil nutr ient and soil moisture at a local scale, accounted for 56% of the vari ation in herbivory at a regional scale among 22 sites. Within and acro ss sites, increased stem resin flow was positively associated with red uced moth attack. On the basis of moth distribution across a stress gr adient, we predicted that pinyons growing in highly stressful environm ents would show increased resistance to herbivores if supplemented wit h water and/or nutrients. We conducted a 6-year experiment at a high-s tress site where individual trees received water only, fertilizer only , and water + fertilizer. Relative to control trees, stem growth and r esin flow increased in all three treatments, but only significantly in the water + fertilizer treatment. Although there was no significant d ifference in herbivore damage among these three treatments, there was an overall reduction in herbivore damage on all treatment trees combin ed, compared to control trees. This experiment suggests that release f rom stress leads to increased resistance to insect attack and is consi stent with our observational data. While other studies have predicted that short-term stress will result in herbivore outbreaks, our studies extend this prediction to chronically stressed host populations. Fina lly, while flush-feeders are not predicted to respond positively to st ressed host plants, we found a positive association between herbivore attack and stressed pinyon populations.