Seed Selection by the Harvester Ant Pogonomyrmex rugosus (Hymenoptera: Formicidae) in Coastal Sage Scrub: Interactions With Invasive Plant Species.

Harvester ants can be the dominant seed predators on plants by collecting and eating seeds and are known to influence plant communities. Harvester ants are abundant in coastal sage scrub (CSS), and CSS is frequently invaded by several exotic plant species. This study used observations of foraging and cafeteria-style experiments to test for seed species selection by the harvester ant Pogonomyrmex rugosus Emery (Hymenoptera: Formicidae) in CSS. Analysis of foraging behavior showed that P. rugosus carried seeds of exotic Erodium cicutarium (L.) and exotic Brassica tournefortii (Gouan) on 85 and 15% of return trips to the nest (respectively), and only a very few ants carried the native seeds found within the study areas. When compared with the availability of seeds in the field, P. rugosus selected exotic E. cicutarium and avoided both native Encelia farinosa (Torrey & A. Gray) and exotic B. tournefortii. Foraging by P. rugosus had no major effect on the seed bank in the field. Cafeteria-style experiments confirmed that P. rugosus selected E. cicutarium over other available seeds. Native Eriogonum fasciculatum (Bentham) seeds were even less selected than E. farinosa and B. tournefortii.

Parasitism of an insect Manduca sexta L. alters feeding behaviour and nutrient utilization to influence developmental success of a parasitoid.

The effects of macronutrient balance on nutrient intake and utilization were examined in Manduca sexta larvae parasitized by Cotesia congregata. Insects fed an artificial diet having constant total macronutrient, but with varied ratios of protein and carbohydrate, with altered diet consumption in response to excesses and deficiencies of the individual macronutrients. Bivariate plots of protein and carbohydrate consumption for non-parasitized larvae demonstrated a curvilinear relationship between points of nutrient intake for the various diets, and the larvae grew best on carbohydrate-biased diets. The relationship was linear for parasitized larvae with the growth uniform across diets. On protein-biased diets, the larvae regulated the nitrogen content, containing similar amounts of nitrogen regardless of consumption. Efficiency of nitrogen conversion in non-parasitized larvae was greatest on carbohydrate-biased diets, while nitrogen conversion by parasitized larvae was greatest with intermediate nutrient ratios. Accounting for carbohydrate consumption, the lipid content decreased as dietary carbohydrate increased, but parasitized larvae contained significantly less lipid. The total biomass of parasites developing in individual host larvae was positively correlated with host protein consumption, but the individual parasites were similar in size. Parasitism influences host nutrient consumption in a manner that achieves uniform host growth under diverse nutritional regimes, thereby constraining blood nutrient concentrations within limits suitable for parasite growth and development.

Nicotine moderates the effects of macronutrient balance on nutrient intake by parasitized Manduca sexta L.

Effects of dietary nicotine and macronutrient ratio on M. sexta larvae were examined. Larvae were fed a carbohydrate-biased, protein-biased or diet having equal amounts of casein and sucrose, with and without nicotine. Without nicotine, larvae displayed compensatory feeding on the low protein diet, but despite consuming more, grew least on this diet. Nicotine at 0.5% had no effect on nutrient consumption. Nicotine at 1.0 and 2.0% reduced overall consumption and thereby also reduced nicotine consumption. Larvae parasitized by C. congregata displayed reduced nutrient intake and growth on all diets. Parasitized larvae responded to 1% nicotine similarly to unparasitized larvae. At 0.5% nicotine, they displayed reduced consumption on all diets, possibly due to altered chemoreceptor sensitivity to nicotine. When offered a choice of two diets having different macronutrient ratios, one with and the other without 0.1% nicotine, all larvae preferred the diet lacking nicotine and failed to regulate nutrient intake such that the nutrient intake target, a ratio of nutrients supporting optimal growth, was achieved. Parasitized larvae consumed less nicotine on a fresh weight basis than unparasitized insects, suggesting that the feeding response of parasitized larvae to nicotine minimizes the exposure of nicotine to developing parasites.

Impact of a screen barrier on dispersion behavior of Homalodisca coagulata (Hemiptera: Cicadellidae).

The recent incursion of the sharpshooter Homalodisca coagulata (Say) (Hemiptera: Cicadellidae), a vector of the plant pathogenic bacterium Xylellafastidiosa, into southern California has caused new epidemics of plant diseases. The potential of H. coagulata to spread throughout the state and disseminate disease has encouraged the development of techniques to limit further spread of the vector and to manage disease epidemics where the insect already exists. We evaluated a unique tactic to curtail the immigration of H. coagulata into high-value crops, including nursery stock, where they can be spread via commercial transportation throughout the state and into disease-susceptible vineyards. This tactic consists of a 5-m-high screen barrier surrounding the crop that is impenetrable to H. coagulata. We examined H. coagulata orientation and flight direction when placed near or on the screen barrier, and determined the proportion of insects that flew over it. When released midway between a barrier and adjacent vegetation 71.5% of H. coagulata flew away from the barrier and in the direction of the vegetation; whereas 29.5% flew in the direction of the barrier. Of the total number of H. coagulata released, 7.5% flew over the barrier. When placed on the barrier, H. coagulata generally climbed up an average of 1.16 m before flying away from the structure. Of the total number of H. coagulata placed on the barrier, 6% flew over it. These results suggest that a screen barrier can be a part of a management strategy to reduce number of H. coagulata in high-value crops.

Feeding behaviour and nutrient selection in an insect Manduca sexta L. and alterations induced by parasitism.

Manduca sexta L. larvae exhibit broad food acceptance with regard to nutrient content during the first 3 days of the last stadium. Larvae fed diets with a constant combined level of casein and sucrose, but variable ratios, display a linear relationship between protein and carbohydrate intake. Larvae grow best on a diet with equal nutrients, but will consume an excess of one nutrient in order to obtain an adequate amount of the other, as nutrient ratio shifts. Parasitized larvae feed similarly, but the nutrient ratio does not affect growth. Unparasitized larvae regulate intake of protein and carbohydrate when offered choices of protein-biased and carbohydrate-biased diets having combined nutrient levels of 120 g/l, but with variable ratios. Larvae normally consume equal amounts of nutrients, regardless of ratio, and grow similarly. As combined nutrient level is reduced in one diet, larvae abandon regulation and feed randomly. Parasitized larvae offered choice diets with 120 g/l combined nutrients do not regulate nutrient intake. Consumption of nutrients varies widely, but growth is unaffected. Larvae offered choices of diets having equal amounts of casein and sucrose but variable fat (corn oil), fail to regulate fat intake, although both unparasitized and parasitized larvae prefer a diet containing higher fat.

Host nutrition determines blood nutrient composition and mediates parasite developmental success: Manduca sexta L. parasitized by Cotesia congregata (Say).

This investigation examined the influence of dietary protein and carbohydrate balance in a chemically defined artificial diet for Manduca sexta larvae on development of the gregarious parasite Cotesia congregata. Normal unparasitized larvae and larvae superparasitized in the fourth stadium were reared to the end of the fifth stadium on six diets, each having the same total amount of casein and sucrose but with different ratios ranging from high protein/no carbohydrate through to low protein/high carbohydrate. Levels of blood protein nitrogen and trehalose, nutrients supporting growth and development of C. congregata, varied with diet and were influenced by parasitism. Different levels of blood metabolites reflected differences in diet consumption, and the relationships between protein nitrogen and trehalose were very similar to those for protein and carbohydrate intake by parasitized and normal larvae on various diets. Dietary nutrient ratio had a significant effect on parasite burden, the numbers of parasites developing in individual host larvae and on parasite biomass. Parasites included individuals that developed and eventually emerged as second instar larvae, moulted to third instars and pupated. Many apparently mature second instar parasites, however, failed to emerge. The proportion of non-emerging individuals varied with diet, and in some cases, parasites failing to emerge were greater in number and total biomass than those that did emerge to complete development. On most diets, the mass of individual parasites was similar regardless of dietary nutrient ratio. Three dimensional models developed to demonstrate the relationships between blood protein nitrogen and trehalose levels and parasite burden and biomass established that the levels of both metabolites are important for supporting growth and development of emerged and non-emerged parasites. In the case of emerged parasites, however, the relationships are linear, and a quadratic function best describes the relationships with non-emerged parasites. Blood metabolite levels supporting the greatest parasite burden and biomass of emerged and non-emerged parasites occupy a region of two dimensional space corresponding to approximately 60-200 mg per insect of protein nitrogen and 60-100 mg per insect of trehalose. Despite the differences in the response of emerged and non-emerged parasites to host nutrition, the present results indicate that host nutrition is not the critical factor determining parasite emergence. The significance of these findings to the biology of C. congregata is discussed.

Nutrition interacts with parasitism to influence growth and physiology of the insect Manduca sexta L.

The influence and interaction of dietary protein:carbohydrate balance and parasitism by Cotesia congregata on nutrient intake and growth were examined over the last two larval stadia of Manduca sexta. Effects of nutritional status on host blood metabolite concentrations were also determined. Six fat-free chemically defined diets were tested, each having the same total level of casein and sucrose, but with casein to sucrose ratios varying from low protein/high carbohydrate to equal levels of both nutrients through to high protein/no carbohydrate. Nutrient ratio and parasitism each affected nutrient consumption and growth. Feeding responses differed between normal and parasitized larvae, as illustrated by nutrient arrays, two-dimensional plots of protein and carbohydrate consumption on diets having different nutrient ratios. Normal larvae consumed more nutrients and took longer to develop as dietary nutrient ratio was displaced from equal levels of both nutrients. Except on the diet having the same amount of protein and carbohydrate, parasitized larvae consumed less nutrients than normal larvae, although on all diets parasitized larvae took longer to develop. When the contribution of parasite biomass was excluded, parasitized larvae showed lower mass gain than normal larvae on all diets. Total mass gain by normal and parasitized larvae with parasite biomass included, however, was similar on diets having intermediate nutrient ratios. Differences in mass gain between diets relative to nutrient consumption were evident from multi-dimensional representations of mass gain with protein and carbohydrate consumption. Three-dimensional plots and contour maps of normal and parasitized larvae were different. When differences in nutrient consumption between diets were taken into account, protein consumption had a greater effect on growth than carbohydrate consumption and normal larvae generally displayed greater mass gain than parasitized larvae on the same diets. Utilization efficiency, the efficiency of conversion of ingested food to body mass, was, therefore, generally reduced in parasitized insects. Concentrations of blood protein, total free amino acids and trehalose were each influenced by dietary nutrient ratio and parasitism. Concentrations of protein and free amino acids generally increased and trehalose concentration decreased as dietary protein increased and carbohydrate decreased. The opposite was the case as dietary carbohydrate increased and protein decreased. Dietary nutrient ratio, however, affected normal and parasitized larvae differently. Parasitized larvae had higher overall trehalose concentrations while normal larvae had higher protein and total free amino acid concentrations. When differences in nutrient consumption between diets were accounted for, protein consumption had a greater effect on blood protein and free amino acid concentrations than did dietary nutrient ratio or parasitism. Protein consumption, however, did not affect trehalose concentration. Carbohydrate consumption had no effect on the concentration of any of the metabolites after differences in nutrient consumption were taken into account. Effects of nutrient consumption on trehalose concentration, therefore, were due to dietary nutrient ratio and parasitism. The potential relevance of the above findings to the biology of parasitized M. sexta larvae is discussed.

The glucogenic response of a parasitized insect Manduca sexta L. is partially mediated by differential nutrient intake.

Induction of gluconeogenesis is accelerated in larvae of the insect Manduca sexta L. parasitized by Cotesia congregata (Say), maintaining the concentration of the blood sugar trehalose, an important nutrient for parasite development. Investigation has demonstrated that when host larvae are offered a choice of diets with varying levels of sucrose and casein, parasitized insects consume a different balance of these nutrients, principally due to a decrease in protein consumption. The result is metabolic homeostasis, with normal unparasitized and parasitized larvae exhibiting similar levels of gluconeogenesis and blood sugar level. In the present study, normal unparasitized and parasitized larvae were maintained on individual chemically defined diets having the balance of protein and carbohydrate consumed by each when offered a dietary choice. Total dietary nutrient, the sum of carbohydrate and protein, was provided at six levels, composed of three pairs of diets. Each diet pair consisting of diets having equivalent overall nutrient ratios of 2:1 and 1:1 casein/sucrose. Host growth and diet consumption were significantly affected by dietary nutrient level and the magnitude of these effects was influenced by parasitism. Due to the effects of dietary nutrient level on diet consumption, none of the unparasitized and parasitized larvae within any of the three diet pairs consumed protein and carbohydrate at the levels predicted by the earlier choice experiments. Among insects on all of the diets, however, two groups of unparasitized and parasitized larvae consumed the expected levels of protein and carbohydrate. In each case, gluconeogenesis, as measured by 13C nuclear magnetic resonance spectroscopy (NMR) analysis of pyruvate cycling and trehalose synthesis from [2-13C]pyruvate, was evident in unparasitized and parasitized insects, confirming the conclusions of the earlier experiments. Generally, all larvae that consumed less than approximately 250 mg of sucrose over the 3-day feeding period, were gluconeogenic, regardless of diet. Differential carbohydrate consumption, therefore, was an important factor in inducing gluconeogenesis in both unparasitized and parasitized insects. The selective 13C enrichment in trehalose displayed by non-gluconeogenic larvae on some diets demonstrated trehalose formation from [2]pyruvate. The absence of net carbohydrate synthesis in these insects was likely due to an elevation of glycolysis. There was no significant effect of diet consumption or parasitism on blood trehalose level. Parasitized larvae displayed higher levels of gluconeogenesis than did unparasitized insects, a finding consistent with the conclusion that blood sugar is rapidly sequestered by developing parasites. The parasite burden, the total number of parasites developing within host larvae, as well as the number of parasites emerging from host larvae to complete development, was significantly less at the lowest dietary nutrient level, but was otherwise similar at all dietary nutrient levels. Moreover, the number of parasites that emerged increased with increasing diet consumption as reflected by host final weight.

Effect of selected insecticides on Homalodisca coagulata (Homoptera: Cicadellidae) and transmission of oleander leaf scorch in a greenhouse study.

Homalodisca coagulata (Say) is a recent introduction to California. It is known to spread a strain of the bacterium Xylella fastidiosa Wells, Raju, Hung, Weisberg, Mandelco-Paul & Brenner that induces oleander leaf scorch disease in oleander, Nerium oleander L. Oleander leaf scorch is lethal to oleander and threatens to decimate one of the most important landscape shrubs in California. Towards developing a management strategy for H. coagulata-spread oleander leaf scorch, we documented the affects of selected insecticides on H. coagulata mortality, feeding behavior, and disease transmission in a greenhouse study. Oleanders treated with fenpropathrin, fenpropathrin + acephate, and imidacloprid caused significant mortality to caged H. coagulata within 4 h of exposure. Within 24 h, these pesticides caused nearly 100% mortality 3 wk after treatment. In other experiments, acetamiprid and fenpropathrin treatments reduced time spent feeding and total time on plants. H. coagulata on fenpropathrin-, acetamiprid-, and imidacloprid-treated oleander died in less than 13 min on average. Oleander leaf scorch transmission by H. coagulata was blocked by applications of foliar-applied acetamiprid, and soil-applied imidacloprid and thiamethoxam.

Altered dietary nutrient intake maintains metabolic homeostasis in parasitized larvae of the insect Manduca sexta L.

Manduca sexta larvae exhibited altered food selection over a 2- or 3-day feeding period when parasitized by Cotesia congregata, and offered a choice of two chemically defined diets, one containing casein without sucrose and a second with sucrose but no casein. While normal larvae consumed the diets in a ratio of approximately 2:1 protein:carbohydrate (w/w), parasitized insects consumed a ratio of approximately 1:1. The altered nutrient ratio consumed by parasitized insects was principally due to a decrease in consumption of the protein diet, and was only partially explained by their lower growth. Conditioning larvae for 1 day to either one of the choice diets had little effect on subsequent dietary intake over a 2-day feeding period. Conditioned larvae, regardless of parasitism, initially fed on the opposite diet immediately after conditioning. Although this suggests that the altered nutrient intake displayed by parasitized insects was not due to any failure in their capacity for dietary selection, these results do not definitively demonstrate an altered nutrient intake target by parasitized larvae. Rather, parasitism may compromise dietary selection, resulting in random feeding. When parasitized larvae were maintained on several isocaloric diets with a varying ratio of casein and sucrose, those larvae feeding on the diet with a ratio of 1:1 of these nutrients supported the largest parasite population. Previous investigation of larvae maintained on a single artificial diet established that parasitized insects display an aberrant induction of gluconeogenesis, so that haemolymph trehalose is maintained at a level equivalent to that of normal insects. In contrast, the present results demonstrated that parasitized larvae offered a choice of diets, and feeding at the altered nutrient ratio above, maintain haemolymph sugar but have the same level of gluconeogenesis as normal larvae given the same dietary choice. These investigations suggest that altered food selection by parasitized M. sexta larvae maintains metabolic homeostasis and, moreover, may be adaptive for C. congregata, potentially maximizing the number of parasites developing in a single host larva.

Seasonal flight activity of two Homalodisca species (Homoptera: Cicadellidae) that spread Xylella fastidiosa in southern California.

Homalodisca coagulata (Say) and Homalodisca lacerta (Fowler) are vectors of a new bacterial disease of oleander in California known as oleander leaf scorch, induced by the bacterium Xylella fastidiosa. H. coagulata also has been implicated in the spread of the strain of X. fastidiosa that induces Pierce's disease of grapevines in California. We monitored the flight activity of H. coagulata and H. lacerta in oleander and citrus by using yellow sticky cards at three southern California locations where outbreaks of oleander leaf scorch have been documented, and where vector compliments are different. Areas sampled included a mesic coastal area (Irvine, CA) that supports predominantly H. coagulata and few H. lacerta, a dry inland location (Palm Desert, CA) that supports predominantly H. lacerta and few H. coagulata, and an intermediate area (Riverside, CA) supporting both Homalodisca species. From November 1996 to October 1999 peak catches of both Homalodisca species occurred during the midsummer at all locations. H. coagulata was trapped in greater numbers in citrus than in oleander at both the Riverside and the Irvine sites. Likewise, H. lacerta in Riverside was more associated with citrus than oleander, yet H. lacerta in Palm Desert was trapped in greater numbers in oleander than citrus.

Interactions of dietary protein and carbohydrate determine blood sugar level and regulate nutrient selection in the insect Manduca sexta L.

The non-homeostatic regulation of blood sugar concentration in the insect Manduca sexta L. was affected by nutritional status. Larvae maintained on diets lacking sucrose displayed low concentrations of trehalose, the blood sugar of insects, which varied from 5 to 15 mM with increasing dietary casein level between 12.5 and 75 g/l. These insects were glucogenic, as demonstrated by the selective 13C enrichment of trehalose synthesized from [3-13C]alanine, and de novo synthesis was the sole source of blood sugar. The distribution of 13C in glutamine established that following transamination of the 13C substituted substrate, [3-13C]pyruvate carboxylation rather than decarboxylation was the principal pathway of Pyr metabolism. The mean blood trehalose level was higher in insects maintained on diets with sucrose. At the lowest dietary casein level blood trehalose was approximately 50 mM, and declined to 20 mM at the highest casein level. Gluconeogenesis was detected in insects maintained on sucrose-free diets at the higher protein levels examined, but [3-13C]pyruvate decarboxylation and TCA cycle metabolism was the principal fate of [3-13C]alanine following transamination, and dietary carbohydrate was the principal source of glucose for trehalose synthesis. Feeding studies established a relationship between nutritional status, blood sugar level and dietary self-selection. Insects preconditioned by feeding on diets without sucrose had low blood sugar levels regardless of dietary casein level, and when subsequently given a choice between a sucrose diet or a casein diet, selected the former. Larvae preconditioned on a diet containing sucrose and the lowest level of casein had high blood sugar levels and subsequently selected the casein diet. Larvae maintained on the sucrose diet with the highest casein level had low blood sugar and self-selected the sucrose diet. When preconditioned on diets with sucrose and intermediate levels of casein, insects selected more equally between the sucrose and the casein diets. It is concluded that blood sugar level may be intimately involved in dietary self-selection by M. sexta larvae, and that in the absence of dietary carbohydrate, gluconeogenesis provides sufficient blood sugar to ensure that larvae choose a diet or diets that produce an optimal intake of dietary protein and carbohydrate.

Interactions between the Encelia leaf beetle and its host plant, Encelia farinosa: The influence of acidic fog on insect growth and plant chemistry.

The impact of acidic deposition on interactions between the plant Encelia farinosa and the herbivorous beetle Trirhabda geminata (Chrysomelidae) was determined under greenhouse conditions. Acidic fogs (pH 2.75) did not significantly affect the overall foliar concentrations of water or soluble protein as compared with control fogs (pH 5.6). Nonetheless, E. farinosa foliage was altered by exposure to three 3-h acidic fogs such that growth and biomass gain by T. geminata increased by more than 30% as compared to beetles feeding on control-fogged plants. Thus, previous indications that changes in soluble proteins or water content were responsible for increased biomass gain and growth of T. geminata cannot be substantiated by this study. Additionally, changes in the plant defensive chemistry were not responsible for increased herbivore growth, as farinosin, encecalin, and euparin foliar concentrations did not vary significantly between fog treatments. Significant increases in CO2 assimilation rates of E. farinosa exposed to acidic fogs were documented at 3, 7, and 21 days following treatment, suggesting that carbohydrate-based products of increased plant metabolism may have played a role (e.g. soluble carbohydrates). However, the key factors responsible for increasing herbivore performance on acidic-fogged E. farinosa remain largely unknown.

Trirhabda geminata (Coleoptera: Chrysomelidae) resistance to the direct impact of simulated acidic fog on larval growth and mortality.

The direct effects of acidic fog (pH 2.75) upon the mortality and growth of Trirhabda geminata Horn (Coleoptera: Chrysomelidae), the dominant herbivore of the California coastal sage scrub, Encelia farinosa (Compositae: Asteraceae), were evaluated. Although there was a consistent pattern among and between experiments, suggesting that acidic fogs could reduce insect growth and survivorship within the first few days following application of treatments, an exposure to three consecutive, 3-h fogs over a five day period did not significantly affect mortality, biomass gain, or larval growth rate. There are two important implications from this study. First, even the highly acidic fogs found in southern California will have minimal direct effects on T. geminata performance. Second, the impacts on T. geminata biology observed in previous studies were likely mediated by host-plant responses to acidic-fog episodes.

Changes in western wheatgrass foliage quality following defoliation: consequences for a graminivorous grasshopper.

We determined the effects of defoliation by a graminivorous grasshopper on the foliage quality of the C3 plant, western wheatgrass (Pascopyrum smithii [Rydb] A. Love). Additionally, we determined the effects of this defoliation upon the subsequent feeding of the graminivorous grasshopper Phoetaliotes nebrascensis Thomas (Orthoptera: Acrididae). In field and greenhouse studies, graminivorous grasshopper herbivory altered the quality of remaining western wheatgrass foliage. In the greenhouse, severe (50% foliage removal) grasshopper grazing (638 grasshoppers/m2 for 72h) resulted in decreased foliar nitrogen (-12%), carbohydrate (-11%) and water (-2.5%) concentrations, and increased phenolic concentrations (+43%). These changes were associated with decreased adult female grasshopper mass gain, consumption rate, approximate digestibility, and food conversion efficiencies. In the field, moderate (14% foliage removal) grasshopper grazing (20 grasshoppers/m2 for 20 days) led to a 10% reduction in foliar nitrogen concentrations. Foliage quality changes in the field were not associated with any reductions in grasshopper mass gain, consumption rates, food digestibility, or conversion efficiencies. The results presented here are consistent with the hypothesis that defoliation leads to a reallocation of carbon and nitrogen compounds within the plant such that foliage quality for P. nebrascensis is reduced.

Impact of acidic deposition onEncelia farinosa gray (Compositae: Asteraceae) and feeding preferences ofTrirhabda geminata horn (Coleoptera: Chrysomelidae).

Container grownEncelia farinosa were exposed to three 3-hr episodes of acidic fog (pH 2.5) typical of events in southern California. Adults and larvae of the specialist leaf-feeding herbivore,Trirhabda geminata, preferred to feed on the acidic-treated foliage compared to control fogged (pH 6.3-6.5) foliage. Previous feeding damage on the plants did not affect feeding preference. The acidic-fogged foliage was significantly higher in total nitrogen and soluble protein but not different from control-treated tissue in water content. Stress on native populations of this drought-deciduous shrub caused by atmospheric pollutants may also result in altered feeding ecology of the beetle.

Responses of the western spruce budworm to varying levels of nitrogen and terpenes.

An agar diet study using western spruce budworm populations from Idaho and New Mexico was carried out to determine the effects of varying concentrations of nitrogen, beta-pinene, and bornyl acetate on larval growth and survival. Increased availability of nitrogen resulted in increased larval growth rate and survival to the adult stage. Larval growth rates from the Idaho population were higher on the high-nitrogen diet than were growth rates from the New Mexico population. The high level of beta-pinene improved larval growth at the high-nitrogen concentration. Bornyl acetate significantly reduced larval growth at both the low and high levels of nitrogen although the effect was greatest with the high-nitrogen diet. High bornyl acetate concentrations also significantly reduced survival to the adult stage. At high-nitrogen levels, a high concentration of bornyl acetate reduced larval growth rates and adult survival to a level similar to that occurring at the low nitrogen and low bornyl acetate concentrations.

Douglas-fir (Pseudotsuga menziesii)-spruce budworm (Choristoneura occidentalis) interactions: the effect of nutrition, chemical defenses, tissue phenology, and tree physical parameters on budworm success.

Measurements of foliage quality, physiological, and phenological condition of sample trees were used as independent variables in multiple correlation analyses to determine their effect on female and male spruce budworm larval dry weights. Female budworm from trees high in foliar concentrations of beta-pinene, myrcene and total nitrogen weighed less than those from trees lacking these characteristics. Male budworm from trees high in foliar concentrations of alpha-pinene, myrcene, terpinolene, citronellyl acetate, and bornyl acetate weighted less than those from trees lacking these characteristics. Additionally, relatively vigorous and productive trees tended to be less susceptible (as evidenced by reduced larval weight) to budworm of either sex.