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The nitrogen content of plant tissue is low relative to that of herbivores; as a consequence, dietary N can limit the growth and reproduction of herbivores and select for attributes that increase N acquisition. Bark beetles face a particularly severe challenge because the phloem that they consume is very low in nitrogen and phosphorus relative to their requirements. We quantified variation in the phloem concentrations of N and P in the host tree, Pinus taeda, and evaluated the following hypotheses regarding the role of symbiotic fungi in nutrient budgets of the herbivore Dendroctonus frontalis: D. frontalis experience variation in phloem nutrient concentrations across several spatial scales (H1); mycangial fungi enhance the diet of D. frontalis larvae by contributing to the acquisition of N and P (H2); Ophiostoma minus, an apparently antagonistic fungal symbiont, hinders D. frontalis larvae because it does not enhance nutrient concentrations of the phloem as much as mycangial fungi do (H3); and larvae of bark beetle species that lack mycangial fungi must consume more phloem to accomplish the same growth as larvae of D. frontalis (H4). In addition, we developed a general model for the N budgets of herbivorous insects that identifies the possible combinations of dietary and physiological parameters that can allow developmental success on low‐nutrient diets. Spatial variation in phloem N was mostly at the level of trees within sites (a scale of meters) while P mostly varied among sites (a scale of kilometers). Trees with higher N content produced larger D. frontalis adults. Prior to infestation by beetles, phloem nutrient concentrations were very uniform within trees and very low relative to that of the bark beetles (N and P concentrations of D. frontalis adults were 28 and 8 times greater, respectively). During infestation, phloem nutrient concentrations increased overall and became highly variable within trees. Nitrogen concentrations increased from 0.40 ± 0.01% (mean ± 1 se) in uninfested phloem to 0.86 ± 0.03% in the phloem surrounding successfully developing D. frontalis larvae, which are typically associated with one or two species of mutualistic mycangial fungi. Nitrogen concentrations were intermediate in other microhabitats within infested trees, including regions with no adult colonization, with failed larval development, or colonized by the antagonistic bluestain fungus O. minus. We parameterized a general nutrient‐budget model for D. frontalis and a sympatric non‐mycangial bark beetle, Ips grandicollis, which indicated that (1) mycangial fungi provide their benefits by concentrating dietary N for larvae; (2) O. minus may exert its antagonistic effects on D. frontalis larvae by failing to concentrate dietary N as much as mycangial fungi do; (3) non‐mycangial bark beetles meet their N budgets through high consumption of unaltered, low‐N phloem; and (4) larvae should easily meet their P requirements with any combination of consumption rate and development time that allows them to meet their N requirements. A...

The ecological literature abounds with studies of environmental effects on plant antiherbivore defences. While various models have been proposed (e.g. plant stress, optimal allocation, growth‐differentiation balance), each has met with mixed support. One possible explanation for the mixed results is that constitutive and induced defences are differentially affected by environmental conditions. In this study, constitutive oleoresin flow from Pinus tadea was least during periods of rapid tree growth and most when drought conditions limited growth; this is as expected if constitutive secondary metabolism is a function of the carbohydrate pool size after growth has been maximised. Induced increases in resin flow, however, were greatest in the fastest growing trees during the season of greatest growth. Apparently, resin production becomes an allocation priority after wounding but not before. Understanding environmental effects on plant antiherbivore defences requires physiological and evolutionary models that account for the differences between constitutive and induced secondary metabolism.

The southern pine beetle, Dendroctonus frontalis, is among the most important agents of ecological disturbance and economic loss in forests of the south-eastern United States. We combined physiological measurements of insect temperature responses with climatic analyses to test the role of temperature in determining the northern distribution limits of D. frontalis. Laboratory measurements of lower lethal temperatures and published records of mortality in wild populations indicated that air temperatures of −16°should result in almost 100% mortality of D. frontalis. The distribution limits for D. frontalis approximate the isoline corresponding to an annual probability of 0.90 of reaching ≤−16°C. Thus, D. frontalis have been found about as far north as they could possibly occur given winter temperature regimes. At latitudes from 39°N (southern Ohio) to 33°N (central Alabama), winter temperatures must exert high mortality on D. frontalis populations in at least one year out of ten. In contrast, we reject the hypotheses that summer temperatures or the distribution of host trees constrain the northern distribution of D. frontalis. Because of the short generation time of D. frontalis, its high dispersal abilities, and the cosmopolitan distribution of suitable host trees, changes in either the mean or variance of minimum annual temperatures could have almost immediate effects on regional patterns of beetle infestations. We estimate that an increase of 3°C in minimum annual temperature could extend the northern distribution limits by 170 km. Increases or decreases in the variance of minimum annual temperatures would further relax climatic constraints on the northern distribution limits of D. frontalis. Results emphasize the ecological importance of spatial and temporal variability in minimum annual temperatures. The physiologically based models provide a tool for guiding land management decisions in forests and illustrate a general approach for predicting the regional effects of climatic patterns on the distribution of organisms.

Strong indirect interactions of Tarsonemus mites (Acarina: Tarsonemidae) and Dendroctonus frontalis (Coleoptera: Scolytidae). -Oikos 102: 243-252. Phoretic mites of bark beetles are classic examples of commensal ectosymbionts.However, many such mites appear to have mutualisms with fungi that could themselves interact with beetles. We tested for indirect effects of phoretic mites on Dendroctonus frontalis, which attacks and kills pine trees in North America. Tarsonemus mites are known to carry ascospores of Ophiostoma minus, which tends to outcompete the mutualistic fungi carried by D. frontalis. Experimental additions and removals of mites from beetles demonstrated that Tarsonemus propagate O. minus in beetle oviposition galleries. Furthermore, the abundance of Tarsonemus and O. minus tended to covary in nature. These results verified a strong mutualism between Tarsonemus and O. minus. Results also indicated that O. minus is an antagonist of D. frontalis: beetle larvae seldom survived in the presence of O. minus (compared to 83% survival elsewhere). Apparently, this is an indirect result of O. minus outcompeting the two species of mycangial fungi that are critical to beetle nutrition. Thus, Tarsonemus mites close a loop of species interactions that includes a commensalism (mites and beetles), a mutualism (mites and O. minus), asymmetric competition (O.minus and mycangial fungi), and another mutualism (mycangial fungi and beetles). This interaction system produces negative feedback that could contribute to the endogenous population dynamics of D. frontalis. Reproductive rate of Tarsonemus was more temperature-sensitive than beetle generation time (which constrains the time for mite reproduction within a tree). This differential temperature sensitivity produces a narrow range of temperatures (centred at 27°C) in which mite reproduction per D. frontalis generation can attain its maximum of 100 mites/beetle. Consequently, seasonal oscillations in temperature are predicted to produce oscillations in the D. frontalis community, and climatic differences between regions could influence the community to dampen or exacerbate the cyclical outbreak dynamics of D. frontalis.