S tudies of the effects of climate change on forests have focused on the ability of species to tolerate temperature and moisture changes and to disperse, but they have ignored the effects of disturbances caused by climate change (e.g., Ojima et al. 1991). Yet modeling studies indicate the importance of climate effects on disturbance regimes (He et al. 1999). Local, regional, and global changes in temperature and precipitation can influence the occurrence, timing, frequency, duration, extent, and intensity of disturbances (Baker 1995, Turner et al. 1998). Because trees can survive from decades to centuries and take years to become established, climate-change impacts are expressed in forests, in part, through alterations in disturbance regimes (Franklin et al. 1992, Dale et al. 2000). Disturbances, both human-induced and natural, shape forest systems by influencing their composition, structure, and functional processes. Indeed, the forests of the United States are molded by their land-use and disturbance history. Within the United States, natural disturbances having the greatest effects on forests include fire, drought, introduced species, insect and pathogen outbreaks, hurricanes, windstorms, ice storms, and landslides (Figure 1). Each disturbance affects forests differently. Some cause large-scale tree mortality, whereas others affect community structure and organization without causing massive mortality (e.g., ground fires). Forest disturbances influence how much carbon is stored in trees or dead wood. All these natural disturbances interact with human-induced effects on the environment, such as air pollution and land-use change resulting from resource extraction, agriculture, urban and suburban expansion, and recreation. Some disturbances can be functions of both natural and human conditions (e.g., forest fire ignition and spread) (Figure 2).

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...

We characterized the structure of condensed tannins from 16 woody plant species (seven genera, six families) and determined their effects on six herbivorous insect species (four genera, two families). There were major differences in tannin structure, even between congeneric plant species. Condensed tannins differed markedly in their antiherbivore activity, averaged over these herbivores, and the herbivores differed in their sensitivity, averaged over these tannins. Furthermore, the same tannin can have different effects on different herbivores, presumably because of interactions between tannin structure and gut physiology. Results challenge the view that tannins provide an evolutionarily stable plant defense because of their uniform chemical properties. Condensed tannin can sometimes impact herbivore fitness through effects on survival and growth, but the largest effects in 45 insect-tannin combinations were less than that of many other plant metabolites at lower doses. Even at high doses, condensed tannins frequently had no strong antiherbivore activity, even against insects with no evolutionary history of encountering the tannin (Ͻ10% reduction in growth rate in 24 of 45 experiments). Most condensed tannins apparently do not have broad-spectrum antiherbivore activity. We doubt that selective pressures from folivorous insects can be the main explanation for the diversion of so much carbon, in so many plant species, into the synthesis of condensed tannins.

The bark beetles Ips pini, I. perroti, and I. grandicollis are sympatric in pine forests of the north-central United States. They share the same limited phloem resource and often coexist within the same host trees. We tested whether phloem resources are partitioned in time and space by measuring spatial and seasonal colonization of logs. Differences among species in flight phenology, development time, voltinism, and spatial colonization patterns within logs reduce, but do not eliminate, species overlap. The bark beetle species share predation by Thanasimus dubius (Cleridae) and Platysoma cylindrica (Histeridae), which exploit pheromone signals for prey location. We employed pheromone traps to test for chemical communication among bark beetle species. Heterospecific signals tend to be deterrents when they are added to conspecific signals but attractants when they are alone, indicating that the communication system can both reduce and increase species overlap in resource use depending upon relative abundance of the species. Deterrence by heterospecific signals is probably a result of selection for minimizing interspecific competition. However, individuals may sometimes benefit from joining aggregations of other species because of (1) predator swamping, (2) improved success in attacking live trees, and (3) location of suitable, recently dead, trees. These benefits should be greatest for males (which locate and colonize host trees before signalling females) and indeed males tended to be more attracted than females by heterospecific signals. Shared resources, shared predators, and heterospecific pheromone communication all contribute to species interactions in this guild of bark beetles, but predicting whether the removal of one species will tend to increase or decrease the abundance of remaining species remains difficult. Species interactions are likely conditional and coexistence is probably promoted by benefits to rare species of multispecies associations.

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