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Unlike most phytophagous insects, the reproduction of primary bark beetles (Coleoptera: Scolytidae) is contingent on host mortality. Consequently, there have been intense selective pressures on trees for properties which confer resistance to attack, and likewise, on the insect for increased behavioral complexity by which to overcome these defenses. In this study, we examined the relationship between the physiology of Pinus contorta var. latifolia and the behavior of the bark beetle, Dendroctonus ponderosae. Host mortality is a discrete outcome which is contingent on the quantitative interaction between host resistance and beetle numbers. At low attack densities, trees respond by confining beetles and their associated fungi within necrotic lesions containing toxic or inhibitory compounds. Beyond a critical "threshold of attack," however, the defensive capacity of the tree is exhausted, and mortality occurs. This threshold occurred at °40 galleries/m2 in our experimental stands. The reproductive success of the bark beetle is directly related to the depletion of host defenses through concentrated attack. At sufficiently high attack densities, °80 galleries/m2, the potential suppressive effect of the host on brood development is not manifested. Increased beetle numbers, however, cause an exponential decline in brood production due to intraspecific competition. Consequently, there is an optimal density, °62 attacks/m2 at which reproduction and survival of the beetle is maximized. Dendroctonus ponderosae utilize defensive secretions as precursors and synergists of aggregation pheromones, and thereby, elicit maximum responses from flying beetles while the host's production of toxins is at its maximum. This mechanism also ensures that the attack will terminate once the tree has been rendered suitable for brood development and, thus, minimizes the deleterious effect of intraspecific competition. Such an interaction, in which all trees could theoretically be colonized regardless of their resistance capacity, cannot result in a stable host—parasite system. Conifers, however, seem to be able to interfere with bark beetle communication. Pheromone synthesis within the hindgut and emission from the entrance site are distinct events, and beetles contained within a copious flow of resin seem unable to elicit attraction. This ability to interrupt the beetles' communication sequence, however, is inversely related to the number of beetles initiating localized attacks. Consequently, the colonization sequence reflects the outcome of a dynamic interaction between the tree and the initial "pioneer" beetles.

Scolytid bark beetles that colonize living conifers are frequently associated with specific fungi that are carried in specialized structures or on the body surface. These fungi are introduced into the tree during the attack process. The continuing association suggests that there is mutual benefit to the fitness of both beetles and fungi. The fungal species may benefit from the association with the beetles by transport to new host trees. Beetle species may benefit from the association with fungi by feeding on the fungi, or by the fungi contributing to the death of the host trees through mycelial penetration of host tissue, toxin release, interactions with preformed and induced conifer defenses, or the combined action of both beetles and fungi during colonization. Extensive research has been directed towards characterizing the interactions of beetle-fungal complexes with live host conifers and determining the ecological advantages for maintaining the associations. However, differences among systems and how species interact under different population and environmental conditions make it difficult to generalize about the importance of the separate biological components in successful host colonization.

SummaryClimate change is expected to drive increased tree mortality through drought, heat stress, and insect attacks, with manifold impacts on forest ecosystems. Yet, climate-induced tree mortality and biotic disturbance agents are largely absent from process-based ecosystem models. Using data sets from the western USA and associated studies, we present a framework for determining the relative contribution of drought stress, insect attack, and their interactions, which is critical for modeling mortality in future climates. We outline a simple approach that identifies the mechanisms associated with two guilds of insects -bark beetles and defoliators -which are responsible for substantial tree mortality. We then discuss cross-biome patterns of insect-driven tree mortality and draw upon available evidence contrasting the prevalence of insect outbreaks in temperate and tropical regions. We conclude with an overview of tools and promising avenues to address major challenges. Ultimately, a multitrophic approach that captures tree physiology, insect populations, and tree-insect interactions will better inform projections of forest ecosystem responses to climate change.