Warming climate has increased access of native bark beetles to high-elevation pines that historically received only intermittent exposure to these tree-killing herbivores. Here we show that a dominant, relatively naïve, high-elevation species, whitebark pine, has inferior defenses against mountain pine beetle compared with its historical lower-elevation host, lodgepole pine. Lodgepole pines respond by exuding more resin and accumulating higher concentrations of toxic monoterpenes than whitebark pine, where they co-occur. Furthermore, the chemical composition of whitebark pine appears less able to inhibit the pheromonal communication beetles use to jointly overcome tree defenses. Despite whitebark pine's inferior defenses, beetles were more likely to attack their historical host in mixed stands. This finding suggests there has been insufficient sustained contact for beetles to alter their complex behavioral mechanisms driving host preference. In no-choice assays, however, beetles readily entered and tunneled in both hosts equally, and in stands containing less lodgepole pine, attacks on whitebark pines increased. High-elevation trees in pure stands may thus be particularly vulnerable to temperature-driven range expansions. Predators and competitors were more attracted to volatiles from herbivores attacking their historical host, further increasing risk in less coevolved systems. Our results suggest cold temperatures provided a sufficient barrier against herbivores for high-elevation trees to allocate resources to other physiological processes besides defense. Changing climate may reduce the viability of that evolutionary strategy, and the life histories of high-elevation trees seem unlikely to foster rapid counter adaptation. Consequences extend from reduced food supplies for endangered grizzly bears to altered landscape and hydrological processes.climate change | coevolution | disturbance | plant-insect interactions | forest insects

Aim  Bark beetle outbreaks have recently affected extensive areas of western North American forests, and factors explaining landscape patterns of tree mortality are poorly understood. The objective of this study was to determine the relative importance of stand structure, topography, soil characteristics, landscape context (the characteristics of the landscape surrounding the focal stand) and beetle pressure (the abundance of local beetle population eruptions around the focal stand a few years before the outbreak) to explain landscape patterns of tree mortality during outbreaks of three species: the mountain pine beetle, which attacks lodgepole pine and whitebark pine; the spruce beetle, which feeds on Engelmann spruce; and the Douglas‐fir beetle, which attacks Douglas‐fir. A second objective was to identify common variables that explain tree mortality among beetle–tree host pairings during outbreaks. Location  Greater Yellowstone ecosystem, Wyoming, USA. Methods  We used field surveys to quantify stand structure, soil characteristics and topography at the plot level in susceptible stands of each forest type showing different severities of infestation (0–98% mortality; n= 129 plots). We then used forest cover and beetle infestation maps derived from remote sensing to develop landscape context and beetle pressure metrics at different spatial scales. Plot‐level and landscape‐level variables were used to explain outbreak severity. Results  Engelmann spruce and Douglas‐fir mortality were best predicted using landscape‐level variables alone. Lodgepole pine mortality was best predicted by both landscape‐level and plot‐level variables. Whitebark pine mortality was best – although poorly – predicted by plot‐level variables. Models including landscape context and beetle pressure were much better at predicting outbreak severity than models that only included plot‐level measures, except for whitebark pine. Main conclusions  Landscape‐level variables, particularly beetle pressure, were the most consistent predictors of subsequent outbreak severity within susceptible stands of all four host species. These results may help forest managers identify vulnerable locations during ongoing outbreaks.

Bark beetle outbreaks and wildfire are important disturbances in conifer ecosystems, yet their interactions are not well understood. We evaluated whether fire injury increased susceptibility of lodgepole pines (Pinus contorta) to mountain pine beetle (Dendroctonus ponderosae Hopkins), how it influenced beetle reproductive success, and whether beetle population phase altered this interaction. Eight sites that experienced wildfire and eight unburned sites were examined in the Greater Yellowstone Ecosystem (USA). Half were in areas where D. ponderosae was undergoing outbreaks, and half were in areas with low populations. We examined 2056 trees one year after fire for burn injury and beetle attack. We quantified beetle reproductive success in a random sample of 106 trees, and measured gallery areas of D. ponderosae and competing subcortical herbivores in 79 additional trees. Baited flight traps sampled stand-level populations of subcortical herbivores and predators.Wildfire predisposed trees to D. ponderosae attack, but nonlinearly, with moderately injured trees being most preferred. This tree-level interaction was influenced by stand-level beetle population size, in that both healthy and fire-injured trees of all classes were attacked where populations were high, but no healthy trees, and only low and moderately injured trees were killed where populations were low. The number of adult brood produced per female was likewise curvilinear, being highest in moderately injured trees. This reflected an apparent trade-off, with high intraspecific competition arising from the large number of beetles needed to overcome defenses in healthy trees, vs. high interspecific competition and low substrate quality in more injured trees.These results suggest that fire-injured trees can provide a reservoir for D. ponderosae during periods when populations are too low to overcome defenses of healthy trees, and might otherwise face localized extinction. However, the likelihood of populations increasing from endemic to outbreak levels in response to increased susceptibility is offset by the opposing constraints of lower substrate quality and higher competitor load in severely injured hosts, and the relative scarcity of moderately injured trees. Wildfire may confer some reproductive increases to populations already outbreaking. We present a conceptual model of how these disturbances and inherent feedbacks interact to affect beetle population dynamics.

We examined the effect of wildfire injury on lodgepole pine chemical defenses against mountain pine beetle. We compared the constitutive phloem chemistry among uninjured, lightly-, moderately-, and severely-injured trees, and the induced chemistry elicited by simulated beetle attack, among these same categories. We also compared the entry rates of caged female beetles into trees of these categories. The volatiles we studied included thirteen monoterpene hydrocarbons, four allylic monoterpene alcohols, one ester, and one phenyl propanoid, of which the monoterpene hydrocarbons always comprised 96% or more of the total. Fire injury reduced the total concentration of these compounds in the induced but not constitutive phloem tissue of lodgepole pines. Fire injury also affected the relative composition of some volatiles in both induced and constitutive phloem. For example, increased fire injury reduced 4-allylanisole, a host compound that inhibits mountain pine beetle aggregation. Increased fire injury also increased (-) α-pinene, which can serve as precursor of pheromone communication. However, it also reduced myrcene and terpinolene, which can serve as stimulants and synergists of pheromone communication. Beetle entry did not show statistical differences among fire injury categories, although there was a trend to increased entry with fire injury. These results suggest that the reduced ability of trees to mobilize induced chemical defenses is an important mechanism behind the higher incidence of attack on fire-injured trees in the field. Future studies should concentrate on whether beetles that enter fire-injured trees are more likely to elicit aggregation, based on the differences we observed in volatile composition.

Cotesia flavipes (Hymenoptera:Braconidae) is a gregarious endoparasitoid of several pyralid stemborer larvae of economic significance including the sugarcane borer, Diatraea saccharalis. In this study, the ability of this parasitoid to develop in a sphingid host, Manduca sexta, was tested. First, second, third, fourth, and even pharate fifth instar host tobacco hornworm larvae were readily parasitized by the female C. flavipes parasitoids but no wasp larvae hatched from the eggs in this refractory host. Instead, the parasitoid eggs were invariably encapsulated by the host's hemocytes and, ultimately, no parasitoids emerged from tobacco hornworm hosts. The first stages of encapsulation were evident at 2 h post-parasitization of the host M. sexta larvae, when the beginning stages of capsule formation were seen. The developmental fate of the host larvae with encapsulated parasitoids was variable. Most succumbed as abnormally small fifth instars or as post-wandering prepupal animals, while a few developed normally to the pupal stage. Dissection of all the larvae or pupae with encapsulated wasp eggs showed evidence of hemocytic encapsulation and melanization of the C. flavipes eggs. This report describes the association between C. flavipes and M. sexta, which appears to be an excellent model system for studying the physiological processes accompanying wasp egg encapsulation that result in death of the host as well as the parasitoid. Since the parasitoid egg never hatches, the system offers an excellent opportunity to identify and study the effects of parasitoid-injected polydnavirus and venom on host physiology.

Wildfire and bark beetle epidemics are two ecologically important natural disturbances in the Intermountain West, yet we know very little about how these two phenomena interact. It is widely believed that beetle-killed trees increase the risk of severe fires; and trees that are weakened, but not killed by fire, are thought to be more susceptible to beetle invasion. However, few studies have rigorously tested these hypotheses. The GYE is currently experiencing an outbreak of unprecedented intensity and complexity, involving several species of bark beetles, including the mountain pine beetle. The outbreak is affecting multiple species of coniferous trees in and near recently burned areas, providing a timely opportunity to investigate these interactions at multiple scales.

Population eruptions by native bark beetles are intermittent and can cause wide spread forest disturbance. The mountain pine beetle (MPB), Dendroctonus ponderosae Hopkins, is one such bark beetle currently affecting all species of pine in the Greater Yellowstone Ecosystem (GYE). MPB populations erupt after generating positive feedback once thresholds are surpassed. Population eruptions require a combination of factors such as favorable weather, tree susceptibility, and the reduced competitors and predators.

We examined whether wildfire injury increased lodgepole pine, Pinus contorta, susceptibility to mountain pine beetle, Dendroctonus ponderosae, how it affects beetle reproduction, whether this interaction differs between endemic and epidemic populations, and how wildfire influences tree defense physiology. Wildfire predisposed trees to mountain pine beetle attack. In particular, fire-injured trees had a lower ability in synthesized monoterpenes in response to simulated attacks than did non-injured trees. However, beetles responded in a non-linear fashion; moderately-injured trees were most preferred. This interaction was influenced by beetle population size. Healthy and fire-injured trees were attacked when populations were high, but no healthy trees and no severely-injured trees were killed when populations were low. Beetle brood production per female was also curvilinear being highest in moderately-injured trees. This reflected a trade-off between high intraspecific competition arising from the large number of beetles needed to overcome defenses in healthy trees, and high interspecific competition and low substrate quality in severely injured trees. These results suggest that fire-injured trees can provide a resource for mountain pine beetles during the extended periods when populations are not high enough to overcome defenses of vigorous trees. But the likelihood that populations could transition from endemic to epidemic levels due to increased tree susceptibility from wildfire is constrained by the opposing factors of lower nutritional quality and more competition load in severely-injured trees, and the relatively low incidence of moderately-injured trees. Wildfire may cause some reproductive increases in populations that are already in outbreak mode.