Purpose of Review Climate change affects populations of forest insect pests in a number of ways. We reviewed the most recent literature (2013-2017) on this subject including previous reviews on the topic. We provide a comprehensive discussion of the subject, with special attention to insect range expansion, insect abundance, impacts on forest ecosystems, and effects on forest insect communities. We considered forest insects according to their major guilds and biomes. Recent Findings Effects of climate change on forest insects are demonstrated for a number of species and guilds, although generalizations of results available so far are difficult because of species-specific responses to climate change. In addition, disentangling direct and indirect effects of climate change is complex due to the large number of variables affected. Modeling based on climate projections is useful when combined with mechanistic explanations. Summary Expansion of either the true range or the outbreak range is observed in several model species/groups of major insect guilds in boreal and temperate biomes. Mechanistic explanations are provided for a few species and are mainly based on increase in winter temperatures. In relation to insect abundance, climate change can either promote outbreaks or disrupt trophic interactions and decrease the severity of outbreaks. There is good evidence that some recent outbreaks of bark beetles and defoliating insects are influenced by climate change and are having a large impact on ecosystems as well as on communities of forest insects.
Climate change is altering insect disturbance regimes via temperature-mediated phenological changes and trophic interactions among host trees, herbivorous insects, and their natural enemies in boreal forests. Range expansion and increase in outbreak severity of forest insects are occurring in Europe and North America. The degree to which northern forest ecosystems are resilient to novel disturbance regimes will have direct consequences for the provisioning of goods and services from these forests and for long-term forest management planning. Among major ecological disturbance agents in the boreal forests of North America is a tortricid moth, the eastern spruce budworm, which defoliates fir (Abies spp.) and spruce (Picea spp.). Northern expansion of this defoliator in eastern North America and climateinduced narrowing of the phenological mismatch between the insect and its secondary host, black spruce (Picea mariana), may permit greater defoliation and mortality in extensive northern black spruce forests. Although spruce budworm outbreak centers have appeared in the boreal black spruce zone historically, defoliation and mortality were minor. Potential increases in outbreak severity and tree mortality raise concerns about the future state of this northern ecosystem. Severe spruce budworm outbreaks could decrease stand productivity compared with their occurrence in more diverse, southern balsam fir forest landscapes that have coevolved with outbreaks. Furthermore, depending on the proportion of balsam fir and deciduous species present and fire recurrence, changes in regeneration patterns and in nutrient cycling could alter ecosystem dynamics and replace black spruce by more productive mixedwood forest, or by less productive ericaceous shrublands. Long-term monitoring, manipulative experiments, and process modeling of climate-induced phenological changes on herbivorous insect pests, their host tree species, and natural enemies in northern forests are therefore crucial to predicting species range shifts and assessing ecological and economic impacts.
Three main hypotheses have been postulated over the past century to explain the outbreaking population dynamics of eastern spruce budworm, Choristoneura fumiferana (Clemens). The Silviculture Hypothesis first arose in the 1920s, with the idea that outbreaks were driven by forestry practices favoring susceptible softwood species. In the 1960s, it was proposed that populations were governed by Multiple Equilibria, with warm weather conditions releasing low-density populations from the regulatory control of natural enemies. Dispersal from outbreak foci, or "epicenters," was seen as causing widespread outbreaks that eventually collapsed following resource depletion. However, in the 1980s, following the re-analysis of data from the 1940s outbreak in New Brunswick, this interpretation was challenged. The alternative Oscillatory Hypothesis proposed that budworm population dynamics were governed by a second-order density-dependent process, with oscillations being driven by natural enemy-victim interactions. Under this hypothesis, weather and resource availability contribute to secondary fluctuations around the main oscillation, and weather and moth dispersal serve to synchronize population cycles regionally. Intensive, independent population studies during the peak and declining phases of the 1980s outbreak supported the principal tenet of the Oscillatory Hypothesis, but concluded that host plant quality played a more important role than this hypothesis proposed. More recent research on the early phase of spruce budworm cycles suggests that mate-finding and natural-enemy-driven Allee effects in low-density populations might be overcome by immigration of moths, which can facilitate the onset of outbreaks. Even more recent research has supported components of all three hypotheses attempting to explain spruce budworm dynamics. In the midst of a new rising outbreak (2006-present), we discuss the evolution of debates surrounding these hypotheses from a historic perspective, examine gaps in current knowledge, and suggest avenues for future research (e.g., intensive studies on low-density populations) to better understand and manage spruce budworm populations.
Previous research indicated that the aggregation pheromone of the southern pine beetle, Dendroctonus frontalis, is produced only by females, the sex that initiates attacks. We provide evidence indicating that secondarily arriving males augment mass aggregation by releasing the attractive synergist (+)-endo-brevicomin. Healthy pines artificially infested with both sexes of D. frontalis were significantly more attractive to conspecifics than trees infested solely with females. Coupled gas chromatography-electroantennographic detection (GC-EAD) analyses of volatiles isolated from male beetles revealed substantially greater olfactory sensitivity by D, frontalis to endo-brevicomin than to any other component. The threshold of detection of both sexes for (+)-endo-brevicomin was four orders of magnitude lower than for its antipode and at least one order of magnitude lower than for either enantiomer of frontalin, the major female-produced aggregation pheromone component. Pairing with a female in a gallery stimulated individual male beetles to produce hundreds of nanograms of (+)-endo-brevicomin. (4-)-endo-Brevicomin was detected in a small percentage of female D. frontalis, whereas (-)-endo-brevicomin was never detected in either sex. In field trapping bioassays, we confirmed that (+)-endo-brevicomin is a potent synergist for attractive combinations of frontalin and pine turpentine. However, (+)-endobrevicomin failed to attract D. frontalis either when presented alone or in combination with turpentine. We postulate that mass colonization of host trees by D. frontalis is mediated by distinct semiochemicals from both sexes rather than females alone. Our discovery of a key aggregation pheromone component in such an apparently well-studied species implies that the pheromone models of other bark beetles could benefit from systematic reexamination using newer technologies. Additionally, baits fortified with (+)-endo-brevicomin may enhance pest management strategies that exploit attractants for D. frontalis.
Disturbance regimes are changing in forests across the world in response to global climate change. Despite the profound impacts of disturbances on ecosystem services and biodiversity, assessments of disturbances at the global scale remain scarce. Here, we analyzed natural disturbances in boreal and temperate forest ecosystems for the period 2001–2014, aiming to 1) quantify their within‐ and between‐biome variation and 2) compare the climate sensitivity of disturbances across biomes. We studied 103 unmanaged forest landscapes with a total land area of 28.2 × 106 ha, distributed across five continents. A consistent and comprehensive quantification of disturbances was derived by combining satellite‐based disturbance maps with local expert knowledge of disturbance agents. We used Gaussian finite mixture models to identify clusters of landscapes with similar disturbance activity as indicated by the percent forest area disturbed as well as the size, edge density and perimeter–area‐ratio of disturbed patches. The climate sensitivity of disturbances was analyzed using Bayesian generalized linear mixed effect models and a globally consistent climate dataset. Within‐biome variation in natural disturbances was high in both boreal and temperate biomes, and disturbance patterns did not vary systematically with latitude or biome. The emergent clusters of disturbance activity in the boreal zone were similar to those in the temperate zone, but boreal landscapes were more likely to experience high disturbance activity than their temperate counterparts. Across both biomes high disturbance activity was particularly associated with wildfire, and was consistently linked to years with warmer and drier than average conditions. Natural disturbances are a key driver of variability in boreal and temperate forest ecosystems, with high similarity in the disturbance patterns between both biomes. The universally high climate sensitivity of disturbances across boreal and temperate ecosystems indicates that future climate change could substantially increase disturbance activity.
We studied the host selection behavior and feeding preference of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae). A. planipennis is an exotic forest insect pest native to Asia that was discovered in North America in 2002 and is causing widespread mortality of ash trees (Fraxinus spp.) in southeast Michigan and surrounding states. We compared host selection and feeding behavior on different species of ash including Manchurian ash (F. mandshurica Rupr.), green ash (F. pennsylvanica Marsh), white ash (F. americana L.), black ash (F. nigra Marsh), blue ash (F. quadrangulata Michx.), and European ash (F. excelsior L). Manchurian ash is native to Asia, whereas the other species (native to North America or Europe) represent novel hosts for A. planipennis. Beetles distributed themselves more frequently and fed to a greater extent on green, black, and white ash compared with blue, European, and Manchurian ash. Although beetles consumed every ash species offered to them, Manchurian ash and blue ash were least preferred in feeding bioassays. When we analyzed the volatile content of intact and girdled ash for quantitative variation in 11 compounds that elicited antennal activity, we found that the overall volatile profiles of the six ash species differed significantly in their relative amounts of antennally active compounds. Green ash has lower relative amounts of volatiles compared with Manchurian ash, which might render it more attractive and less resistant to A. planipennis. Lower tolerance and resistance of green ash might make it more susceptible to mortality compared with Manchurian ash, which coevolved with the beetle in its native range. Repellent odors, potential antifeedants, and genes for resistance in Manchurian ash could be explored for methods to manage A. planipennis populations.
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