The tree-killing mountain pine beetle (Dendroctonus ponderosae Hopkins) is an important disturbance agent of western North American forests and recent outbreaks have affected tens of millions of hectares of trees. Most western North American pines (Pinus spp.) are hosts and are successfully attacked by mountain pine beetles whereas a handful of pine species are not suitable hosts and are rarely attacked. How pioneering females locate host trees is not well understood, with prevailing theory involving random landings and/or visual cues. Here we show that female mountain pine beetles orient toward volatile organic compounds (VOCs) from host limber pine (Pinus flexilis James) and away from VOCs of non-host Great Basin bristlecone pine (Pinus longaeva Bailey) in a Y-tube olfactometer. When presented with VOCs of both trees, females overwhelmingly choose limber pine over Great Basin bristlecone pine. Analysis of VOCs collected from co-occurring limber and Great Basin bristlecone pine trees revealed only a few quantitative differences. Noticeable differences included the monoterpenes 3-carene and D-limonene which were produced in greater amounts by host limber pine. We found no evidence that 3-carene is important for beetles when selecting trees, it was not attractive alone and its addition to Great Basin bristlecone pine VOCs did not alter female selection. However, addition of D-limonene to Great Basin bristlecone pine VOCs disrupted the ability of beetles to distinguish between tree species. When presented alone, D-limonene did not affect behavior, suggesting that the response is mediated by multiple compounds. A better understanding of host selection by mountain pine beetles could improve strategies for managing this important forest insect. Moreover, elucidating how Great Basin bristlecone pine escapes attack by mountain pine beetles could provide insight into mechanisms underlying the incredible longevity of this tree species.
Abstract:Interior Douglas-fir is a prevalent forest type throughout the central Rocky Mountains. Past management actions, specifically fire suppression, have led to an expansion of this forest type. Although Douglas-fir forests cover a broad geographic range, few studies have described the interactive effects of various disturbance agents on forest health conditions. In this paper, we review pertinent literature describing the roles, linkages, and mechanisms by which disturbances, including insect outbreaks, pathogens, fire, and other abiotic factors, affect the development, structure, and distribution of interior montane forests primarily comprised of Douglas-fir. We also discuss how these effects may influence important resource values such as water, biodiversity, wildlife habitat, timber, and recreation. Finally, we identify gaps where further research may increase our understanding of these disturbance agents, their interacting roles, and how they influence long-term forest health.
Mass attack by tree-killing bark beetles (Curculionidae: Scolytinae) brings about large chemical changes in host trees that can have important ecological consequences. For example, mountain pine beetle (Dendroctonus ponderosae Hopkins) attack increases emission of terpenes by lodgepole pine (Pinus contorta Dougl. ex Loud.), affecting foliage flammability with consequences for wildfires. In this study, we measured chemical changes to Douglas-fir (Pseudotsuga menziesii var. glauca (Mirb.) Franco) foliage in response to attack by Douglas-fir beetles (Dendroctonus pseudotsugae Hopkins) as trees die and crowns transitioned from green/healthy, to green-infested (year of attack), to yellow (year after attack), and red (2 yr after attack). We found large differences in volatile and within-needle terpene concentrations among crown classes and variation across a growing season. In general, emissions and concentrations of total and individual terpenes were greater for yellow and red needles than green needles. Douglas-fir beetle attack increased emissions and concentrations of terpene compounds linked to increased tree flammability in other conifer species and compounds known to attract beetles (e.g., [Formula: see text]-pinene, camphene, and D-limonene). There was little relationship between air temperature or within-needle concentrations of terpenes and emission of terpenes, suggesting that passive emission of terpenes (e.g., from dead foliage) does not fully explain changes in volatile emissions. The potential physiological causes and ecological consequences of these bark beetle-associated chemical changes are discussed.
Downed woody material (DWM) is a key component in forest ecosystems with age, structure, and disturbance described as primary factors that influence DWM dynamics. In particular, much emphasis is placed on large coarse woody debris (CWD). Fine woody debris (FWD) (less than 7.62 cm diameter), duff, and litter also contribute to carbon stocks, provide habitat, add to nutrient cycling, and are often the most available fuels for fire, yet are regularly overlooked in studies describing the forest floor. Throughout the middle montane zone within the Intermountain West region USA, interior Douglas-fir (Pseudotsuga menziesii var. glauca Mirb. Franco) is a predominant forest type, yet little is known about the forest floor complex in these forests. We used a chronosequence approach to compare DWM patterns over the course of stand development among stands with different disturbance histories. Using classification and regression trees, we also evaluated an assemblage of environmental, structural, and disturbance variables to determine factors of most importance for estimating loading for DWM, duff, and litter. We found CWD resembled a U-shaped pattern of buildup while FWD components remained stable over the course of stand development regardless of disturbance history. Our results indicate that large DWM components are most closely associated with the amount of standing dead material in a stand, primarily the density and basal area of snags. Fine woody material was more aligned with live stand components, while duff and litter were more influenced by disturbance.
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