The southern pine beetle, Dendroctonus frontalis Zimmermann is an important mortality agent of Pinus in the eastern United States of America where it commonly shares hosts with the black turpentine beetle, Dendroctonus terebrans (Olivier), which infrequently kills trees. Unlike D. frontalis, which must kill its hosts to become established in the bark and reproduce, D. terebrans can occupy living hosts as a parasite. Olfactory mechanisms whereby D. frontalis initially locates hosts have not been demonstrated, whereas D. terebrans responds strongly to host odors. Because D. terebrans produces frontalin, the primary aggregation pheromone component for D. frontalis, and commonly arrives on hosts prior to D. frontalis, it has been hypothesized that D. terebrans pheromone components can mediate D. frontalis location of suitable, living trees. We assessed this possibility with studies of the semiochemical interactions between D. frontalis and D. terebrans. Coupled gas chromatography-electroantennographic detection analyses indicated that D. terebrans produces nine different olfactory stimulants for D. frontalis, nearly all of them known semiochemicals for D. frontalis. A trapping experiment designed to address the potentially confounding influence of lure contamination confirmed that the D. terebrans pheromone component exo-brevicomin enhances attraction of D. frontalis and thus could be an attractive kairomone. In ambulatory bioassays, male D. frontalis were strongly attracted to odors of frass of solitary female and paired D. terebrans, indicating their attraction to the naturally occurring semiochemicals of D. terebrans. Cues from D. terebrans may influence host and mate-finding success of D. frontalis and, thereby, the latter's virulence.
AbstractMacroclimate drives vegetation distributions, but fine‐scale topographic variation can generate microclimate refugia for plant persistence in unsuitable areas. However, we lack quantitative descriptions of topography‐driven microclimatic variation and how it shapes forest structure, diversity, and composition. We hypothesized that topographic variation and the presence of the forest overstory cause spatiotemporal microclimate variation affecting tree performance, causing forest structure, diversity, and composition to vary with topography and microclimate, and topography and the overstory to buffer microclimate. In a 20.2‐ha inventory plot in the North American Great Plains, we censused woody stems ≥1 cm in diameter and collected detailed topographic and microclimatic data. Across 59‐m of elevation, microclimate covaried with topography to create a sharp desiccation gradient, and topography and the overstory buffered understory microclimate. The magnitude of microclimatic variation mirrored that of regional‐scale variation: with increasing elevation, there was a decrease in soil moisture corresponding to the difference across ~2.1° of longitude along the east‐to‐west aridity gradient and an increase in air temperature corresponding to the difference across ~2.7° of latitude along the north‐to‐south gradient. More complex forest structure and higher diversity occurred in moister, less‐exposed habitats, and species occupied distinct topographic niches. Our study demonstrates how topographic and microclimatic gradients structure forests in putative climate‐change refugia, by revealing ecological processes enabling populations to be maintained during periods of unfavorable macroclimate.
Climate change is causing rapid shifts in the abiotic and biotic environmental conditions experienced by plant populations, but we lack generalizable frameworks for predicting the consequences for species. These changes may cause individuals to become poorly matched to their environments, potentially inducing shifts in the distributions of populations and altering species’ habitat and geographic ranges. We present a trade-off-based framework for understanding and predicting whether plant species may undergo range shifts, based on ecological strategies defined by functional trait variation. We define a species’ capacity for undergoing range shifts as the product of its colonization ability and the ability to express a phenotype well-suited to the environment across life stages (phenotype–environment matching), which are both strongly influenced by a species’ ecological strategy and unavoidable trade-offs in function. While numerous strategies may be successful in an environment, severe phenotype–environment mismatches result in habitat filtering: propagules reach a site but cannot establish there. Operating within individuals and populations, these processes will affect species’ habitat ranges at small scales, and aggregated across populations, will determine whether species track climatic changes and undergo geographic range shifts. This trade-off-based framework can provide a conceptual basis for species distribution models that are generalizable across plant species, aiding in the prediction of shifts in plant species’ ranges in response to climate change.
Globally, forests provide important ecosystem services, but anthropogenic change may shift the boundaries of forested biomes, because small-scale environmental changes govern biome transitions. This is especially true in semi-arid forests, where minor topographic and microclimatic changes influence forest functioning and transitions to open biomes such as grasslands. However, we lack quantitative descriptions of topographically driven microclimate variation and how it shapes forest structure, diversity, and composition in these transition zones. Leveraging a 20.2-ha forest inventory plot (Niobrara plot) at a semi-arid forest-grassland transition zone in the Niobrara River valley (Nebraska, USA), we paired data on abundances and distributions of seedlings, saplings, and adults of woody species with topographic and microclimate data to test the hypothesis that if topographic variation causes variation in microclimate that affects forest function, then forest structure, diversity, and composition should vary significantly with topography and microclimate. Microclimatic variation within the Niobrara plot strongly corresponded with topography, creating a sharp water availability and exposure gradient from the river floodplain to the forest-grassland transition zone. The magnitude of microclimate variation corresponded to that of regional macroclimate variation. Mean soil moisture was 10.2% lower along the higher-elevation transition zone than in the canyon bottoms, corresponding to variation across approximately 2.5 degrees of longitude. Mean air temperature increased by 2.2 °C from the canyon bottoms to upper canyon, corresponding to variation across approximately 3 degrees of latitude. Forest structure, diversity, and composition correlated strongly with topographic and microclimatic gradients. More complex forest structure and higher species richness of adults and saplings occurred in moister, less exposed habitats with steeper slopes and lower elevations, whereas seedling stem density and richness were higher in higher-light, moister habitats at lower elevations. Species occupied well-defined topographic niches, promoting high beta diversity along topographic and microclimatic gradients and high species turnover from the floodplain to the transition zone. Synthesis : Microclimatic and topographic variation drive patterns of structure, diversity, and composition in the forests at this forest-grassland transition zone. As the macroclimate becomes warmer and drier, topographically mediated microclimatic refuges supporting diverse, structurally complex forested ecosystems may shrink in semi-arid regions.
Climate change is associated with increases in temperature and the frequency and intensity of abiotic disturbances such as drought that can affect forest resilience at multiple scales. Higher temperatures leading to extended growing seasons alter the phenology, distribution, and population dynamics of herbivorous insects. Bark beetles (Coleoptera: Curculionidae: Scolytinae) are among the most ecologically and economically important herbivorous insect taxa and can undergo population eruptions and cause widespread coniferous tree mortality. Specifically, Ips bark beetles preferentially attack conifers that are stressed, dying, or dead—such tree conditions are predicted to become more common due to climate change. Our chapter objectives were to synthesize knowledge on the effects of drought on (1) the incidence of Ips beetle outbreaks and (2) changes in conifer host physiology that increase their susceptibility to bark beetles. We documented 132 Ips beetle outbreaks globally between 1919 and 2018 that were reported in association with drought or water stress. However, very few studies quantitatively demonstrated this relationship and/or defined the mechanisms using experiments. Alterations in host physiology that lead to tree stress will increase colonization by Ips beetles, but their responses may be dependent on attributes at the tree, stand to landscape level. Given the shifts in precipitation and temperature regimes worldwide, we argue for more in-depth studies that quantify the damage and drivers of Ips beetle outbreaks, assessing simultaneous changes in host trees and Ips beetles, ways to mitigate damage for beetle outbreaks, and urgently needed proactive adaptive management strategies in drought-prone forests worldwide.
Several species of aggressive bark beetle in the genus Dendroctonus (Coleoptera: Curculionidae) undergo large fluctuations in population density with distinct outbreak and non-outbreak phases. We investigated attributes we hypothesized as subject to density-dependent variation (in particular, those likely to express phenotypic plasticity) in the southern pine beetle (SPB), Dendroctonus frontalis, as possible indicators of population fluctuations. These traits were morphology (body size and hindwing shape) and the sex ratio of trap-captured, dispersing SPB populations. We compared attributes of beetles from locations that ranged from having zero to high numbers (>1500) of SPB infestations (spots) at the county level for two summers. Southern pine beetle were obtained from six states in the southeastern USA and had been collected during a springtime, region-wide trapping survey used for forecasting outbreaks annually. Although we detected an expected but weak sexual dimorphism in both size and shape-related traits, no morphological differences were found between SPB collected from traps in counties with low or high densities of spots (<= 10 or >10 per county, respectively). We found no relationship between numbers of SPB spots per county and trapped sex ratios in 2016, but we observed a strong trend in 2017, with about three times higher proportions of females trapped in counties with low compared with high numbers of spots. This implies that one or more known or possible factors influencing trapped sex ratios (e.g., disparities between the sexes in their responses to semiochemicals or in their propensity to disperse) can be density-dependent. Including trap-captured sex ratios in prediction models may improve current forecasting of SPB outbreaks in the southeastern USA, informing more timely and effective management of one of the most economically and ecologically important beetle species of this region.
The southeastern U.S. is considered the "wood-basket" of the world where loblolly pine (Pinus taeda L.) plantations provide tremendous ecological and economic benefits to the region. These plantations are susceptible to various natural and anthropogenic abiotic and biotic stressors and disturbances. The southern pine engravers, Ips avulsus (Eichhoff), I. calligraphus (Germar), and I. grandicollis (Eichhoff), are considered secondary colonizers of stressed, damaged, and dying loblolly pines. However, they may undergo outbreaks and colonize live pine hosts if environmental conditions cause physiological stress to trees. In 2016, > 230 concurrent Ips infestations > 2 ha in size were documented in Georgia, U.S., reportedly due to severe drought. In these forests, prescribed burning is often conducted every 2-3 years to reduce fuel-loads, improve wildlife habitat, and manage understory vegetation. However, the effects of low-severity prescribed fire on active southern Ips infestations are unknown; fire may exacerbate or alleviate beetle outbreaks. Our objectives were to: (1) compare Ips infestations between burned and unburned sites to determine the short-term effects of prescribed fire on loblolly pine mortality; and (2) determine which site-level and tree-level variables were the best predictors of short-term levels of tree mortality. We monitored 838 pines on ten sites for eight months following prescribed fire in spring 2017. Overall, 69 (8%) trees died with 3.6 times higher tree mortality on unburned sites, and a higher probability of survival on burned sites. At the site-level, binomial logistic regression models including treatment (unburned versus burned) and time since burn were the best predictors of loblolly pine mortality. At the tree-level, model selection showed that treatment, crown mortality level (1-5), Ips activity level (none, low, medium, and high), and tree diameter provided the best predictions of mortality. Prescribed burning may thus help alleviate pest pressure and increase tree resilience in loblolly pine forests in the southeastern U.S.
Flume tests and in situ jet erosion tests (JETs) allow for the process-based quantification of erodibility parameters of cohesive soils. In the excess shear stress model used to predict soil detachment, the critical shear stress (tau(c)) corresponds to the stress at which fluvial forces can detach soil particles or aggregates and the erodibility coefficient (k(d)) governs the rate at which detachment occurs when the imposed shear exceeds tau(c). The primary objective of this research was to derive and compare erodibility parameters from flume tests and JETs on various root-permeated soil samples. Based on statistical analysis, erodibility coefficients from the flume tests were typically statistically similar to those derived with JETs. In order to further determine the capability of the JET to indicate differences in erodibility of root-permeated soils, this research then performed JETs on side-by-side in situ soils. The JETs on root-permeated soils estimated significantly higher tau(c) but insignificant differences in k(d) compared to JETs on adjacent bare soil. A significant correlation was observed between exposed root surface area (TRSA) and tau(c), but no correlation between TRSA and k(d) or the equilibrium scour depth was observed. More flume and JET experiments are needed to clarify the relationship between root-permeated soils and erodibility parameters, and address a wider range of average root diameters than was considered in this research.