海南尖峰岭热带山地雨林不同空间尺度和径级水平的物种丰富度与个体密度关联

以海南尖峰岭热带山地雨林为研究对象, 基于尖峰岭原始林60 hm²大样地, 分析了7个不同空间尺度(5 m × 5 m、10 m × 10 m、20 m × 20 m、40 m × 40 m、60 m × 60 m、80 m × 80 m、100 m × 100 m)、3个不同径级(胸径(DBH) ≥ 1.0 cm、DBH ≥ 2.5 cm、DBH ≥ 7.5 cm)物种丰富度与个体密度之间的关联性, 以期为负密度制约效应等研究的尺度选择提供参考依据。研究结果表明, 物种丰富度和个体密度之间的关联性是基于空间尺度存在的, 并受分析植株径级大小的影响。DBH ≥ 1.0 cm时, 在4个空间尺度下(5 m × 5 m、10 m × 10 m、20 m × 20 m、40 m × 40 m), 物种丰富度随个体密度增加而增加, 但物种丰富度和个体密度两者间的相关性随取样空间尺度的增大而逐渐下降; 在其他3个空间尺度下(60 m × 60 m、80 m × 80 m、100 m × 100 m), 两者间没有显著相关性。当DBH ≥ 2.5 cm和DBH ≥ 7.5 cm时, 物种丰富度和个体密度间的相关性随空间尺度变化的趋势也与上述结果基本一致; 在60 m × 60 m的空间尺度上, DBH ≥ 1.0 cm时物种丰富度和个体密度相关性不显著, DBH ≥ 2.5 cm和DBH ≥ 7.5 cm两个径级的物种丰富度和个体密度表现出弱相关性。DBH ≥ 2.5 cm和DBH ≥ 7.5 cm时, 各空间尺度下物种丰富度和个体密度之间的决定系数几乎一致, 但高于DBH ≥ 1.0 cm时两者之间的决定系数, 说明大径级植株的个体密度和物种丰富度之间有更强的关联性, 负密度制约效应和自疏效应可能是产生这种现象的主要原因。     

Are bark beetle outbreaks less synchronous than forest Lepidoptera outbreaks?

Comparisons of intraspecific spatial synchrony across multiple epidemic insect species can be useful for generating hypotheses about major determinants of population patterns at larger scales. The present study compares patterns of spatial synchrony in outbreaks of six epidemic bark beetle species in North America and Europe. Spatial synchrony among populations of the Eurasian spruce bark beetle Ips typographus was significantly higher than for the other bark beetle species. The spatial synchrony observed in epidemic bark beetles was also compared with previously published patterns of synchrony in outbreaks of defoliating forest Lepidoptera, revealing a marked difference between these two major insect groups. The bark beetles exhibited a generally lower degree of spatial synchrony than the Lepidoptera, possibly because bark beetles are synchronized by different weather variables that are acting on a smaller scale than those affecting the Lepidoptera, or because inherent differences in their dynamics leads to more cyclic oscillations and more synchronous spatial dynamics in the Lepidoptera.     

Drought and stand susceptibility to attacks by the European spruce bark beetle: A remote sensing approach

1. Several time-series analyses have demonstrated that after extreme summer drought bark beetle damage increased. However, studies predicting stand susceptibility over large spatial extents are limited by technical constraints in obtaining detailed, spatially-explicit data on infestation spot occurrence.
2. Using a unique dataset of georeferenced bark beetle infestation data, we tested whether the spatial variation of local growing conditions of forest stands, topography, and landscape variables modified the local occurrence of Ips typographus infestations after a severe hot drought in Central Europe.
3. Bark beetle infestation occurrence depended on soil-related aridity intensity, elevation, slope, and soil conditions. We showed that elevation interacted with growing conditions and topography. At low elevations, spruce forests growing on flat areas and wetter soils were more sensitive to the infestations. On the contrary, forests on steep slopes and soils with low water availability were rarely attacked. At the landscape scale, bark beetle damage increased with host tree cover but decreased with compositional diversity.
4. Our findings are generally consistent with the growth-differentiation balance hypothesis that predicts that trees growing under chronic dry conditions tend to be more resistant against biotic disturbances.
5. Spruce stands at low elevations located in homogeneous landscapes dominated by spruce were those more exposed to bark beetles in the initial phase of a drought-induced outbreak.

     

黄土高原马栏林区基于不同植被组织尺度的群落物种多样性

物种多样性格局随着时空尺度的变化而变化, 同时也与植被组织尺度的变化密切相关, 基于多组织尺度的研究能更好地揭示一个地区的物种多样性规律。在应用数量分类(TWINSPAN)和主成分分析法(PCA)确定黄土高原马栏林区不同组织尺度的群落类型及其相互关系的基础上, 采用加性分配法分析该区域物种多样性与植被型、群系和群丛3种植被组织尺度之间的关系, 结果表明: (1) 区域物种多样性(γ)可加性分配分为群丛内(α₁)、群丛间(β₁)、群系间(β₂)和植被型间(β₃) 4个多样性成分, 无论用物种丰富度指数还是Shannon-Wiener多样性或Simpson多样性指数, 乔木、灌木和草本植物的最大物种多样性都存在于群丛内(草本层的物种丰富度除外), 说明群丛尺度是度量该区物种多样性的最佳尺度。(2)植被型、群系和群丛3种组织尺度的Shannon-Wiener多样性百分比均大于Simpson多样性百分比, 说明稀有种的分布对马栏林区各植被组织尺度的物种多样性格局起主要作用。(3)各尺度间的β多样性大小顺序在乔木、灌木、草本植物3层以不同多样性指数表示时各不相同, 这与乔木、灌木、草本植物3层的物种组成和分布, 以及主导不同植被组织尺度的物种多样性的因素差异有密切联系。     

Influence of tree hollow characteristics and forest structure on saproxylic beetle diversity in tree hollows in managed forests in a regional comparison

Tree hollows are among the rarest habitats in today''s Central European managed forests but are considered key structures for high biodiversity in forests. To analyze and compare the effects of tree hollow characteristics and forest structure on diversity of saproxylic beetles in tree hollows in differently structured managed forests, we examined between 41 and 50 tree hollows in beech trees in each of three state forest management districts in Germany. During the two‐year study, we collected 283 saproxylic beetle species (5880 individuals; 22% threatened species), using emergence traps. At small spatial scales, the size of hollow entrance and the number of surrounding microhabitat structures positively influenced beetle diversity, while the stage of wood mould decomposition had a negative influence, across all three forest districts. We utilized forest inventory data to analyze the effects of forest structure in radii of 50–500 m around tree hollows on saproxylic beetle diversity in the hollows. At these larger spatial scales, the three forest management districts differed remarkably regarding the parameters that influenced saproxylic beetle diversity in tree hollows. In Ebrach, characterized by mostly deciduous trees, the amount of dead wood positively influenced beetle diversity. In the mostly coniferous Fichtelberg forest district, with highly isolated tree hollows, in contrast, only the proportion of beech trees around the focal tree hollows showed a positive influence on beetle diversity. In Kelheim, characterized by mixed forest stands, there were no significant relationships between forest structure and beetle diversity in tree hollows. In this study, the same local tree hollow parameters influenced saproxylic beetle diversity in all three study regions, while parameters of forest structure at larger spatial scales differed in their importance, depending on tree‐species composition.     

Impacts of silvicultural thinning treatments on beetle trap captures and tree attacks during low bark beetle populations in ponderosa pine forests of northern Arizona

Our research used a combination of passive traps, funnel traps with lures, baited trees, and surveys of long-term thinning plots to assess the impacts of different levels of stand basal area (BA) on bark beetle tree attack and on trap captures of Ips spp., Dendroctonus spp., and their predators. The study occurred at two sites in ponderosa pine, Pinus ponderosa Dougl. ex Laws., forests, from 2004 to 2007 during low bark beetle populations. Residual stand BA ranged from 9.0 to 37.0 m2/ha. More predators and bark beetles were collected in passive traps in stands of lower BA than in stands of higher BA; however, significance varied by species and site, and total number of beetles collected was low. Height of the clear panel passive traps affected trap catches for some species at some sites and years. When pheromone lures were used with funnel traps [Ips pini (Say) lure: lanierone, +03/-97 ipsdienol], we found no significant difference in trap catches among basal area treatments for bark beetles and their predators. Similarly, when trees were baited (Dendroctonus brevicomis LeConte lure: myrcene, exo-brevicomin and frontalin), we found no significant difference for days to first bark beetle attack. Surveys of long-term thinning treatments found evidence of bark beetle attacks only in unthinned plots (approximately 37 m2/ha basal area). We discuss our results in terms of management implications for bark beetle trapping and control.     

Population dynamics in changing environments: the case of an eruptive forest pest species

In recent decades we have seen rapid and co‐occurring changes in landscape structure, species distributions and even climate as consequences of human activity. Such changes affect the dynamics of the interaction between major forest pest species, such as bark beetles (Coleoptera: Curculionidae, Scolytinae), and their host trees. Normally breeding mostly in broken or severely stressed spruce; at high population densities some bark beetle species can colonise and kill healthy trees on scales ranging from single trees in a stand to multi‐annual landscape‐wide outbreaks. In Eurasia, the largest outbreaks are caused by the spruce bark beetle, Ips typographus (Linnaeus), which is common and shares a wide distribution with its main host, Norway spruce (Picea abies Karst.). A large literature is now available, from which this review aims to synthesize research relevant for the population dynamics of I. typographus and co‐occurring species under changing conditions. We find that spruce bark beetle population dynamics tend to be metastable, but that mixed‐species and age‐heterogeneous forests with good site‐matching tend to be less susceptible to large‐scale outbreaks. While large accumulations of logs should be removed and/or debarked before the next swarming period, intensive removal of all coarse dead wood may be counterproductive, as it reduces the diversity of predators that in some areas may play a role in keeping I. typographus populations below the outbreak threshold, and sanitary logging frequently causes edge effects and root damage, reducing the resistance of remaining trees. It is very hard to predict the outcome of interspecific interactions due to invading beetle species or I. typographus establishing outside its current range, as they can be of varying sign and strength and may fluctuate depending on environmental factors and population phase. Most research indicates that beetle outbreaks will increase in frequency and magnitude as temperature, wind speed and precipitation variability increases, and that mitigating forestry practices should be adopted as soon as possible considering the time lags involved.     

Capturing cross-scalar variation of habitat selection with grid sampling: an example with hazel grouse (Tetrastes bonasia L.)

The objective of this study was to investigate a grid-based sampling design to determine the cross-scalar selection of habitat by a territorial animal species: the hazel grouse (Tetrastes bonasia L.). In each of three sites with increasing hazel grouse nest site density, three lattice grids were used to measure both the habitat variables and the species occurrence in 100 30?×?30 m cells. We calculated the average values for habitat variables, as well as use versus non-use by the species, at three spatial scales: small (1?×?1 cell), intermediate (2?×?2 cells) and large (3?×?3 cells). Generalised linear mixed models were integrated into a method of variation and hierarchical partitioning and used to assess the relationship between the habitat variables and the species preferences at each scale. In all scales, species selection was associated with ground layer composition. Selection was also associated with the composition of the woody layer and negatively associated with dominance of tor grass (Brachypodium rupestre (Host) Roem. & Schult.) at the two larger scales. Both litter cover and thinning contributed positively to the habitat selection at the two smaller scales. The other variables were significant only at one scale or explained a relatively low proportion of the variation at multiple scales. Neither the management nor the stand structure variables played a significant independent role across scales when compared with ground layer variables. The total variation explained was highest (ca. 90 %) at the large scale. This finding indicates the possibility of obtaining cross-scalar hazel grouse preferences from grid-based sampling, provided that spatial autocorrelation in the data is handled appropriately.     

Space-for-time substitution reveals a hump-shaped distribution of dung beetles

Unraveling how climate change impacts the diversity and distribution patterns of organisms is a major concern in ecology, especially with climate-sensitive species, such as dung beetles. Often found in warmer weather conditions, beetles are used as bio-indicators of environmental conditions. By using an altitudinal gradient as a proxy for climate change (i.e., space-for-time substitution), we assessed how changes in climatic variables, such as temperature and precipitation, impact patterns of dung beetle diversity and distribution in the Peruvian Andes. We recorded dung beetle diversity using three different types of baits, feces, carrion, and fruits, distributed in 18 pitfall traps in five different altitudinal sites (from 900 to 2500 m, 400 m apart from each other) in the rainy and dry season. We found that (i) dung beetle richness and abundance were influenced by the climate gradient, (ii) seasonality influenced beetle richness, which was high in the wet season, but did not influence abundance, (iii) dung beetle richness and abundance fit to a hump-shaped distribution pattern along the altitudinal gradient, and (iv) species richness is the beta-diversity component that best describes the composition of dung beetle species along the altitudinal gradient. Our data show that the distribution and diversity of dung beetles are different at larger scales, with different patterns resulting from the response of species to both abiotic and biotic factors.     

Comparative study of spatial patterns and ecological niches of beetles in two Malaysian mountains elevation gradients

The study of beetle communities is a valuable approach for biogeographical and conservation studies because their species and ecological diversities are very high, and they take different roles in ecosystems. However, beetle macroecology and conservation studies are disproportionately scarce, especially in tropical Asia. The objective of this study is to compare beetle abundance, diversity and species richness along the elevation gradients in two mountains in Peninsular Malaysia. Three passive sampling methods were utilized for beetle sampling with four marked elevation gradients: 500 m, 1000 m, 1500 m and 1800 m. Species richness in Fraser’s Hill was higher at highest elevation, but this value was not-significantly different from these in other elevations, except for the site at 1000 m with significantly lower estimates. Genting Highlands showed a significant decrease in species richness with the increase in elevation, without differences between the higher elevation sites. Pairwise comparison of species richness, Simpson Dominance and Shannon diversity between same elevation sites of Fraser’s Hill and Genting Highlands were all significantly different. The levels of vertical and horizontal colonization have had comparatively different weights in terms of their effect on the pattern of diversity and the integration of the beetle community in these two localities. At Fraser’s Hill, similar conditions at different elevations drives different responses, whereas at Genting Highlands contrasting and different environmental conditions at each elevation, drives different responses. We suggest the potential use of these results for biodiversity conservation in terms of climate variables in accordance with niche patterns.     

Climate change and the outbreak ranges of two North American bark beetles

Abstract

• 1 One expected effect of global climate change on insect populations is a shift in geographical distributions toward higher latitudes and higher elevations. Southern pine beetle Dendroctonus frontalis and mountain pine beetle Dendroctonus ponderosae undergo regional outbreaks that result in large‐scale disturbances to pine forests in the south‐eastern and western United States, respectively.
• 2 Our objective was to investigate potential range shifts under climate change of outbreak areas for both bark beetle species and the areas of occurrence of the forest types susceptible to them.
• 3 To project range changes, we used discriminant function models that incorporated climatic variables. Models to project bark beetle ranges employed changed forest distributions as well as changes in climatic variables.
• 4 Projected outbreak areas for southern pine beetle increased with higher temperatures and generally shifted northward, as did the distributions of the southern pine forests.
• 5 Projected outbreak areas for mountain pine beetle decreased with increasing temperature and shifted toward higher elevation. That trend was mirrored in the projected distributions of pine forests in the region of the western U.S. encompassed by the study.
• 6 Projected outbreak areas for the two bark beetle species and the area of occurrence of western pine forests increased with more precipitation and decreased with less precipitation, whereas the area of occurrence of southern pine forests decreased slightly with increasing precipitation.
• 7 Predicted shifts of outbreak ranges for both bark beetle species followed general expectations for the effects of global climate change and reflected the underlying long‐term distributional shifts of their host forests.

     

Attack density and breeding success of bark beetles (Coleoptera, Scolytidae) at different distances from forest-clearcut edge

1 Attack density and breeding success of bark beetles (Coleoptera: Scolytidae) and the abundance of their predators were studied at forest-clearcut edges in southern Finland on Norway spruce, Picea abies, bolts placed at different distances from a forest-clearcut edge. 2 Attack densities of two species, Hylurgops palliatus and H. glabratus, increased markedly towards the forest interior. The breeding success of H. palliatus was increased with increasing distance from the stand edge. None of the recorded 10 bark beetle species was distinctly abundant near the edge, but Pityogenes chalcographus appeared to prefer this zone. 3 There were no significant differences in numbers of bark beetle predators between the forest interior and the edge. Forest-clearcut edge had a significant effect on the desiccation of sample bolts. 4 Thus, forest edge is likely to affect the colonization behaviour and breeding success of bark beetles via host material properties.     

The tempo and mode of gopher mound production in a tallgrass prairie remnant

To gain better insight into how small-scale disturbances might affect ecological processes, such as the maintenance of plant species diversity, we conducted a two-year study characterizing spatio-temporal patterns of gopher mound production on a tallgrass prairie remnant located at Anderson Prairie, Iowa. USA. The locations of all newly produced gopher mounds were mapped on two 80 × 80 m permanent plots. We used these data to characterize spatio-temporal patterns of mound production across a range of scales. We found that mound production was highly clustered at scales of < 8 m over short periods of time (< 2 weeks), but shifted in location over a 3–4 weeks time period, resulting in a clustered pattern at scales of < 20 m over longer time periods (up to the 2 yr of the study). We also found that patterns of mound production at intermediate spatial scales (> 20 m) remained fairly static over time, although they differed significantly from site to site. The results of this study suggest that small-scale patterns of variability in mound production may increase habitat variability over very short spatial scales, possibly providing a mechanism that can enhance the development and maintenance of species diversity.     

Functional diversity versus species diversity: relationships with habitat heterogeneity at multiple scales in a subtropical evergreen broad-leaved forest

Understanding the relationship between functional and species diversity as well as their association with habitat heterogeneity can help reveal the mechanisms of species coexistence in ecological communities. However, these interactions have been poorly studied in subtropical forests. In this paper, we evaluated functional diversity (as measured by Rao’s Q) and traditional species diversity (based on Simpson’s index) in a 24 ha forest plot in a subtropical evergreen broad-leaved forest (EBLF) in China. We compared the sensitivities of functional and species diversity to topographic variables (elevation, convexity, slope and aspect) at multiple spatial scales based on 10 × 10, 20 × 20, 40 × 40 and 50 × 50 m quadrats. Functional and species diversity were found to have different distribution patterns along a topographical gradient, with functional diversity better explained by topography than was species diversity using a spatial autocorrelation regression error model. Furthermore, functional diversity had a significantly greater association with topographic variables than species diversity in both adult and young trees; in both cases, the strength of the diversity-habitat association increased with quadrat size. We conclude that functional diversity reflects a greater diversity-habitat association in EBLF than does species diversity, and that the association depends on the spatial scale and life stages of the woody plants under evaluation.     

Habitat loss decreases predator–prey ratios in a pine-bark beetle system

Increasing intensity of land use by humans has led to loss of natural habitats, resulting in isolation of remaining habitat fragments. Using a pine-bark beetle ecosystem as a model, we tested the hypothesis that the ratio of abundance of predators to prey should decrease with increasing habitat loss at the landscape scale. We selected ten red pine ( Pinus resinosa ) sites, representing extremes of available habitat within a 2 km radius surrounding each stand. The bark beetle, Ips pini , and its coleopteran predators were sampled using baited multiple funnel traps. Effects of stand isolation were considerable; ratios of predators to prey (mean number of predators/number of prey±SE) were significantly reduced in isolated stands (0.38±0.09) as compared to those with large amounts of surrounding conifer habitat (1.63±0.41). The decline in ratio occurred both because there was: a) a lower abundance of predators (ca 0.5–0.8×) captured in isolated stands; and b) a significantly higher number of prey (ca 2.2×) captured in isolated stands. Isolation or loss of habitat, therefore, differentially affected the two trophic levels, supporting theoretical predictions. Reductions in predator abundance and, presumably, enemy-caused mortality may lead to changes in the population dynamics of their prey species, possibly leading to increased outbreaks as habitat becomes increasingly isolated.     

Spatial genetic structure in seed stands of <Emphasis Type="Italic">Pinus lumholtzii</Emphasis> B.L. Rob. &amp; Fernald in Durango,Mexico

Sad pine is one of the most prominent pine species in Mexico due to its conspicuous pendulous foliage and extreme habitat. However, scientific studies of the species are scarce, and genetic information on sad pine populations is lacking. This endemic tree species occurs naturally on the Sierra Madre Occidental where it covers a total area of about 1,600,000 ha. It typically grows with several species of Quercus and Pinus or in pure stands in uneven-aged forests. Pinus lumholtzii is naturally spatially fragmented, and genetic research on seed and pollen dispersal patterns and spatial genetic structure (SGS)—and the possible implications of these in terms of evolution, conservation and breeding management—is particularly important. Given the fragmented occurrence of the preferred soil type, the goal of this research was to use amplified fragment length polymorphism (AFLP) markers to identify potential differences in spatial genetic structure within and between five P. lumholtzii seed stands at fine and large scales. At the fine scale, we almost always observed non-significant autocorrelation, suggesting that the genetic variants of P. lumholtzii are randomly distributed in space within each sampled seed stand. At the larger scale, our findings provide strong support for the theory of isolation by distance that predicts the expected pattern of SGS at drift–dispersal equilibrium. We recommend a network of P. lumholtzii seed stands of maximum distances of 100 km among stands to prevent greater loss of local genetic variants and use the seeds for reforestations in a radius of maximal 50 km from their proveniences.     

Varying Patterns on Varying Scales: A Metacommunity Analysis of Nematodes in European Lakes

Ecological community patterns are often extremely complex and the factors with the greatest influence on community structure have yet to be identified. In this study we used the elements of metacommunity structure (EMS) framework to characterize the metacommunities of freshwater nematodes in 16 European lakes at four geographical scales (radius ranging from 80 m to 360 km). The site characteristics associated with site scores indicative of the structuring gradient were identified using Spearman rank correlations. The metacommunities of the 174 nematode species included in this analysis mostly had a coherent pattern. The degree of turnover increased with increasing scale. Ordination scores correlated with geographical variables on the larger scales and with the trophic state index on a regional scale. The association of the structuring gradient with spatial variables and the scale-dependent increase in turnover showed that nematode dispersal was limited. The different metacommunity patterns identified at the increasing geographical scales suggested different, scale-related mechanisms of species distribution, with species sorting dominating on smaller and mass effects on larger geographical scales.     

Does functional homogenization accompany taxonomic homogenization of British birds and how do biotic factors and climate affect these processes?

Environmental change has reshuffled communities often causing taxonomic homogenization rather than differentiation. Some studies suggest that this increasing similarity of species composition between communities is accompanied by an increase in similarity of trait composition—functional homogenization—although different methodologies have failed to come to any consistent conclusions. Functional homogenization could have a large effect on ecosystem functioning and stability. Here, we use the general definition of homogenization as “reduced spatial turnover over time” to compare changes in Simpson's beta diversity (taxonomic turnover) with changes in Rao's quadratic entropy beta diversity (functional turnover) in British breeding birds at three spatial scales. Using biotic and climatic variables, we identify which factors may predispose a site to homogenization. The change in turnover measures between two time periods, 20 years apart, was calculated. A null model approach was taken to identify occurrences of functional homogenization and differentiation independent of changes in taxonomic turnover. We used conditional autoregressive models fitted using integrated nested Laplace approximations to determine how environmental drivers and factors relating to species distributions affect changes in spatial turnover of species and functional diversity. The measurement of functional homogenization affects the chance of rejection of the null models, with many sites showing taxonomic homogenization unaccompanied by functional homogenization, although occurrence varies with spatial scale. At the smallest scale, while temperature‐related variables drive changes in taxonomic turnover, changes in functional turnover are associated with variation in growing degree days; however, changes in functional turnover become more difficult to predict at larger spatial scales. Our results highlight the multifactorial processes underlying taxonomic and functional homogenization and that redundancy in species traits may allow ecosystem functioning to be maintained in some areas despite changes in species composition.     

Beetle diversity in a matrix of old-growth boreal forest: influence of habitat heterogeneity at multiple scales

The relative contribution of compositional and structural heterogeneity on biodiversity is currently ambiguous because field studies generally integrate these two sources of habitat heterogeneity into a single index. We established the relationship between species richness of ground-dwelling and flying beetles and compositional and structural attributes of forest heterogeneity. The relationship was evaluated at two spatial scales: the scale of forest stand, corresponding to an 11.3  m radius, and the scale of landscape, corresponding to either a 400 or 800  m radius. Seventy stands were sampled in the matrix of old-growth boreal forest of the North Shore region of Québec, Canada, during the summers of 2004 and 2005. A total of 133 ground-dwelling beetle species (range: 4–42 species per site) were captured in the pitfall traps and 251 flying species (range 16–58 species per site) in flight-interception traps. We found that the most relevant type of heterogeneity to explain variations in species richness and the significance of landscape scale information varied between groups of beetles. Compositional heterogeneity (i.e. the number of species of forest trees and shrubs) at the stand scale best predicted species richness in ground-dwelling beetles. On the other hand, it was the combined influence of structural and compositional habitat heterogeneity at stand and landscape scales that best explained richness patterns in flying beetles. Our study outlines the significance of considering multiple types and spatial scales of habitat heterogeneity when describing patterns of species richness.     

Anomalous outbreaks of an invasive defoliator and native bark beetle facilitated by warm temperatures,changes in precipitation and interspecific interactions

Biotic disturbance agents such as insects can be highly responsive to climatic change and have widespread ecological and economic impacts on forests. Quantifying the responses of introduced and native insects to climate, including how dynamics of one agent may mediate those of another, is important for forecasting disturbance and associated impacts on forest structure and function. We investigated drivers of outbreaks by larch casebearer Coleophora laricella, an invasive defoliator, and eastern larch beetle Dendroctonus simplex, a native, tree‐killing bark beetle, on tamarack Larix laricina from 2000 to in Minnesota, USA. We evaluated the utility of temporal, spatial and climatic variables in predicting the presence/absence of outbreaks of each insect in cells of rasterized aerial survey data. The role of defoliation by larch casebearer in outbreaks of eastern larch beetle was also investigated. For both species, the most important predictors of outbreak occurrence were proximity of conspecific outbreaks in space and time. For larch casebearer, outbreak occurrence was positively associated with spring precipitation and warmer growing seasons. Outbreak occurrence of eastern larch beetle was positively associated with warmer and dryer years and was more likely in cells with prior defoliation by larch casebearer. Our results demonstrate that climate can drive large scale outbreaks of introduced and non‐native disturbance agents on a single host species, and that interactions at the tree level between such agents may scale up to manifest across large temporal and spatial scales.