Spatial patterns of hyperparasitism along a latitudinal gradient of forest genus diversity: insights from the spruce budworm-parasitoids community.

High-order mobile predators are generally thought to increase ecosystem stability and resilience to natural perturbations. In many insect food-webs, higher trophic positions are occupied by parasitoids, which are themselves hosts for hyperparasitoids that can reduce primary parasitoids' efficiency in controlling insect pests. Hyperparasitoids can thus provide ecosystem disservices by facilitating pest outbreaks, or ecosystem services by stabilizing food web fluctuations over longer time periods. To better understand how hyperparasitism affects multitrophic forest systems, we examined for the first time spatial variations in hyperparasitism associated with the spruce budworm. We examined 2 common primary parasitoids of the spruce budworm during outbreaks (Apanteles fumiferanae and Glypta fumiferanae), and estimated their true and pseudohyperparasitism rates in 2014-2015 from 28 locations across a latitudinal gradient (over 450 km) of forest genus diversity. Hyperparasitoid cryptic diversity was also quantified using DNA-barcoding. We found that A. fumiferanae and G. fumiferanae share at least 2 of 5 common hyperparasitoid species, confirming the connected nature of the spruce budworm-parasitoid food web. Moreover, hyperparasitism is modulated by spatial context as we observed a positive correlation between forest genus diversity and hyperparasitism for A. fumiferanae, but not for G. fumiferanae. Further monitoring hyperparasitism holds significant potential to provide new insights into how forest composition affects multitrophic interactions and spatio-temporal outbreak dynamics.

Quantifying and predicting population connectivity of an outbreaking forest insect pest.

Context: Dispersal has a key role in the population dynamics of outbreaking species such as the spruce budworm (Choristoneura fumiferana) as it can synchronize the demography of distant populations and favor the transition from endemic to epidemic states. However, we know very little about how landscape structure influences dispersal in such systems while such knowledge is essential for better forecasting of spatially synchronous population dynamics and to guide management strategies. Objectives: We aimed to characterize the spatial environmental determinants of spruce budworm dispersal to determine how these features affect outbreak spread in Quebec (Canada). We then apply our findings to predict expected future landscape connectivity and explore its potential consequences on future outbreaks. Methods: We used a machine-learning landscape genetics approach on 447 larvae covering most of the outbreak area and genotyped at 3562 SNP loci to identify the main variables affecting connectivity. Results: We found that the connectivity between outbreak populations was driven by the combination of precipitation and host cover. Our forecasting suggests that between the current and next outbreaks, connectivity may increase between Ontario and Quebec, and might decrease in the eastern part, which could have the effect of limiting outbreak spread from Ontario and Quebec to the eastern provinces. Conclusions: Although we did not identify any discrete barriers, low connectivity areas might constrain dispersal in the current and future outbreaks and should in turn, be intensively monitored. However, continued sampling as the outbreak progresses is needed to confirm the temporal stability of the observed patterns. Supplementary Information: The online version contains supplementary material available at 10.1007/s10980-021-01382-9.

Landscape-scale population connectivity in two parasitoid species associated with the spruce budworm: Testing the birdfeeder effect using genetic data.

Periodic and spatially synchronous outbreaks of insect pests have dramatic consequences for boreal and sub-boreal forests. Within these multitrophic systems, parasitoids can be stabilizing agents by dispersing toward patches containing higher host density (the so-called birdfeeder effect). However, we know little about the dispersal abilities of parasitoids in continuous forested landscapes, limiting our understanding of the spatiotemporal dynamics of host-parasitoid systems, and constraining our ability to predict forest resilience in the context of global changes. In this study, we investigate the spatial genetic structure and spatial variation in genetic diversity of two important species of spruce budworm larval parasitoids during outbreaks: Apanteles fumiferanae Viereck (Braconidae) and Glypta fumiferanae (Viereck) (Ichneumonidae). Using parasitoids sampled in 2014 from 26 and 29 locations across a study area of 350,000 km2 , we identified 1,012 and 992 neutral SNP loci for A. fumiferanae (N = 279 individuals) and G. fumiferanae (N = 382), respectively. Using DAPC, PCA, AMOVA, and IBD analyses, we found evidence for panmixia and high genetic connectivity for both species, matching the previously described genetic structure of the spruce budworm within the same context, suggesting similar effective dispersal during outbreaks and high parasitoid population densities between outbreaks. We also found a significant negative relationship between genetic diversity and latitude for A. fumiferanae but not for G. fumiferanae, suggesting that northern range limits may vary by species within the spruce budworm parasitoid community. These spatial dynamics should be considered when predicting future insect outbreak severities in boreal landscapes.

The Vision of Managing for Pest-Resistant Landscapes: Realistic or Utopic?

Purpose of Review: Forest managers have long suggested that forests can be made more resilient to insect pests by reducing the abundance of hosts, yet this has rarely been done. The goal of our paper is to review whether recent scientific evidence supports forest manipulation to decrease vulnerability. To achieve this goal, we first ask if outbreaks of forest insect pests have been more severe in recent decades. Next, we assess the relative importance of climate change and forest management-induced changes in forest composition/structure in driving these changes in severity. Recent Findings: Forest structure and composition continue to be implicated in pest outbreak severity. Mechanisms, however, remain elusive. Recent research elucidates how forest compositional and structural diversity at neighbourhood, stand, and landscape scales can increase forest resistance to outbreaks. Many recent outbreaks of herbivorous forest insects have been unprecedented in terms of duration and spatial extent. Climate change may be a contributing factor, but forest structure and composition have been clearly identified as contributing to these unprecedented outbreaks. Summary: Current research supports using silviculture to create pest-resistant forest landscapes. However, the precise mechanisms by which silviculture can increase resistance remains uncertain. Further, humans tend to more often create pest-prone forests due to political, economic, and human resistance to change and a short-sighted risk management perspective that focuses on reactive rather than proactive responses to insect outbreak threats. Future research efforts need to focus on social, political, cultural, and educational mechanisms to motivate implementation of proven ecological solutions if pest-resistant forests are to be favoured by management.

Community-level modelling of boreal forest mammal distribution in an oil sands landscape.

Anthropogenic landscape disturbances are known to alter, destroy, and fragment habitat, which typically leads to biodiversity loss. The effects of landscape disturbance generally vary among species and depend on the nature of the disturbances, which may interact and result in synergistic effects. Western Canada's oil sands region experiences disturbances from forestry and energy sector activities as well as municipal and transportation infrastructure. The effects of those disturbances on single species have been studied and have been implicated in declines of the boreal woodland caribou (Rangifer tarandus caribou). Yet, the specific responses of the mammal community, and of functional groups such as prey and predators, to those interacting disturbances are still poorly known. We investigated the responses of black bear, grey wolf, coyote, fisher, lynx, red fox, American red squirrel, white-tailed deer, moose, caribou, and snowshoe hare to both natural habitat and disturbance associated with anthropogenic features within Alberta's northeast boreal forest. We used a novel community-level modelling framework on three years of camera-trap data collected in an oil sands landscape. This framework allowed us to identify the natural and anthropogenic features which explained the most variation in occurrence frequency among functional groups, as well as compare responses to linear and non-linear anthropogenic disturbance. Occurrence frequency by predators was better explained by anthropogenic features than by natural habitat. Both linear and non-linear anthropogenic features helped explain occurrence frequency by prey and predators, although the effects differed in magnitude and spatial scale. To better conserve boreal biodiversity, management actions should extend beyond a focus on caribou and wolves and aim to restore habitat across a diversity of anthropogenic disturbances and monitor the dynamics of the entire mammal community.

Strategies to optimize modeling habitat suitability of Bertholletia excelsa in the Pan-Amazonia.

AIM: Amazon-nut (Bertholletia excelsa) is a hyperdominant and protected tree species, playing a keystone role in nutrient cycling and ecosystem service provision in Amazonia. Our main goal was to develop a robust habitat suitability model of Amazon-nut and to identify the most important predictor variables to support conservation and tree planting decisions. LOCALIZATION: Amazon region, South America. METHODS: We collected 3,325 unique Amazon-nut records and assembled >100 spatial predictor variables organized across climatic, edaphic, and geophysical categories. We compared suitability models using variables (a) selected through statistical techniques; (b) recommended by experts; and (c) integrating both approaches (a and b). We applied different spatial filtering scenarios to reduce overfitting. We additionally fine-tuned MAXENT settings to our data. The best model was selected through quantitative and qualitative assessments. RESULTS: Principal component analysis based on expert recommendations was the most appropriate method for predictor selection. Elevation, coarse soil fragments, clay, slope, and annual potential evapotranspiration were the most important predictors. Their relative contribution to the best model amounted to 75%. Filtering of the presences within a radius of 10 km displayed lowest overfitting, a satisfactory omission rate and the most symmetric distribution curve. Our findings suggest that under current environmental conditions, suitable habitat for Amazon-nut is found across 2.3 million km2, that is, 32% of the Amazon Biome. MAIN CONCLUSION: The combination of statistical techniques with expert knowledge improved the quality of our suitability model. Topographic and soil variables were the most important predictors. The combination of predictor variable selection, fine-tuning of model parameters and spatial filtering was critical for the construction of a reliable habitat suitability model.

Temporal variation in spatial genetic structure during population outbreaks: Distinguishing among different potential drivers of spatial synchrony.

Spatial synchrony is a common characteristic of spatio-temporal population dynamics across many taxa. While it is known that both dispersal and spatially autocorrelated environmental variation (i.e., the Moran effect) can synchronize populations, the relative contributions of each, and how they interact, are generally unknown. Distinguishing these mechanisms and their effects on synchrony can help us to better understand spatial population dynamics, design conservation and management strategies, and predict climate change impacts. Population genetic data can be used to tease apart these two processes as the spatio-temporal genetic patterns they create are expected to be different. A challenge, however, is that genetic data are often collected at a single point in time, which may introduce context-specific bias. Spatio-temporal sampling strategies can be used to reduce bias and to improve our characterization of the drivers of spatial synchrony. Using spatio-temporal analyses of genotypic data, our objective was to identify the relative support for these two mechanisms to the spatial synchrony in population dynamics of the irruptive forest insect pest, the spruce budworm (Choristoneura fumiferana), in Quebec (Canada). AMOVA, cluster analysis, isolation by distance, and sPCA were used to characterize spatio-temporal genomic variation using 1,370 SBW larvae sampled over four years (2012-2015) and genotyped at 3,562 SNP loci. We found evidence of overall weak spatial genetic structure that decreased from 2012 to 2015 and a genetic diversity homogenization among the sites. We also found genetic evidence of a long-distance dispersal event over >140 km. These results indicate that dispersal is the key mechanism involved in driving population synchrony of the outbreak. Early intervention management strategies that aim to control source populations have the potential to be effective through limiting dispersal. However, the timing of such interventions relative to outbreak progression is likely to influence their probability of success.

The role of dispersal, selection, and timing of sampling on the false discovery rate of loci under selection during geographic range expansion.

Identifying adaptive loci is important to understand the evolutionary potential of species undergoing range expansion. However, in expanding populations, spatial demographic processes such as allele surfing can create spatial patterns of neutral genetic variation that appear similar to those generated through adaptive processes. As a result, the false discovery rate of adaptive loci may be inflated in landscape genomic analyses. Here, we take a simulation modelling approach to investigate how range expansion affects our ability to correctly distinguish between neutral and adaptive genetic variation, using the mountain pine beetle outbreak system as a motivating example. We simulated the demographic and population genetic dynamics of populations undergoing range expansion using an individual-based genetic model CDMetaPOP. We investigated how the false discovery rate of adaptive loci is affected by (i) dispersal capacity, (ii) timing of sampling, and (iii) the strength of selection on an adaptive reference locus. We found that a combination of weak dispersal, weak selection, and early sampling presents the greatest risk of misidentifying loci under selection. Expanding populations present unique challenges to the reliable identification of adaptive loci. We demonstrate that there is a need for further methodological development to account for directional demographic processes in landscape genomics.

Parasitism Rates of Spruce Budworm Larvae: Testing the Enemy Hypothesis Along a Gradient of Forest Diversity Measured at Different Spatial Scales.

Vegetational diversity is generally thought to be associated with ecosystem stability and resilience to perturbations such as insect outbreaks. The enemies' hypothesis states that vegetational diversity contributes to greater top-down control of insect pests, by providing further resources to natural enemies than homogeneous environments. However, direct evaluation of this hypothesis is difficult because different species of natural enemies can respond to vegetational diversity in dissimilar manners and at different spatial scales depending on functional traits such as prey/host specificity and dispersal. In this study, we specifically test the enemies' hypothesis at the landscape level in a continuous forest environment. We investigated how parasitism of spruce budworm larvae by the common parasitoids Apanteles fumiferanae and Glypta fumiferanae vary with forest diversity and host larval density at different spatial scales in the province of Quebec (Canada). We found that parasitism rates of the two parasitoid species we examined respond in opposite ways to forest diversity. Parasitism by A. fumiferanae was positively related to forest diversity, whereas parasitism by G. fumiferanae was negatively related to forest diversity. In agreement with the enemies' hypothesis, we also found that spruce budworm larval density decreased with forest diversity. We discuss these results with respect to the enemies' hypothesis and the presumed host range of the parasitoids species we examined, as well as their body size. Because A. fumiferanae kills its host earlier than G. fumiferanae, we conclude that northern forest landscapes could be more affected by spruce budworm defoliation than southern forests during the present and future outbreaks.

Lagged cumulative spruce budworm defoliation affects the risk of fire ignition in Ontario, Canada.

Detailed understanding of forest disturbance interactions is needed for effective forecasting, modelling, and management. Insect outbreaks are a significant forest disturbance that alters forest structure as well as the distribution and connectivity of combustible fuels at broad spatial scales. The effect of insect outbreaks on fire activity is an important but contentious issue with significant policy consequences. The eastern spruce budworm (Choristoneura fumiferana) is a native defoliating insect in eastern North America whose periodic outbreaks create large patches of dead fir and spruce trees. Of particular concern to fire and forest managers is whether these patches represent an increased fire risk, if so, for how long, and how the relationship between defoliation and fire risk varies through space and time. Previous work suggests a temporary increase in flammability in budworm-killed forests, but regional and seasonal variability in these relationships has not been examined. Using an extensive database on historical lightning-caused fire ignitions and spruce budworm defoliation between 1963 and 2000, we assess the relative importance of cumulative defoliation and fire weather on the probability of ignition in Ontario, Canada. We modeled fire ignition using a generalized additive logistic regression model that accounts for temporal autocorrelation in fire weather. We compared two ecoregions in eastern Ontario (Abitibi Plains) and western Ontario (Lake of the Woods) that differ in terms of climate, geomorphology, and forest composition. We found that defoliation has the potential to both increase and decrease the probability of ignition depending on the time scale, ecoregion, and season examined. Most importantly, we found that lagged spruce budworm defoliation (8-10 yr) increases the risk of fire ignition whereas recent defoliation (1 yr) can decrease this risk. We also found that historical defoliation has a greater influence on ignition risk during the spring than during the summer fire season. Given predicted increases in forest insect activity due to global change, these results represent important information for fire management agencies that can be used to refine existing models of fire risk.

Modeling Landscape-Level Spatial Variation in Sex Ratio Skew in the Mountain Pine Beetle (Coleoptera: Curculionidae).

Through their influence on effective population sizes, sex ratio skew affects population dynamics. We examined spatial variation in female-biased sex ratios in the mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak in western Canada to better understand how environmental context affects sex ratio skew. Our specific objectives were to: 1) characterize spatial variation in mountain pine beetle sex ratio; 2) test previously asserted hypotheses that beetle sex ratio varies with tree diameter and year in outbreak; and 3) develop predictive models of sex ratio skew for larval and adult populations. Using logistic regression, we modeled the probability that an individual beetle (n = 2,369) was female as a function of multiple environmental variables across 34 stands in British Columbia and Alberta, Canada. We identified a consistent female-biased sex ratio with significantly greater skew in adults (2:1, n = 713) than in larvae (1.76:1, n = 1,643). We found that the proportion of larval females increased with decreasing tree size and with outbreak age. However, adults did not respond to tree size and larvae did not respond to outbreak age. Predictive models differed between larvae and adults. All identified models perform well and included predictors related to weather, tree diameter, and year in outbreak. Female-biased sex ratios appear to originate from differential male mortality during development rather than from sex-biased oviposition, suggesting sex ratio skew is not the cause of outbreaks, but rather a consequence.

Life-stage differences in spatial genetic structure in an irruptive forest insect: implications for dispersal and spatial synchrony.

Dispersal determines the flux of individuals, energy and information and is therefore a key determinant of ecological and evolutionary dynamics. Yet, it remains difficult to quantify its importance relative to other factors. This is particularly true in cyclic populations in which demography, drift and dispersal contribute to spatio-temporal variability in genetic structure. Improved understanding of how dispersal influences spatial genetic structure is needed to disentangle the multiple processes that give rise to spatial synchrony in irruptive species. In this study, we examined spatial genetic structure in an economically important irruptive forest insect, the spruce budworm (Choristoneura fumiferana) to better characterize how dispersal, demography and ecological context interact to influence spatial synchrony in a localized outbreak. We characterized spatial variation in microsatellite allele frequencies using 231 individuals and seven geographic locations. We show that (i) gene flow among populations is likely very high (Fst  ≈ 0); (ii) despite an overall low level of genetic structure, important differences exist between adult (moth) and juvenile (larvae) life stages; and (iii) the localized outbreak is the likely source of moths captured elsewhere in our study area. This study demonstrates the potential of using molecular methods to distinguish residents from migrants and for understanding how dispersal contributes to spatial synchronization. In irruptive populations, the strength of genetic structure depends on the timing of data collection (e.g. trough vs. peak), location and dispersal. Taking into account this ecological context allows us to make more general characterizations of how dispersal can affect spatial synchrony in irruptive populations.

Exploring the impact of climate variability during the Last Glacial Maximum on the pattern of human occupation of Iberia.

The Last Glacial Maximum (LGM) was a global climate event, which had significant repercussions for the spatial distribution and demographic history of prehistoric populations. In Eurasia, the LGM coincides with a potential bottleneck for modern humans and may mark the divergence date for Asian and European populations (Keinan et al., 2007). In this research, the impact of climate variability on human populations in the Iberian Peninsula during the Last Glacial Maximum (LGM) is examined with the aid of downscaled high-resolution (16 × 16 km) numerical climate experiments. Human sensitivity to short time-scale (inter-annual) climate variability during this key time period, which follows the initial modern human colonisation of Eurasia and the extinction of the Neanderthals, is tested using the spatial distribution of archaeological sites. Results indicate that anatomically modern human populations responded to small-scale spatial patterning in climate variability, specifically inter-annual variability in precipitation levels as measured by the standard precipitation index. Climate variability at less than millennial scale, therefore, is shown to be an important component of ecological risk, one that played a role in regulating the spatial behaviour of prehistoric human populations and consequently affected their social networks.

Spatial genetic structure of the mountain pine beetle (Dendroctonus ponderosae) outbreak in western Canada: historical patterns and contemporary dispersal.

Environmental change has a wide range of ecological consequences, including species extinction and range expansion. Many studies have shown that insect species respond rapidly to climatic change. A mountain pine beetle epidemic of record size in North America has led to unprecedented mortality of lodgepole pine, and a significant range expansion to the northeast of its historic range. Our goal was to determine the spatial genetic variation found among outbreak population from which genetic structure, and dispersal patterns may be inferred. Beetles from 49 sampling locations throughout the outbreak area in western Canada were analysed at 13 microsatellite loci. We found significant north-south population structure as evidenced by: (i) Bayesian-based analyses, (ii) north-south genetic relationships and diversity gradients; and (iii) a lack of isolation-by-distance in the northernmost cluster. The north-south structure is proposed to have arisen from the processes of postglacial colonization as well as recent climate-driven changes in population dynamics. Our data support the hypothesis of multiple sources of origin for the outbreak and point to the need for population specific information to improve our understanding and management of outbreaks. The recent range expansion across the Rocky Mountains into the jack/lodgepole hybrid and pure jack pine zones of northern Alberta is consistent with a northern British Columbia origin. We detected no loss of genetic variability in these populations, indicating that the evolutionary potential of mountain pine beetle to adapt has not been reduced by founder events. This study illustrates a rapid range-wide response to the removal of climatic constraints, and the potential for range expansion of a regional population.

Characterizing the physical and genetic structure of the lodgepole pine × jack pine hybrid zone: mosaic structure and differential introgression.

Understanding the physical and genetic structure of hybrid zones can illuminate factors affecting their formation and stability. In north-central Alberta, lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia) and jack pine (Pinus banksiana Lamb) form a complex and poorly defined hybrid zone. Better knowledge of this zone is relevant, given the recent host expansion of mountain pine beetle into jack pine. We characterized the zone by genotyping 1998 lodgepole, jack pine, and hybrids from British Columbia, Alberta, Saskatchewan, Ontario, and Minnesota at 11 microsatellites. Using Bayesian algorithms, we calculated genetic ancestry and used this to model the relationship between species occurrence and environment. In addition, we analyzed the ancestry of hybrids to calculate the genetic contribution of lodgepole and jack pine. Finally, we measured the amount of gene flow between the pure species. We found the distribution of the pine classes is explained by environmental variables, and these distributions differ from classic distribution maps. Hybrid ancestry was biased toward lodgepole pine; however, gene flow between the two species was equal. The results of this study suggest that the hybrid zone is complex and influenced by environmental constraints. As a result of this analysis, range limits should be redefined.

Spatial genetic structure of a symbiotic beetle-fungal system: toward multi-taxa integrated landscape genetics.

Spatial patterns of genetic variation in interacting species can identify shared features that are important to gene flow and can elucidate co-evolutionary relationships. We assessed concordance in spatial genetic variation between the mountain pine beetle (Dendroctonus ponderosae) and one of its fungal symbionts, Grosmanniaclavigera, in western Canada using neutral genetic markers. We examined how spatial heterogeneity affects genetic variation within beetles and fungi and developed a novel integrated landscape genetics approach to assess reciprocal genetic influences between species using constrained ordination. We also compared landscape genetic models built using Euclidean distances based on allele frequencies to traditional pair-wise Fst. Both beetles and fungi exhibited moderate levels of genetic structure over the total study area, low levels of structure in the south, and more pronounced fungal structure in the north. Beetle genetic variation was associated with geographic location while that of the fungus was not. Pinevolume and climate explained beetle genetic variation in the northern region of recent outbreak expansion. Reciprocal genetic relationships were only detectedin the south where there has been alonger history of beetle infestations. The Euclidean distance and Fst-based analyses resulted in similar models in the north and over the entire study area, but differences between methods in the south suggest that genetic distances measures should be selected based on ecological and evolutionary contexts. The integrated landscape genetics framework we present is powerful, general, and can be applied to other systems to quantify the biotic and abiotic determinants of spatial genetic variation within and among taxa.

Two-dimensional wavelet analysis of spruce budworm host basal area in the Border Lakes landscape.

Increases in the extent and severity of spruce budworm (Choristoneura fumiferana Clem.) outbreaks over the last century are thought to be the result of changes in forest structure due to forest management. A corollary of this hypothesis is that manipulations of forest structure and composition can be used to reduce future forest vulnerability. However, to what extent historical forest management has influenced current spatial patterns of spruce budworm host species is unknown. To identify landscape-scale spatial legacies of forest management in patterns of spruce budworm host species (i.e., Abies balsamea and Picea spp.), we analyzed remotely sensed forest data from the Border Lakes landscape of northern Minnesota and northwestern Ontario. Our study area contains three regions with different management histories: (1) fine-scale logging patterns in Minnesota, (2) coarse-scale logging patterns in Ontario, and (3) very limited logging history in the Boundary Waters Canoe Area and adjacent Quetico Provincial Park. We analyzed forest basal-area data using wavelets and null models to identify: (1) at which scales forest basal area is structured, (2) where those scales of pattern are significantly present, and (3) whether regions of local significance correspond to regional boundaries that separate the study area. Results indicate that spatial patterns in host basal area are created by nonstationary processes and that these processes are further constrained by lakes and wetlands. Wavelet analysis combined with significance testing revealed a bimodal distribution of scale-specific wavelet variance and separate zones of host species basal area that partially correspond with regional boundaries, particularly between Minnesota and the Wilderness region. This research represents one of the first comparisons of forest spatial structure in this region across an international border and presents a novel method of two-dimensional wavelet analysis that can be used to identify significant scale-specific structure in spatial data.

Spatial community structure of mountain pine beetle fungal symbionts across a latitudinal gradient.

Symbiont redundancy in obligate insect-fungal systems is thought to buffer the insect host against symbiont loss and to extend the environmental conditions under which the insect can persist. The mountain pine beetle is associated with at least three well-known and putatively obligate ophiostomatoid fungal symbionts that vary in their environmental tolerances. To better understand the spatial variation in beetle-fungal symbiotic associations, we examined the community composition of ophiostomatoid fungi associated with the mountain pine beetle as a function of latitude and elevation. The region investigated represents the leading edge of a recent outbreak of mountain pine beetle in western Canada. Using regression and principal components analysis, we identified significant spatial patterns in fungal species abundances that indicate symmetrical replacement between two of the three fungi along a latitudinal gradient and little variation in response to elevation. We also identified significant variation in the prevalence of pair-wise species combinations that occur within beetle galleries. Frequencies of pair-wise combinations were significantly different from what was expected given overall species abundances. These results suggest that complex processes of competitive exclusion and coexistence help determine fungal community composition and that the consequences of these processes vary spatially. The presence of three fungal symbionts in different proportions and combinations across a wide range of environmental conditions may help explain the success of mountain pine beetle attacks across a broad geographic range.

Considering spatial and temporal scale in landscape-genetic studies of gene flow.

Landscape features exist at multiple spatial and temporal scales, and these naturally affect spatial genetic structure and our ability to make inferences about gene flow. This article discusses how decisions about sampling of genotypes (including choices about analytical methods and genetic markers) should be driven by the scale of spatial genetic structure, the time frame that landscape features have existed in their current state, and all aspects of a species' life history. Researchers should use caution when making inferences about gene flow, especially when the spatial extent of the study area is limited. The scale of sampling of the landscape introduces different features that may affect gene flow. Sampling grain should be smaller than the average home-range size or dispersal distance of the study organism and, for raster data, existing research suggests that simplifying the thematic resolution into discrete classes may result in low power to detect effects on gene flow. Therefore, the methods used to characterize the landscape between sampling sites may be a primary determinant for the spatial scale at which analytical results are applicable, and the use of only one sampling scale for a particular statistical method may lead researchers to overlook important factors affecting gene flow. The particular analytical technique used to correlate landscape data and genetic data may also influence results; common landscape-genetic methods may not be suitable for all study systems, particularly when the rate of landscape change is faster than can be resolved by common molecular markers.

Utility of computer simulations in landscape genetics.

Population genetics theory is primarily based on mathematical models in which spatial complexity and temporal variability are largely ignored. In contrast, the field of landscape genetics expressly focuses on how population genetic processes are affected by complex spatial and temporal environmental heterogeneity. It is spatially explicit and relates patterns to processes by combining complex and realistic life histories, behaviours, landscape features and genetic data. Central to landscape genetics is the connection of spatial patterns of genetic variation to the usually highly stochastic space-time processes that create them over both historical and contemporary time periods. The field should benefit from a shift to computer simulation approaches, which enable incorporation of demographic and environmental stochasticity. A key role of simulations is to show how demographic processes such as dispersal or reproduction interact with landscape features to affect probability of site occupancy, population size, and gene flow, which in turn determine spatial genetic structure. Simulations could also be used to compare various statistical methods and determine which have correct type I error or the highest statistical power to correctly identify spatio-temporal and environmental effects. Simulations may also help in evaluating how specific spatial metrics may be used to project future genetic trends. This article summarizes some of the fundamental aspects of spatial-temporal population genetic processes. It discusses the potential use of simulations to determine how various spatial metrics can be rigorously employed to identify features of interest, including contrasting locus-specific spatial patterns due to micro-scale environmental selection.