Context

While remote sensing imagery is effective for quantifying land cover changes across large areas, its utility for directly assessing the response of animals to disturbance is limited. Soundscapes approaches—the recording and analysis of sounds in a landscape—could address this shortcoming.

Objectives

In 2011, a massive wildfire named “the Horseshoe 2 Burn” occurred in the Chiricahua National Monument, Arizona, USA. We evaluated the impact of this wildfire on acoustic activity of animal communities.

Methods

In 2013, soundscape recordings were collected over 9 months in 12 burned and 12 non-burned sites in four ecological systems. The seasonal and diel biological acoustic activity were described using the “Bioacoustic Index”, a detailed aural analysis of sound sources, and a new tool called “Sonic Timelapse Builder” (STLB).

Results

Seasonal biophony phenology showed a diurnal peak in June and a nocturnal peak in October in all ecological systems. On June mornings, acoustic activity was lower at burned than at non-burned sites in three of four ecological systems, due to a decreased abundance of cicadas directly impacted by the death of trees. Aural analyses revealed that 55% of recordings from non-burned sites contained insect sounds compared to 18% from burned sites. On October nights, orthopteran activity was more prevalent at some burned sites, possibly due to post-fire emergence of herbaceous.

Conclusions

Soundscape approaches can help address long-term conservation issues involving the responses of animal communities to wildfire. Acoustic methods can serve as a valuable complement to remote sensing for disturbance-based landscape management.

Context

Growing evidence suggests that climate change could substantially alter forest disturbances. Interactions between individual disturbance agents are a major component of disturbance regimes, yet how interactions contribute to their climate sensitivity remains largely unknown.

Objectives

Here, our aim was to assess the climate sensitivity of disturbance interactions, focusing on wind and bark beetle disturbances.

Methods

We developed a process-based model of bark beetle disturbance, integrated into the dynamic forest landscape model iLand (already including a detailed model of wind disturbance). We evaluated the integrated model against observations from three wind events and a subsequent bark beetle outbreak, affecting 530.2 ha (3.8 %) of a mountain forest landscape in Austria between 2007 and 2014. Subsequently, we conducted a factorial experiment determining the effect of changes in climate variables on the area disturbed by wind and bark beetles separately and in combination.

Results

iLand was well able to reproduce observations with regard to area, temporal sequence, and spatial pattern of disturbance. The observed disturbance dynamics was strongly driven by interactions, with 64.3 % of the area disturbed attributed to interaction effects. A +4 °C warming increased the disturbed area by +264.7 % and the area-weighted mean patch size by +1794.3 %. Interactions were found to have a ten times higher sensitivity to temperature changes than main effects, considerably amplifying the climate sensitivity of the disturbance regime.

Conclusions

Disturbance interactions are a key component of the forest disturbance regime. Neglecting interaction effects can lead to a substantial underestimation of the climate change sensitivity of disturbance regimes.

Context

Primates are an important component of biodiversity in tropical regions. However, many studies on the effects of habitat change on primates ignore the relative influence of landscape composition and configuration.

Objectives

This study addresses the question: how important are landscape-scale forest area and composition relative to patch-scale (1–1080 ha) and site-scale (transect of 1 km) habitat variables for the occupancy and abundance of four primate species in the Colombian Llanos.

Methods

Using a randomly stratified survey design, 81 fragments were surveyed for primate occupancy and abundance. We used zero-inflated models to test the relative influence of landscape-scale, patch-scale and site-scale variables on occupancy and abundance for each species. A 95% confidence set of models was constructed using the cumulative Akaike weight for each model and the relative importance of each set of variables calculated for each primate species.

Results

Occupancy was determined by a combination of site-scale, patch-scale and landscape-scale variables but this varied substantially among the primate species.

Conclusion

Our study highlights the importance of managing primates at a range of scales that considers the relative importance of site-, patch- and landscape-scale variables.

Context

Despite decades of research, there is an intense debate about the consistency of the hump-shaped pattern describing the relationship between diversity and disturbance as predicted by the intermediate disturbance hypothesis (IDH). Previous meta-analyses have not explicitly considered interactive effects of disturbance frequency and intensity of disturbance on plant species diversity in terrestrial landscapes.

Objective

We conducted meta-analyses to test the applicability of IDH by simultaneously examining the relationship between species richness, disturbance frequency (quantified as time since last disturbance as originally proposed) and intensity of disturbance in forest landscapes.

Methods

The effects of disturbance frequency, intensity, and their interaction on species richness was evaluated using a mixed-effects model.

Results

We found that species richness peaks at intermediate frequency after both high and intermediate disturbance intensities, but the richness-frequency relationship differed between intensity classes.

Conclusions

Our study highlights the need to measure multiple disturbance components that could help reconcile conflicting empirical results on the effect of disturbance on plant species diversity.

Context

No single model can capture the complex species range dynamics under changing climates—hence the need for a combination approach that addresses management concerns.

Objective

A multistage approach is illustrated to manage forested landscapes under climate change. We combine a tree species habitat model—DISTRIB II, a species colonization model—SHIFT, and knowledge-based scoring system—MODFACs, to illustrate a decision support framework.

Methods

Using shortleaf pine (Pinus echinata) and sugar maple (Acer saccharum) as examples, we project suitable habitats under two future climate change scenarios (harsh, Hadley RCP8.5 and mild CCSM RCP4.5 at ~2100) at a resolution of 10 km and assess the colonization likelihood of the projected suitable habitats at a 1 km resolution; and score biological and disturbance factors for interpreting modeled outcomes.

Results

Shortleaf pine shows increased habitat northward by 2100, especially under the harsh scenario of climate change, and with higher possibility of natural migration confined to a narrow region close to the current species range boundary. Sugar maple shows decreased habitat and has negligible possibility of migration within the US due to a large portion of its range being north of the US border. Combination of suitable habitats with colonization likelihoods also allows for identification of potential locations appropriate for assisted migration, should that be deemed feasible.

Conclusion

The combination of these multiple components using diverse approaches leads to tools and products that may help managers make management decisions in the face of a changing climate.

Context

Interactions among disturbances, climate, and vegetation influence landscape patterns and ecosystem processes. Climate changes, exotic invasions, beetle outbreaks, altered fire regimes, and human activities may interact to produce landscapes that appear and function beyond historical analogs.

Objectives

We used the mechanistic ecosystem-fire process model FireBGCv2 to model interactions of wildland fire, mountain pine beetle (Dendroctonus ponderosae), and white pine blister rust (Cronartium ribicola) under current and future climates, across three diverse study areas.

Methods

We assessed changes in tree basal area as a measure of landscape response over a 300-year simulation period for the Crown of the Continent in north-central Montana, East Fork of the Bitterroot River in western Montana, and Yellowstone Central Plateau in western Wyoming, USA.

Results

Interacting disturbances reduced overall basal area via increased tree mortality of host species. Wildfire decreased basal area more than beetles or rust, and disturbance interactions modeled under future climate significantly altered landscape basal area as compared with no-disturbance and current climate scenarios. Responses varied among landscapes depending on species composition, sensitivity to fire, and pathogen and beetle suitability and susceptibility.

Conclusions

Understanding disturbance interactions is critical for managing landscapes because forest responses to wildfires, pathogens, and beetle attacks may offset or exacerbate climate influences, with consequences for wildlife, carbon, and biodiversity.

Context

Recent conceptual developments in ecosystem services research have revealed the need to elucidate the complex and unintended relationships between humans and the environment if we are to better understand and manage ecosystem services in practice.

Objectives

This study aimed to develop a model that spatially represents a complex human–environment (H–E) system consisting of heterogeneous social–ecological components and feedback mechanisms at multiple scales, in order to assess multi-dimensional (spatial, temporal, and social) trade-offs in ecosystem services.

Methods

We constructed an agent-based model and empirically calibrated it for a semi-arid region in Northeast China, and examined ecosystem service trade-offs derived from the Sloping Land Conversion Program (SLCP), which is based on payment for ecosystem services. This paper describes our model, named Inner Mongolia Land Use Dynamic Simulator (IM-LUDAS), using the overview, design concepts, and details + decision (ODD + D) protocol and demonstrates the capabilities of IM-LUDAS through simulations.

Results

IM-LUDAS represented typical characteristics of complex H–E systems, such as secondary and cross-scale feedback loops, time lags, and threshold change, revealing the following results: tree plantations expanded by the SLCP facilitated vegetation and soil restoration and household change toward off-farm livelihoods, as expected by the government; conversely, the program caused further land degradation outside the implementation plots; moreover, the livelihood changes were not large enough to compensate for income deterioration by policy-induced reduction in cropland.

Conclusions

IM-LUDAS proved itself to be an advanced empirical model that can recreate essential features of complex H–E systems and assess multi-dimensional trade-offs in ecosystem services.

Context

The positive correlation between landscape area of semi-natural habitat and wild pollinator richness and abundance in agroecosystems has been well studied. However, we lack a deep understanding of local scale floral resource and nest provisioning for wild bees necessary to optimize implementation of pollinator conservation practices.

Objectives

The primary objective of this study was to use a spatially interactive landscape pollination model (hereafter, the Lonsdorf model) to represent field scale spatial patterns of wild bee abundance and richness within a heterogeneous landscape in the mid-Atlantic USA.

Methods

We parameterized the Lonsdorf model with high resolution aerial imagery and insight from a previously published floristic study. To test the Lonsdorf model predictions, field studies were conducted to measure wild bee abundance and species richness in apple orchards as a function of distance from a forest edge.

Results

Field measurements indicated apple pollinator abundance and species richness significantly decreased with increasing distance from the forest edge. The Lonsdorf model pollination service score was highly sensitive to changes in resource provisioning in orchard and non-crop areas, and including resource rich forest and forest edge habitats in the model significantly improved pollination service estimates.

Conclusions

We demonstrated a novel application of the Lonsdorf model at a field scale to predict trends in pollination service provisioning as a factor of local habitat features. With sufficiently detailed inputs, the Lonsdorf model is a promising tool to quantify field scale pollination service deficits, guiding more cost effective habitat supplementation and other conservation efforts.

Context

Forests throughout eastern North America continue to recover from broad-scale intensive land use that peaked in the nineteenth century. These forests provide essential goods and services at local to global scales. It is uncertain how recovery dynamics, the processes by which forests respond to past forest land use, will continue to influence future forest conditions. Climate change compounds this uncertainty.

Objectives

We explored how continued forest recovery dynamics affect forest biomass and species composition and how climate change may alter this trajectory.

Methods

Using a spatially explicit landscape simulation model incorporating an ecophysiological model, we simulated forest processes in New England from 2010 to 2110. We compared forest biomass and composition from simulations that used a continuation of the current climate to those from four separate global circulation models forced by a high emission scenario (RCP 8.5).

Results

Simulated forest change in New England was driven by continued recovery dynamics; without the influence of climate change forests accumulated 34 % more biomass and succeed to more shade tolerant species; Climate change resulted in 82 % more biomass but just nominal shifts in community composition. Most tree species increased AGB under climate change.

Conclusions

Continued recovery dynamics will have larger impacts than climate change on forest composition in New England. The large increases in biomass simulated under all climate scenarios suggest that climate regulation provided by the eastern forest carbon sink has potential to continue for at least a century.

Context

Forest landscape models (FLMs) are important tools for simulating forest changes over broad spatial and temporal scales. The ability of FLMs to accurately predict forest changes may be significantly influenced by the formulations of site-scale processes including seedling establishment, tree growth, competition, and mortality.

Objective

The objectives of this study were to investigate the effects of site-scale processes and interaction effects of site-scale processes and harvest on landscape-scale forest change predictions.

Methods

We compared the differences in species’ distribution (quantified by species’ percent area), total aboveground biomass, and species’ biomass derived from two FLMs: (1) a model that explicitly incorporates stand density and size for each species age cohort (LANDIS PRO), and (2) a model that explicitly tracks biomass for each species age cohort (LANDIS-II with biomass succession extension), which are variants from the LANDIS FLM family with different formulations of site-scale processes.

Results

For early successional species, the differences in simulated distribution and biomass were small (mostly less than 5 %). For mid- to late-successional species, the differences in simulated distribution and biomass were relatively large (10–30 %). The differences in species’ biomass predictions were generally larger than those for species’ distribution predictions. Harvest mediated the differences on landscape-scale predictions.

Conclusions

The effects of site-scale processes on landscape-scale forest change predictions are dependent on species’ ecological traits such as shade tolerance, seed dispersal, and growth rates.

Context

Policy decisions form a major driver of land use change, with important implications for socially and environmentally susceptible regions. It is well known that there can be major unintended consequences, especially where policies are not tailored to regionally specific contexts.

Objectives

In this paper we assess the implications of 60 years of agricultural policies on soil erosion prevention (SEP) by vegetation, an essential regulating ecosystem service in Mediterranean Europe.

Methods

To assess these implications we produced and analysed a time series of land cover/use and environmental conditions datasets (from 1951 to 2012) in relation to changing agricultural policies for a specific region in the southern Portugal. A set of indicators related to SEP allowed us to identify that land use intensification as increased soil erosion in the last 60 years.

Results

Particularly in the last 35 years, as a consequence of headage payments for cattle, the agricultural policy had a significant effect in the density and renewal of the tree cover, resulting in drastic effects for the provision of the SEP service. These are more significant after 1986, coinciding with the implementation of several Common Agricultural Policy instruments focused on increasing the modernization and productivity capacity of farm systems.

Conclusions

The results show some unintended effects of agricultural policy mechanisms on ecosystem service provision and highlight the need for context-based policies, tailored to the environmental constrains and potentials of each region.

Context

Urbanisation places increasing stress on ecosystem services; however existing methods and data for testing relationships between service delivery and urban landscapes remain imprecise and uncertain. Unknown impacts of scale are among several factors that complicate research. This study models ecosystem services in the urban area comprising the towns of Milton Keynes, Bedford and Luton which together represent a wide range of the urban forms present in the UK.

Objectives

The objectives of this study were to test (1) the sensitivity of ecosystem service model outputs to the spatial resolution of input data, and (2) whether any resultant scale dependency is constant across different ecosystem services and model approaches (e.g. stock- versus flow-based).

Methods

Carbon storage, sediment erosion, and pollination were modelled with the InVEST framework using input data representative of common coarse (25 m) and fine (5 m) spatial resolutions.

Results

Fine scale analysis generated higher estimates of total carbon storage (9.32 vs. 7.17 kg m^?2) and much lower potential sediment erosion estimates (6.4 vs. 18.1 Mg km^?2 year^?1) than analyses conducted at coarser resolutions; however coarse-scale analysis estimated more abundant pollination service provision.

Conclusions

Scale sensitivities depend on the type of service being modelled; stock estimates (e.g. carbon storage) are most sensitive to aggregation across scales, dynamic flow models (e.g. sediment erosion) are most sensitive to spatial resolution, and ecological process models involving both stocks and dynamics (e.g. pollination) are sensitive to both. Care must be taken to select model data appropriate to the scale of inquiry.

Context

An increase in the incidence of large wildfires worldwide has prompted concerns about the resilience of forest ecosystems, particularly in the western U.S., where recent changes are linked with climate warming and 20th-century land management practices.

Objectives

To study forest resilience to recent wildfires, we examined relationships among fire legacies, landscape features, ecological conditions, and patterns of post-fire conifer regeneration.

Methods

We quantified regeneration across 182 sites in 21 recent large fires in dry mixed-conifer forests of the U.S. northern Rockies. We used logistic and negative binomial regression to predict the probability of establishment and abundance of conifers 5–13 years post-fire.

Results

Seedling densities varied widely across all sites (0–127,500 seedlings ha^?1) and were best explained by variability in distance to live seed sources (β = ?0.014, p = 0.002) and pre-fire tree basal area (β = 0.072, p = 0.008). Beyond 95 m from the nearest live seed source, the probability of seedling establishment was low. Across all the fires we studied, 75 % of the burned area with high tree mortality was within this 95-m threshold, suggesting the presence of live seed trees to facilitate natural regeneration.  

Conclusions

Combined with the mix of species present within the burn mosaic, dry mixed-conifer forests will be resilient to large fires across our study region, provided that seedlings survive, fire do not become more frequent, high-severity patches do not get significantly larger, and post-fire climate conditions remain suitable for seedling establishment and survival.

Context

Global temperatures are projected to increase and affect forests and wildlife populations. Forest management can potentially mitigate climate-induced changes through promoting carbon sequestration, forest resilience, and facilitated change.

Objectives

We modeled direct and indirect effects of climate change on avian abundance through changes in forest landscapes and assessed impacts on bird abundances of forest management strategies designed to mitigate climate change effects.

Methods

We coupled a Bayesian hierarchical model with a spatially explicit landscape simulation model (LANDIS PRO) to predict avian relative abundance. We considered multiple climate scenarios and forest management scenarios focused on carbon sequestration, forest resilience, and facilitated change over 100 years.

Results

Management had a greater impact on avian abundance (almost 50% change under some scenarios) than climate (<3% change) and only early successional and coniferous forest showed significant change in percent cover across time. The northern bobwhite was the only species that changed in abundance due to climate-induced changes in vegetation. Northern bobwhite, prairie warbler, and blue-winged warbler generally increased in response to warming temperatures but prairie warbler exhibited a non-linear response and began to decline as summer maximum temperatures exceeded 36 °C at the end of the century.

Conclusion

Linking empirical models with process-based landscape change models can be an effective way to predict climate change and management impacts on wildlife, but time frames greater than 100 years may be required to see climate related effects. We suggest that future research carefully consider species-specific effects and interactions between management and climate.

Context

Pasture-woodlands are semi-natural landscapes that result from the combined influences of climate, management, and intrinsic vegetation dynamics. These landscapes are sensitive to future changes in land use and climate, but our ability to predict the impact on ecosystem service provisioning is limited due to the disparate scales in time and space that govern their dynamics.

Objectives

To develop a process-based model to simulate pasture-woodland landscapes and the provisioning of ecosystem services (i.e., livestock forage, woody biomass and landscape heterogeneity).

Methods

We modified a dynamic forest landscape model to simulate pasture-woodland landscapes in Switzerland. This involved including an annual herbaceous layer, selective grazing from cattle, and interactions between grazing and tree recruitment. Results were evaluated within a particular pasture, and then the model was used to simulate regional vegetation patterns and livestock suitability for a ~198,000 ha landscape in the Jura Vaudois region.

Results

The proportion of vegetation cover types at the pasture level (i.e., open, semi-open and closed forests) was well represented, but the spatial distribution of trees was only broadly similar. The entire Jura Vaudois region was simulated to be highly suitable for livestock, with only a small proportion being unsuitable due to steep slopes and high tree cover. High and low elevation pastures were equally suitable for livestock, as lower forage production at higher elevations was compensated by reduced tree cover.

Conclusions

The modified model is valuable for assessing landscape to regional patterns in vegetation and livestock, and offers a platform to evaluate how climate and management impact ecosystem services.

Context

Interactions between landscape-scale processes and fine-grained habitat heterogeneity are usually invoked to explain species occupancy in fragmented landscapes. In variegated landscapes, however, organisms face continuous variation in micro-habitat features, which makes necessary to consider ecologically meaningful estimates of habitat quality at different spatial scales.

Objectives

We evaluated the spatial scales at which forest cover and tree quality make the greatest contribution to the occupancy of the long-horned beetle Microplophorus magellanicus (Coleoptera: Cerambycidae) in a variegated forest landscape.

Methods

We used averaged data of tree quality (as derived from remote sensing estimates of the decay stage of single trees) and spatially independent pheromone-baited traps to model the occurrence probability as a function of multiple cross-scale combinations between forest cover and tree quality (with scales ranging between 50 and 400 m).

Results

Model support and performance increased monotonically with the increasing scale at which tree quality was measured. Forest cover was not significant, and did not exhibit scale-specific effects on the occurrence probability of M. magellanicus. The interactive effect between tree quality and forest cover was stronger than the independent (additive) effects of tree quality and particularly forest cover. Significant interactions included tree quality measured at spatial scales ≥200 m, but cross-scale interactions occurred only in four of the seven best-supported models.

Conclusions

M. magellanicus respond to the high-quality trees available in the landscape rather than to the amount of forest per se. Conservation of viable metapopulations of M. magellanicus should consider the quality of trees at spatial scales >200 m.

Context

Although forest fragmentation is generally thought to impact tree growth and mortality negatively, recent work suggests some forests are resilient. Experimental forests provide an opportunity to examine the timing and extent of forest tree resilience to disturbance from fragmentation.

Objectives

We used the Wog Wog Habitat Fragmentation Experiment in southeastern Australia to test Eucalyptus growth and survivorship responses to forest fragmentation over a 26 year period.

Methods

We measured 2418 tree diameters and used spline-regression techniques to examine non-monotonic fragmentation effect over two time periods.

Results

Over the first 4 years after fragmentation, individual eucalypt tree growth was greater than in continuous forest for large trees and mortality rates were higher only within 10 m of edges. Over the following 22 years only the effects on tree growth remained and on average all fragments rebounded so that their biomass and mortality rates were equivalent to continuous forest. Importantly non-monotonic patterns were observed in growth and mortality with respect to area and distance from edge in both study periods, demonstrating that fragmentation impacts on trees can be strong in localized areas (greatest in 3 ha fragments and 0–30 m edges) and over short time periods.

Conclusions

Dry-sclerophyll eucalypt forests join the set of forest types that display resilient growth dynamics post fragmentation. Moreover, persistent non-monotonic impacts on tree growth with respect to tree size, fragment area, and fragment distance from edge, highlighting landscape fragmentation as a driver of habitat heterogeneity within remnant forest fragments.

Context

Habitat destruction is the leading threat to terrestrial biodiversity, isolating remnant habitat in a matrix of modified vegetation.

Objectives

Our goal was to determine how species richness in several broad taxonomic groups from remnant forest was influenced by matrix quality, which we characterized by comparing plant biomass in forest and the surrounding matrix.

Methods

We coupled data on species-area relationships (SARs) in forest remnants from 45 previously published studies with an index of matrix quality calculated using new estimates of plant biomass derived from satellite imagery.

Results

The effect size of SARs was greatest in landscapes with low matrix quality and little forest cover. SARs were generally stronger for volant than for non-volant species. For the terrestrial taxa included in our analysis, matrix quality decreased as the proportion of water, ice, or urbanization in a landscape increased.

Conclusions

We clearly demonstrate that matrix quality plays a major role in determining patterns of species richness in remnant forest. A key implication of our work is that activities that increase matrix quality, such as active and passive habitat restoration, may be important conservation measure for maintaining and restoring biodiversity in modified landscapes.

Context

Landscape heterogeneity (the composition and configuration of different landcover types) plays a key role in shaping woodland bird assemblages in wooded-agricultural mosaics. Understanding how species respond to landscape factors could contribute to preventing further decline of woodland bird populations.

Objective

To investigate how woodland birds with different species traits respond to landscape heterogeneity, and to identify whether specific landcover types are important for maintaining diverse populations in wooded-agricultural environments.

Methods

Birds were sampled from woodlands in 58 2 × 2 km tetrads across southern Britain. Landscape heterogeneity was quantified for each tetrad. Bird assemblage response was determined using redundancy analysis combined with variation partitioning and response trait analyses.

Results

For woodland bird assemblages, the independent explanatory importance of landscape composition and landscape configuration variables were closely interrelated. When considered simultaneously during variation partitioning, the community response was better represented by compositional variables. Different species responded to different landscape features and this could be explained by traits relating to woodland association, foraging strata and nest location. Ubiquitous, generalist species, many of which were hole-nesters or ground foragers, correlated positively with urban landcover while specialists of broadleaved woodland avoided landscapes containing urban areas. Species typical of coniferous woodland correlated with large conifer plantations.

Conclusions

At the 2 × 2 km scale, there was evidence that the availability of resources provided by proximate landcover types was highly important for shaping woodland bird assemblages. Further research to disentangle the effects of composition and configuration at different spatial scales is advocated.

Context

Forest insect outbreaks are influenced by ecological processes operating at multiple spatial scales, including host-insect interactions within stands and across landscapes that are modified by regional-scale variations in climate. These drivers of outbreak dynamics are not well understood for the western spruce budworm, a defoliator that is native to forests of western North America.

Objectives

Our aim was to assess how processes across multiple spatial scales drive western spruce budworm outbreak dynamics. Our objective was to assess the relative importance and influence of a set of factors covering the stand, landscape, and regional scales for explaining spatiotemporal outbreak patterns in British Columbia, Canada.

Methods

We used generalized linear mixed effect models within a multi-model interference framework to relate annual budworm infestation mapped from Landsat time series (1996–2012) to sets of stand-, landscape-, and regional-scale factors derived from forest inventory data, GIS analyses, and climate models.

Results

Outbreak patterns were explained well by our model (R ² = 93%). The most important predictors of infestation probability were the proximity to infestations in the previous year, landscape-scale host abundance, and dry autumn conditions. While stand characteristics were overall less important predictors, we did find infestations were more likely amongst pure Douglas-fir stands with low site indices and high crown closure.

Conclusions

Our findings add to growing empirical evidence that insect outbreak dynamics are driven by multi-scaled processes. Forest management planning to mitigate the impacts of budworm outbreaks should thus consider landscape- and regional-scale factors in addition to stand-scale factors.