Since the late 1990s, extensive outbreaks of native bark beetles (Curculionidae: Scolytinae) have affected coniferous forests throughout Europe and North America, driving changes in carbon storage, wildlife habitat, nutrient cycling, and water resource provisioning. Remote sensing is a crucial tool for quantifying the effects of these disturbances across broad landscapes. In particular, Landsat time series (LTS) are increasingly used to characterize outbreak dynamics, including the presence and severity of bark beetle-caused tree mortality, though broad-scale LTS-based maps are rarely informed by detailed field validation. Here we used spatial and temporal information from LTS products, in combination with extensive field data and Random Forest (RF) models, to develop 30-m maps of the presence (i.e., any occurrence) and severity (i.e., cumulative percent basal area mortality) of beetle-caused tree mortality 1997–2019 in subalpine forests throughout the Southern Rocky Mountains, USA. Using resultant maps, we also quantified spatial patterns of cumulative tree mortality throughout the region, an important yet poorly understood concept in beetle-affected forests. RF models using LTS products to predict presence and severity performed well, with 80.3% correctly classified (Kappa = 0.61) and R2 = 0.68 (RMSE = 17.3), respectively. We found that ≥10,256 km2 of subalpine forest area (39.5% of the study area) was affected by bark beetles and 19.3% of the study area experienced ≥70% tree mortality over the twenty-three year period. Variograms indicated that severity was autocorrelated at scales < 250 km. Interestingly, cumulative patch-size distributions showed that areas with a near-total loss of the overstory canopy (i.e., ≥90% mortality) were relatively small (<0.24 km2) and isolated throughout the study area. Our findings help to inform an understanding of the variable effects of bark beetle outbreaks across complex forested regions and provide insight into patterns of disturbance legacies, landscape connectivity, and susceptibility to future disturbance.

Changes in climate are altering disturbance regimes in forests of western North America, leading to increases in the potential for disturbance events to overlap in time and space. Though interactions between abiotic and biotic disturbance (e.g., the effect of bark beetle outbreak on subsequent wildfire) have been widely studied, interactions between multiple biotic disturbances are poorly understood. Defoliating insects, such as the western spruce budworm (WSB; Choristoneura freemanni), have been widely suggested to predispose trees to secondary colonization by bark beetles, such as the Douglas-fir beetle (DFB; Dendroctonus pseudotsugae). However, there is little quantitative research that supports this observation. Here, we asked: Does previous WSB damage increase the likelihood of subsequent DFB outbreak in Douglas-fir (Pseudotsuga menziesii) forests of the Southern Rocky Mountains, USA? To quantify areas affected by WSB and then DFB, we analyzed Aerial Detection Survey data from 1999–2019. We found that a DFB presence followed WSB defoliation more often than expected under a null model (i.e., random distribution). With climate change expected to intensify some biotic disturbances, an understanding of the interactions between insect outbreaks is important for forest management planning, as well as for improving our understanding of forest change.

The spatial overlap of multiple ecological disturbances in close succession has the capacity to alter trajectories of ecosystem recovery. Widespread bark beetle outbreaks and wildfire have affected many forests in western North America in the past two decades in areas of important habitat for native ungulates. Bark beetle outbreaks prior to fire may deplete seed supply of the host species, and differences in fire-related regeneration strategies among species may shift the species composition and structure of the initial forest trajectory. Subsequent browsing of postfire tree regeneration by large ungulates, such as elk (Cervus canadensis), may limit the capacity for regeneration to grow above the browse zone to form the next forest canopy. Five stand-replacing wildfires burned~60,000 ha of subalpine forest that had previously been affected by severe (>90% mortality) outbreaks of spruce beetle (SB, Dendroctonus rufipennis) in Engelmann spruce (Picea engelmannii) in 2012-2013 in southwestern Colorado. Here we examine the drivers of variability in abundance of newly established conifer tree seedlings [spruce and subalpine fir (Abies lasiocarpa)] and resprouts of quaking aspen (Populus tremuloides) following the short-interval sequence of SB outbreaks and wildfire (2-8 yr between SB outbreak and fire) at sites where we previously reconstructed severities of SB and fire. We then examine the implications of ungulate browsing for forest recovery. We found that abundances of postfire spruce seedling establishment decreased substantially in areas of severe SB outbreak. Prolific aspen resprouting in stands with live aspen prior to fire will favor an initial postfire forest trajectory dominated by aspen. However, preferential browsing of postfire aspen resprouts by ungulates will likely slow the rate of canopy recovery but browsing is unlikely to alter the species composition of the future forest canopy. Collectively, our results show that SB outbreak prior to fire increases the vulnerability of spruce-fir forests to shifts in forest type (conifer to aspen) and physiognomic community type (conifer forest to non-forest). By identifying where compounded disturbance interactions are likely to limit recovery of forests or tree species, our findings are useful for developing adaptive management strategies in the context of warming climate and shifting disturbance regimes.

Biotic disturbances that overlap in space and time may result in important shifts in forest structure and composition, with potential effects on many ecosystem services. Starting in the late 1990s, outbreaks of multiple bark beetle species caused widespread mortality of three co‐occurring conifer species in the ca. 40,000‐km2 subalpine zone of the southern Rocky Mountains (SRM), USA. To better understand the implications of such outbreaks, our goal was to determine if overlapping outbreaks of multiple bark beetle species caused greater tree mortality than single‐species outbreaks in stands with multiple susceptible host tree species. We mapped stand susceptibility to outbreaks of spruce beetle (SB, Dendroctonus rufipennis), mountain pine beetle (MPB, Dendroctonus ponderosae), and western balsam bark beetle (WBBB, Dryocoetes confusus) by combining aerial survey data and forest composition variables in a random forest modeling framework. Then, we used existing maps of cumulative forest mortality from bark beetles to investigate the extent and severity of overlapping outbreaks from 1999 to 2019. We found that 46% of stands with two or more of the three studied hosts species—Engelmann spruce (Picea engelmannii), lodgepole pine (Pinus contorta var. latifolia), or subalpine fir (Abies lasiocarpa)—were susceptible to overlapping outbreaks (25% of all sampled stands). Of those stands, 31% experienced outbreaks of two or more beetle species. Stands affected by outbreaks of both MPB and SB had higher tree mortality than stands affected by one species alone, though stands susceptible to both MPB and SB were uncommon (<4% of all sampled stands). No other combinations of beetle outbreaks increased tree mortality above levels caused by single‐species outbreaks. Thus, contrary to expectations, overlapping outbreaks were rarely more severe than single‐species outbreaks in the SRM. This suggests that diverse forest communities may buffer against the most severe effects of bark beetle outbreaks, even during warm, dry conditions.