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Over the past several decades, forests worldwide have experienced increases in biotic disturbances caused by insects and plant pathogens – a trend that is expected to continue with climate warming. Whereas the causes and effects of individual biotic disturbances are well studied, spatiotemporal interactions among multiple biotic disturbances are less so, despite their importance to ecosystem function and resilience. Here, we highlight an emerging phenomenon of “hotspots” of biotic disturbances (that is, two or more biotic disturbances that overlap in space and time), documenting trends in recent decades in temperate conifer forests of the western US. We also explore potential mechanisms behind and effects of biotic disturbance hotspots, with particular focus on how altered post‐disturbance recovery (successional pathways) can have profound consequences for ecosystem resilience and biodiversity conservation. Finally, we propose research directions that can elucidate drivers of biotic disturbance hotspots and their ecological effects at various spatial scales, and provide insight into this new knowledge frontier.

 

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‐km²subalpine 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 contortavar.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.

 

Tree die-off, driven by extreme drought and exacerbated by a warming climate, is occurring rapidly across every wooded continent—threatening carbon sinks and other ecosystem services provided by forests and woodlands. Forecasting the spatial patterns of tree die-off in response to drought is a priority for the management and conservation of forested ecosystems under projected future hotter and drier climates. Several thresholds derived from drought-metrics have been proposed to predict mortality ofPinus edulis,a model tree species in many studies of drought-induced tree die-off. To improve future capacity to forecast tree mortality, we used a severe drought as a natural experiment. We compared the ability of existing mortality thresholds derived from four drought metrics (the Forest Drought Severity Index (FDSI), the Standardized Precipitation Evapotranspiration Index, and raw values of precipitation (PPT) and vapor pressure deficit, calculated using 4 km PRISM data) to predict areas ofP. edulisdie-off following an extreme drought in 2018 across the southwestern US. Using aerial detection surveys of tree mortality in combination with gridded climate data, we calculated the agreement between these four proposed thresholds and the presence and absence of regional-scale tree die-off using sensitivity, specificity, and the area under the curve (AUC). Overall, existing mortality thresholds tended to over predict the spatial extent of tree die-off across the landscape, yet some retain moderate skill in discriminating between areas that experienced and did not experience tree die-off. The simple PPT threshold had the highest AUC score (71%) as well as fair sensitivity and specificity, but the FDSI had the greatest sensitivity to die-off (85.9%). We highlight that empirically derived climate thresholds may be useful forecasting tools to identify vulnerable areas to drought induced die-off, allowing for targeted responses to future droughts and improved management of at-risk areas.

 

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. 

1. Amplified by warming temperatures and drought, recent outbreaks of native bark beetles (Curculionidae: Scolytinae) have caused extensive tree mortality throughout Europe and North America. Despite their ubiquitous nature and important effects on ecosystems, forest recovery following such disturbances is poorly understood, particularly across regions with varying abiotic conditions and outbreak effects. 2. To better understand post-outbreak recovery across a topographically complex region, we synthesized data from 16 field studies spanning subalpine forests in the Southern Rocky Mountains, USA. From 1997 to 2019, these forests were heavily affected by outbreaks of three native bark beetle species (Dendroctonus ponderosae, Dendroctonus rufipennis and Dryocoetes confusus). We compared pre- and post-outbreak forest conditions and developed region-wide predictive maps of post-outbreak (1) live basal areas, (2) juvenile densities and (3) height growth rates for the most abundant tree species – aspen (Populus tremuloides), Engelmann spruce (Picea engelmannii), lodgepole pine (Pinus contorta) and subalpine fir (Abies lasiocarpa). 3. Beetle-caused tree mortality reduced the average diameter of live trees by 28.4% (5.6 cm), and species dominance was altered on 27.8% of field plots with shifts away from pine and spruce. However, most plots (82.1%) were likely to recover towards pre-outbreak tree densities without additional regeneration. Region-wide maps indicated that fir and aspen, non-host species for bark beetle species with the most severe effects (i.e. Dendroctonus spp.), will benefit from outbreaks through increased compositional dominance. After accounting for individual size, height growth for all conifer species was more rapid in sites with low winter precipitation, high winter temperatures and severe outbreaks. 4. Synthesis. In subalpine forests of the US Rocky Mountains, recent bark beetle outbreaks have reduced tree size and altered species composition. While eventual recovery of the pre-outbreak forest structure is likely in most places, changes in species composition may persist for decades. Still, forest communities following bark beetle outbreaks are widely variable due to differences in pre-outbreak conditions, outbreak severity and abiotic gradients. This regional variability has critical implications for ecosystem services and susceptibility to future disturbances. 

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. 

Understanding how severe disturbances and their interactions affect forests is key to projecting ecological change under a warming climate. Substantial increases in some biotic disturbances, such as bark beetle outbreaks, in temperate forest ecosystemsmay compromise recovery to a forest vegetation type (i.e., physiognomic recovery or resilience), especially if subsequent biotic disturbances (e.g., herbivory) alter recovery mechanisms. From 2005 to 2017, severe outbreaks (>90% mortality) of spruce bark beetles (SB,Dendroctonus rufipennis) affected Engelmann spruce (Picea engelmannii) across 325,000 ha of spruce and subalpine fir (Abies lasiocarpa) forest in the southern Rocky Mountains, USA. Concurrently, an outbreak of western balsam bark beetle (WBBB,Dryocoetes confuses) infested subalpine fir across at least 47,000 of these hectares. We explored the capacity of 105 stands affected by one or two bark beetle outbreaks and browsing of juvenile trees by ungulates to return to a forest vegetation type in the context of pre‐outbreak forest conditions and topography. Nine initial forest trajectories (i.e., at least several decades) were identified from four pre‐outbreak forest types affected by three biotic disturbances that occurred at different spatial scales and severities. Most stands (86%) contained surviving nonhost adult trees in the main canopy (fir and aspen [Populus tremuloides]) and many surviving juveniles of all species, implying that they are currently on a trajectory for physiognomic recovery. Stands composed exclusively of large‐diameter spruce were affected by a severe SB outbreak and were most vulnerable to a transition to a low‐density forest, below regional stocking levels (<370 trees/ha). Greater pre‐outbreak stand structural complexity and species diversity were key traits of stands with a higher potential for physiognomic recovery. However, all multispecies stands shifted in relative composition of the main canopy to nonhost species, suggesting low potential for compositional recovery over the next several decades. Most post‐outbreak stands (86%) exceeded regional stocking levels with trees taller than the browse zone (<2 m). As such, ungulate browsing on over half of all juveniles will primarily affect the rate of infilling of the forest canopy and preferential browsing of more palatable species will influence the composition of the future forest canopy.

 

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.