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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. 

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. 

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. 

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. 

Abstract Warming temperatures and rising moisture deficits are expected to increase the rates of background tree mortality–low amounts of tree mortality (~0.5%–2% year⁻¹), characterizing the forest demographic processes in the absence of abrupt, coarse‐scale disturbance events (e.g. fire). When compounded over multiple decades and large areas, even minor increases in background tree mortality (e.g. <0.5% year⁻¹) can cause changes to forest communities and carbon storage potential that are comparable to or greater than those caused by disturbances.We examine how temporal variability in rates of background tree mortality for four subalpine conifers reflects variability in climate and climate teleconnections using observations of tree mortality from 1982 to 2019 at Niwot Ridge, Colorado, USA. Individually marked trees (initial population 5,043) in 13 permanent plots—located across a range of site conditions, stand ages and species compositions—were censused for new mortality nine times over 37 years.Background tree mortality was primarily attributed to stress from unfavourable climate and competition (71.2%) and bark beetle activity (23.3%), whereas few trees died from wind (5.3%) and wildlife impacts (0.2%). Mean annualized tree mortality attributed to tree stress and bark beetles more than tripled across all stands between initial censuses (0.26% year⁻¹, 1982–1993/1994) and recent censuses (0.82% year⁻¹, 2008–2019). Higher rates of tree mortality were related to warmer maximum summer temperatures, greater summer moisture deficits, and negative anomalies in ENSO (La Niña), with greater effects of drought in some subpopulations (tree size, age and species). For example, in older stands (>250 years), larger and older trees were more likely to die than smaller and younger trees. Differences in tree mortality rates and sensitivity to climate among subpopulations that varied by stand type may lead to unexpected shifts in stand composition and structure.Synthesis. A strong relationship between higher rates of tree mortality and warmer, drier summer climate conditions implies that climate warming will continue to increase background mortality rates in subalpine forests. Combined with increases in disturbances and declining frequency of moist‐cool years suitable for seedling establishment, increasing rates of tree mortality have the potential to drive declines in subalpine tree populations. 

Abstract Changing disturbance regimes and climate can overcome forest ecosystem resilience. Following high-severity fire, forest recovery may be compromised by lack of tree seed sources, warmer and drier postfire climate, or short-interval reburning. A potential outcome of the loss of resilience is the conversion of the prefire forest to a different forest type or nonforest vegetation. Conversion implies major, extensive, and enduring changes in dominant species, life forms, or functions, with impacts on ecosystem services. In the present article, we synthesize a growing body of evidence of fire-driven conversion and our understanding of its causes across western North America. We assess our capacity to predict conversion and highlight important uncertainties. Increasing forest vulnerability to changing fire activity and climate compels shifts in management approaches, and we propose key themes for applied research coproduced by scientists and managers to support decision-making in an era when the prefire forest may not return. 

Tree fecundity and recruitment have not yet been quantified at scales needed to anticipate biogeographic shifts in response to climate change. By separating their responses, this study shows coherence across species and communities, offering the strongest support to date that migration is in progress with regional limitations on rates. The southeastern continent emerges as a fecundity hotspot, but it is situated south of population centers where high seed production could contribute to poleward population spread. By contrast, seedling success is highest in the West and North, serving to partially offset limited seed production near poleward frontiers. The evidence of fecundity and recruitment control on tree migration can inform conservation planning for the expected long-term disequilibrium between climate and forest distribution.