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1. Rocky Mountain forests are poised to recover following bark beetle outbreaks but with altered composition

   https://doi.org/10.1111/1365-2745.13999  

   Rodman, Kyle C.; Andrus, Robert A.; Carlson, Amanda R.; Carter, Trevor A.; Chapman, Teresa B.; Coop, Jonathan D.; Fornwalt, Paula J.; Gill, Nathan S.; Harvey, Brian J.; Hoffman, Ashley E.; et al (December 2022, Journal of Ecology)

   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. 

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2. Overlapping outbreaks of multiple bark beetle species are rarely more severe than single‐species outbreaks

   https://doi.org/10.1002/ecs2.4478  

   Tutland, Niko_J; Rodman, Kyle_C; Andrus, Robert_A; Hart, Sarah_J (March 2023, Ecosphere)

   Abstract 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. 

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3. Effects of Bark Beetle Outbreaks on Forest Landscape Pattern in the Southern Rocky Mountains, U.S.A.

   https://doi.org/10.3390/rs13061089  

   Rodman, Kyle C.; Andrus, Robert A.; Butkiewicz, Cori L.; Chapman, Teresa B.; Gill, Nathan S.; Harvey, Brian J.; Kulakowski, Dominik; Tutland, Niko J.; Veblen, Thomas T.; Hart, Sarah J. (March 2021, Remote Sensing)

   null (Ed.)

   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. 

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4. Spruce Beetle Outbreak Increases Streamflow From Snow‐Dominated Basins in Southwest Colorado, USA

   https://doi.org/10.1029/2021WR029964  

   Manning, Aidan L.; Harpold, Adrian; Csank, Adam (April 2022, Water Resources Research)

   Abstract Bark beetles have impacted over 58 million acres of coniferous forest in the Western US since 2000. Most beetle impacted forests are in snow dominated, water limited headwater basins, which generate a disproportionate fraction of water supplies. Previous studies show mixed impacts of bark beetle outbreaks on streamflow with the potential to cause increased or decreased flows, but these studies either predate long‐term snowpack data, are model‐based, or examine only mountain pine beetle outbreaks. Ours is the first study to use an empirical, climate‐normalized paired catchment approach to quantify streamflow response to spruce beetle kill. Using 27 years of climate and streamflow observations from southwest Colorado, we show that in three of the six beetle impacted study basins, annual climate‐normalized streamflow increased by 22%–37% for at least three to 6 years after the beetle outbreak. Impacted basins exhibited no decreased flows and flows in unimpacted control basins remained unchanged. Among impacted basins, no single basin characteristic clearly explained variation of streamflow response. Higher runoff ratios during snowmelt contribute anywhere from 9% to 64% of streamflow increases, implying the importance of both snow and growing season processes in driving streamflow increases. These findings show variable, sometimes substantial streamflow increases in critical water supply basins following beetle kill in subalpine spruce forests, and contrast with evidence of unchanged or decreased streamflow following beetle kill in lower elevation pine forests in colder northern Colorado basins, highlighting the importance of climate and forest composition in refining hydrologic predictions following mountain forest disturbances. 

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5. Increasing rates of subalpine tree mortality linked to warmer and drier summers

   https://doi.org/10.1111/1365-2745.13634  

   Andrus, Robert A.; Chai, Rachel K.; Harvey, Brian J.; Rodman, Kyle C.; Veblen, Thomas T.; Battipaglia, ed., Giovanna (March 2021, Journal of Ecology)

   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. 

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