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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Bayesian Predictions of Bark Beetle Attack and Mortality of Three Conifer Species During Epidemic and Endemic Population Stages
Bark beetles naturally inhabit forests and can cause large-scale tree mortality when they reach epidemic population numbers. A recent epidemic (1990s–2010s), primarily driven by mountain pine beetles ( Dendroctonus ponderosae ), was a leading mortality agent in western United States forests. Predictive models of beetle populations and their impact on forests largely depend on host related parameters, such as stand age, basal area, and density. We hypothesized that bark beetle attack patterns are also dependent on inferred beetle population densities: large epidemic populations of beetles will preferentially attack large-diameter trees, and successfully kill them with overwhelming numbers. Conversely, small endemic beetle populations will opportunistically attack stressed and small trees. We tested this hypothesis using 12 years of repeated field observations of three dominant forest species (lodgepole pine Pinus contorta , Engelmann spruce Picea engelmannii , and subalpine fir Abies lasiocarpa ) in subalpine forests of southeastern Wyoming paired with a Bayesian modeling approach. The models provide probabilistic predictions of beetle attack patterns that are free of assumptions required by frequentist models that are often violated in these data sets. Furthermore, we assessed seedling/sapling regeneration in response to overstory mortality and hypothesized that higher seedling/sapling establishment occurs in areas with highest overstory mortality because resources are freed from competing trees. Our results indicate that large-diameter trees were more likely to be attacked and killed by bark beetles than small-diameter trees during epidemic years for all species, but there was no shift toward preferentially attacking small-diameter trees in post-epidemic years. However, probabilities of bark beetle attack and mortality increased for small diameter lodgepole pine and Engelmann spruce trees in post-epidemic years compared to epidemic years. We also show an increase in overall understory growth (graminoids, forbs, and shrubs) and seedling/sapling establishment in response to beetle-caused overstory mortality, especially in lodgepole pine dominated stands. Our observations provide evidence of the trajectories of attack and mortality as well as early forest regrowth of three common tree species during the transition from epidemic to post-epidemic stages of bark beetle populations in the field.
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- Award ID(s):
- 1904421
- NSF-PAR ID:
- 10232668
- Date Published:
- Journal Name:
- Frontiers in Forests and Global Change
- Volume:
- 4
- ISSN:
- 2624-893X
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Abstract 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. -
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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 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. -
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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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Abstract 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.
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Accepted Manuscript1.0
- Journal Article:
- https://doi.org/10.3389/ffgc.2021.679104
