MPB Outbreaks Research Articles

Soil Nitrogen Five Years after Bark Beetle Infestation in Lodgepole Pine Forests

The unprecedented scale of recent mountain pine beetle (Dendroctonus ponderosae, MPB) infestation in lodgepole pine (Pinus contorta ex Loudon var. latifolia Engelm. ex S. Watson) forests of western North America may significantly impact the role of forest soils as important sinks for global C and N. The aim of this study was to explain soil and litter N responses in stands experiencing total mortality during the first 5 yr of MPB outbreak. Four-year monitoring (2009–2012) of needlefall deposition, soil water-filled pore space (WFPS), NH₄, NO₃, potentially mineralizable N (PMN), dissolved organic N (DON), N₂O, and CH₄ was established in Medicine Bow National Forest in stands of different infestation histories (infested in 2007, 2008, and uninfested). Results suggest that total stand mortality demonstrated strong temporal dynamics. First-year soil responses included increases in PMN and were followed by WFPS and NH₄ increases in the second year, NO₃ increase in the third year, and DON increase in in the fourth year of the infestation. Litter NH₄ increased within 1 yr, but litter PMN, NO₃, and DON increased 3 yr after the infestation, suggesting enhanced litter decomposability. Soil N₂O and CH₄ fluxes also started to increase 4 yr after the infestation. These changes were indicative of ongoing disequilibrium for the period of at least 5 yr after MPB infestation. The magnitude of N mineralization and N₂O and CH₄ fluxes may continue to increase beyond the initial 5 yr of monitoring because litter-derived labile N will probably contribute to the second pulse of N mineralization.

Harvesting the dead and decaying forests: Potential carbon storage in harvested wood products

British Columbia (BC)'s forests and wood products industry face tremendous adversity in the face of the mountain pine beetle (MPB; Dendroctonus ponderosae) epidemic. The death and decay of one of BC's leading merchantable tree species (lodgepole pine; Pinus contorta Dougl. ex Loud. var. latifolia Engelm.) threatens both the sustainability and climate mitigation opportunities in BC forests. Our study investigates the potential for carbon (C) storage in wood products in the post-epidemic phase of the MPB outbreak. In particular, we contrast the C-balance implications of converting MPB timber into long-lived wood products (i.e., lumber) (scenario 1) versus softwood pellets for bioenergy (scenario 2). Our findings highlight the importance of the C-cycle neutral assumption in determining the net C-balance of lumber versus pellets. Timber harvests for lumber resulted in a net C storage with this assumption (case 1), and net C emissions without this assumption (case 2). With uncertainty surrounding the future C-balance of BC forests, the current assumption of biomass-derived emissions being completely C-neutral may need to be re-evaluated.

Ecological Consequences of Mountain Pine Beetle Outbreaks for Wildlife in Western North American Forests

Mountain pine beetle (Dendroctonus ponderosae) (MPB) outbreaks are increasingly prevalent in western North America, causing considerable ecological change in pine (Pinus spp.) forests with important implications for wildlife. We reviewed studies examining wildlife responses to MPB outbreaks and postoutbreak salvage logging to inform forest management and guide future research. Our review included 16 studies describing MPB outbreak relationships with 89 bird species and 6 studies describing relationships with 11 mammalian species, but no studies of reptiles or amphibians. We included studies that compared wildlife response metrics temporally (before versus after the outbreak) and spatially (across sites that varied in severity of outbreak) in relation to beetle outbreaks. Outbreaks ranged in size from 20,600 to ≥107 ha and studies occurred 1–30 years after the peak MPB outbreak, but most studies were conducted over the short-term (i.e., ≤6 years after the peak of MPB-induced tree mortality). Birds were the only taxa studied frequently; however, high variability existed among those studies to allow many inferences, although some patterns were evident. Avian studies concluded that cavity-nesting species responded more favorably to beetle-killed forests than species with open-cup nests, and species nesting in the shrub layer favored outbreak forests compared with ground and open-cup canopy nesters that generally showed mixed relationships. Bark-drilling species as a group clearly demonstrated a positive short-term association with MPB epidemics compared with that of other foraging assemblages. Cavity-nesting birds that do not consume bark beetles (i.e., secondary cavity-nesting species and nonbark-drilling woodpeckers) also exhibited some positive responses to MPB outbreaks, although not as pronounced or consistent as those of bark-drilling woodpeckers. Mammalian responses to MPB outbreaks were mixed. Studies consistently reported negative effects of MPB outbreaks on red squirrels (Tamiasciurus hudsonicus). However, there is evidence that red squirrels can persist after an outbreak under some conditions, e.g., when nonhost tree species are present. For small mammal species associated with forest understories, responses may be most pronounced during the postepidemic period (>6 years after the peak of beetle-induced tree mortality) when snags fall to produce coarse woody debris. Postoutbreak salvage logging studies (n = 6) reported results that lacked consensus. Postoutbreak salvage logging may have an impact on fewer wildlife species than postfire salvage logging, probably because only host-specific tree species are removed after beetle outbreaks.

An innovative aerial assessment of Greater Yellowstone Ecosystem mountain pine beetle‐caused whitebark pine mortality

An innovative aerial survey method called the Landscape Assessment System (LAS) was used to assess mountain pine beetle (MPB; Dendroctonus ponderosae)-caused mortality of whitebark pine (Pinus albicaulis) across the species distribution in the Greater Yellowstone Ecosystem (GYE; 894 774 ha). This large-scale implementation of the LAS method consisted of 8673 km of flight lines, along which 4653 geo-tagged, oblique aerial photos were captured at the catchment level (a subset of 12-digit USGS hydrologic units) and geographic information system (GIS) processed. The Mountain Pine Beetle-caused Mortality Rating System, a landscape-scale classification system designed specifically to measure the cumulative effects of recent and older MPB attacks on whitebark pine, was used to classify mortality with a rating from 0 to 6 based on the amount of red (recent attack) and gray (old attack) trees visible. The approach achieved a photo inventory of 79% of the GYE whitebark pine distribution. For the remaining 21%, mortality levels were estimated based on an interpolated surface. Results that combine the photo-inventoried and interpolated mortality indicate that nearly half (46%) of the GYE whitebark pine distribution showed severe mortality (3-4 or 5.3-5.4 rating), 36% showed moderate mortality (2-2.9 rating), 13% showed low mortality (1-1.9 rating), and 5% showed trace levels of mortality (0-0.9). These results reveal that the proliferation of MPB in the subalpine zone of the GYE due to climate warming has led to whitebark pine mortality that is more severe and widespread than indicated from either previous modeling research or USDA Forest Service Aerial Detection surveys. Sixteen of the 22 major mountain ranges of the GYE have experienced widespread moderate-to-severe mortality. The majority of catchments in the other six mountain ranges show low-to-moderate mortality. Refugia from MPB outbreaks, at least for now, also exist and correspond to locations that have colder microclimates. The spatially explicit mortality information produced by this project has helped forest managers develop and implement conservation strategies that include both preservation and restoration efforts. Future research aimed at documenting and quantifying the ecological impacts of widespread decline and collapse of this foundation and keystone species is warranted.

Spatiotemporal patterns of mountain pine beetle activity in the southern Rocky Mountains

The current mountain pine beetle (MPB; Dendroctonus ponderosae) outbreak in the southern Rocky Mountains has impacted approximately 750 000 ha of forest. Weather and habitat heterogeneity influence forest insect population dynamics at multiple spatial and temporal scales. Comparison of forest insect population dynamics in two principal host species may elucidate the relative contribution of weather and landscape factors in initiating and driving extensive outbreaks. To investigate potential drivers of the current MPB outbreak, we compared broadscale spatiotemporal patterns of MPB activity in lodgepole pine (Pinus contorta) and ponderosa pine (Pinus ponderosa) from 1996 to 2010 in Colorado and southern Wyoming with regional weather fluctuations, and then tracked the annual meso-scale progression of the epidemic in lodgepole pine with respect to weather, topographic, previous MPB activity, and forest stand attributes. MPB activity in lodgepole pine compared to ponderosa pine showed higher magnitude and extent of spatial synchrony. Warm temperatures and low annual precipitation favorable to beetle populations showed high regional synchrony across areas of both pine species, suggesting that habitat interacts with weather in synchronizing MPB populations. Cluster analysis of time series patterns identified multiple, disjunct locations of incipient MPB activity (epicenters) in lodgepole pine, which overlapped an earlier 1980s MPB outbreak, and suggests a regional trigger (drought) across this homogenous forest type. Negative departures from mean annual precipitation played a key role in subsequent spread of MPB outbreak. Development of the outbreak was also associated with lower elevations, greater dominance by lodgepole pine, stands of larger tree size, and stands with higher percentage canopy cover. After epidemic levels of MPB activity were attained, MPB activity was less strongly associated with stand and weather variables. These results emphasize the importance of considering differences in patterns of MPB dynamics for different host pine species even under similar regional-scale weather variation and the nonstationarity of outbreak dynamics over time.

Lodgepole pine, jack pine, and their hybrids: molecular markers reveal mountain pine beetle host-range expansion into jack pine of the boreal forest

Background Lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia) is found in western North America, extending from the Yukon into British Columbia and Washington, and along the Rocky Mountains and eastern slopes to Colorado [1]. Jack pine (Pinus banksiana Lamb.) is a closely related species found east of the Rockies, mainly in Canada’s boreal forest from the Northwest Territories to Quebec [1]. Lodgepole and jack pine ranges overlap in northern Alberta and the Northwest Territories, where these species hybridize (Fig. 1). In the absence of reliable molecular markers, morphological characteristics are commonly used to distinguish lodgepole pine, jack pine and hybrids. However, hybrids and pure species can be difficult to visually distinguish, particularly at the presumed periphery of the hybrid zone, which is poorly described. Mountain pine beetle (MPB; Dendroctonus ponderosae Hopkins) is indigenous to western North America. In the current MPB outbreak, more than 14 million hectares of mainly lodgepole pine forests have sustained MPB-caused mortality in British Columbia alone [2]. Following MPB long range dispersal into northwestern Alberta in 2006, MPB has continued its apparently unprecedented eastward spread into the lodgepole x jack pine hybrid zone of Alberta (Fig. 1). To better define this hybrid zone, we developed microsatellite and SNP markers that distinguish the pure species from hybrids. We then used the microsatellite markers to test the hypothesis that the MPB epidemic has spread into jack pine.

Regeneration and Growth Following Mountain Pine Beetle Attack: A Synthesis of Knowledge

The mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB) infestation has altered forests of lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia Engelm.) to an unprecedented extent in British Columbia. After an MPB outbreak, advance regeneration significantly contributed to form a new canopy and stand; however, the time needed to form a new stand depends on site-specific conditions. Assessment of regeneration and the growth of residual trees in stands after MPB attack are critical for three purposes: (1) forecasting long-term development (yield) of attacked stands; (2) selecting stands for growth-improving silvicultural treatments; and (3) forecasting impacts to ecological attributes such as hydrology, habitat, and vegetation types. This article reviews and synthesizes recent research concerning lodgepole pine stand performance after MPB attack in British Columbia. Species composition, abundance, spatial distribution, and overall stand health are described. This information is important for forest managers or practitioners who make decisions regarding management of MPB-attacked stands. Moreover, a number of key gaps exist in our knowledge about factors affecting advance regeneration and the residual trees of MPB-attacked stands. This article presents a list of knowledge gaps for management information and further research initiatives.

Changes in stand structure in uneven-aged lodgepole pine stands impacted by mountain pine beetle epidemics and fires in central British Columbia

We examined the development of lodgepole pine (Pinus contorta Dougl.) in uneven-aged stands in the Interior Douglasfir (IDF) biogeoclimatic zone of central of British Columbia (B.C.), which are currently undergoing a massive outbreak of the mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB). Using historical ecological approaches, dendrochronology, and stand measurement data, we determined the roles MPB and fire disturbances have played in the ecological processes of lodgepole pine in an Interior Douglas-fir zone. We found that multiple mixed-severity fires created patchy uneven-aged stands dominated by lodgepole pine. Since fire suppression in the 20th century, multiple MPB disturbances have maintained the structural complexity of the stands and favoured regeneration of lodgepole pine in the understory despite the absence of fire, resulting in self-perpetuating multi-age lodgepole pine stands. Analysis of the stand structures remaining after multiple MPB outbreaks showed that, even with high overstory mortality, the sample stands contained several MPB-initiated cohorts, consisting of younger and smaller-diameter lodgepole pine. These surviving lodgepole pine layers, which are less susceptible to beetle, will provide important ecological legacies, and could play an important role in the mid-term timber supply chain. We concluded that, in the absence of fire, the MPB plays a more frequent role in directing stand dynamics and structure in uneven-aged lodgepole pine stands resulting in selfperpetuating complex stands in the central interior. We compared and contrasted these findings with those obtained in “even-aged” lodgepole pine stands, also in the Interior Douglas-fir zone in the southern interior, which were investigated in an earlier study. Key words: lodgepole pine, mountain pine beetle, dendroecology, complex stands, mixed-severity fire regime

Stand and landscape-level simulations of mountain pine beetle ( Dendroctonus ponderosae) and salvage logging effects on live tree and deadwood habitats in south-central British Columbia, Canada

The effects of mountain pine beetle (MPB, Dendroctonus ponderosae Hopkins) and salvage logging were modeled on live trees and deadwood habitats, including snags and coarse woody debris (CWD), in stands that varied by lodgepole pine ( Pinus contorta Dougl.) dominance, age and growing site quality. Results were projected on a lodgepole pine-dominated landscape in south-central British Columbia, Canada to examine differences in the density, distribution and type of live tree and deadwood habitats following the MPB outbreak and salvage logging. Tree mortality caused by MPB varied considerably with percent pine and factors that affect tree size, including tree age, size and growing site. Mortality ranged as much as 38–86% even in 100% pine stands, creating a range of residual live tree levels and a pulse of snags that contributes to the increased total and well-decayed CWD volumes over time. In contrast, salvage logging consistently reduces large live trees and snags and contributes to long-term declines in both total and well-decayed snag and CWD volume. Compared to the MPB outbreak, widespread salvage logging will leave a paucity of live trees, snags and CWD in cutover areas on the landscape. This becomes a concern if ongoing natural disturbances and timber harvest deplete remaining mature forests and few intermediate aged forests (40–100 yrs) exist to recruit new live tree and deadwood habitats in the future. Careful planning prior to salvage logging is required to maintain and recruit adequate live tree and deadwood habitats to support native wildlife in future landscapes.

Influence of fire and mountain pine beetle on the dynamics of lodgepole pine stands in British Columbia, Canada

An outbreak of the mountain pine beetle ( Dendroctonus ponderosae Hopkins; MPB), currently affecting over 10.1 million hectares of lodgepole pine forests ( Pinus contorta Dougl.) in British Columbia, Canada, is the largest in recorded history. We examined the dynamics of even-aged lodgepole pine forests in southern British Columbia, which were undergoing this MPB outbreak. Using dendroecology and forest measurements we reconstructed the stand processes of stand initiation, stand disturbances, tree mortality, and regeneration, and explained the current stand structure and the potential MPB impacts in selected stands. Our results indicate that stand-replacing fires initiated even-aged seral lodgepole pine stands in this region. In the absence of fire in the 20th century, multiple MPB disturbances, which each resulted in partial canopy removal, modified the simple one-layer structure of the fire-origin stands by the initiation of post-MPB disturbance regeneration layers, transforming the stands into complex, multi-aged stands. Despite high overstory mortality due to the current MPB outbreak, regeneration layers, which are likely to survive the current outbreak, will provide important ecological legacies and will contribute to mid-term timber supply.

Large‐scale climatic patterns and area affected by mountain pine beetle in British Columbia, Canada

We present evidence of high spatial synchrony in an area affected by mountain pine beetle (MPB, Dendroctonus ponderosae) across large distances in British Columbia, Canada, in a study of a spatially explicit database of an area affected by MPB‐caused tree mortality for the period 1959–2002. We further show that large‐scale climatic patterns (Pacific Decadal Oscillation (PDO) and, to a lesser degree, Arctic Oscillation (AO)) are strongly related to the observed MPB synchrony, and that they probably operate through controlling the frequency of extreme cold winter temperatures that affect MPB larvae survival. A smaller portion of the data's variability is linked to the onset of the two largest outbreaks in the studied period and might be attributed to dispersal from outbreak‐prone areas or else to differences in microhabitat (e.g., host availability) in these regions. The onset of a warm PDO phase in 1976 favored MPB outbreaks by reducing the occurrence of extremely low winter temperatures province‐wide. Likewise, the exceptionally high and persistent AO values of the late 1980s and 1990s enhanced MPB activity in the southern and northern parts of the region. Summer warmth cannot be discarded as an important agent at smaller scales.

Effects of mountain pine beetle ( Dendroctonus ponderosae Hopkins) infestations on forest stand structure in the southern Canadian Rocky Mountains

Forests in the montane and lower subalpine ecoregions of the southern Canadian Rocky Mountains may have been more open and structurally diverse at the beginning of the 20th Century than today. Today, the area of mature Pinus contorta subsp. latifolia Dougl. forest that is conducive to mountain pine beetle (MPB, Dendroctonus ponderosae Hopkins) infestations, infrequent high severity fires, and herbivory appears to have increased in Banff and Kootenay National Parks. Based on a review of the literature, we hypothesised that MPB infestations increase forest stand structural diversity and tree species diversity. Stand structure parameters were investigated in mesic montane and lower subalpine stands approximately 15, 25, and 65 years after MPB outbreaks. Parameters measured were stand density (number of trees per ha), diameter at breast height, height class, species, age class distribution, and coarse woody debris mass. Influences of fire frequency, time since fire, and fire severity on these parameters were assessed to determine whether fire history had a confounding influence on stand structure. The Shannon–Wiener index indicated higher stand structural diversity 15 years but not 25 and 65 years after MPB infestations. MPB infestations led to general decreases within stands in the number of living trees, small diameter snags, and Pinus tree species and an increase within stands in the number of large diameter snags. Management that allows the occurrence of the natural fire regime of variable severity fires, in addition to some MPB infestations, would provide for more open and diverse stands. MPB infestations have some effects on stand structure that are similar to those of fire. Changes in stand structure resulting from recent declines in burning rates within Banff and Kootenay National Parks can be reversed to some extent by MPB infestations.

INFLUENCES OF SECONDARY DISTURBANCES ON LODGEPOLE PINE STAND DEVELOPMENT IN ROCKY MOUNTAIN NATIONAL PARK

Although high-severity fire is the primary type of disturbance shaping the structure of lodgepole pine (Pinus contorta) stands in the southern Rocky Mountains, many post-fire stands are also affected by blowdown, low-severity surface fires, and/or outbreaks of mountain pine beetle (MPB; Dendroctonus ponderosae). The ecological effects of these secondary disturbances are poorly understood but are potentially important in the context of managing for ecological restoration and fire hazard mitigation. We investigated the effects of blowdown, surface fires, and MPB outbreaks on demographic processes in post-fire lodgepole pine stands in Rocky Mountain National Park, Colorado, USA. We used dendroecological methods to reconstruct stand characteristics prior to and following secondary disturbances for paired stands with and without secondary disturbances. Surface fire events do not kill canopy trees or trigger pulses of recruitment and as such do not have detectable influences on stand development. In contrast, both MPB and blowdown kill canopy trees and trigger pulses of tree regeneration of lodgepole pine and subalpine fir (Abies lasiocarpa). The amount and species composition of post-disturbance regeneration is dependent on the severity of the disturbance and on the time since stand initiation. Secondary disturbances of higher severity (i.e., killing >50% of the canopy trees) that occur in younger post-fire stands favor new establishment of lodgepole pine. In contrast, secondary disturbances of lower severity in older stands (>250 years) trigger a pulse of establishment of subalpine fir. The results of this study demonstrate that the high tree densities characteristic of lodgepole pine stands in the southern Rockies (southern Wyoming to northern New Mexico) are the result of dense regeneration following stand-replacing fires and that surface fires had little or no thinning effect on tree densities. Thus, current high stand densities in the study area are not the result of suppression of surface fires. Moreover, the strong pulses of regeneration following forest thinning by MPB and blowdowns imply that, depending on the degree of thinning, thinning prescriptions to reduce fuels in the lodgepole pine forest type may have the unintended consequence of increasing ladder fuels 15-20 years following treatments.

Model Analysis of Spatial Patterns in Mountain Pine Beetle Outbreaks

The mountain pine beetle [MPB,Dendroctonus ponderosaeHopkins (Coleoptera: Scolytidae)] is an aggressive bark beetle, one that typically needs to kill host trees in order to successfully reproduce. This ecological adaptation has resulted in an organism that is both economically important and ecologically significant. Even though significant resources have been expended on MPB research, and a great deal of knowledge exists regarding individual aspects of MPB ecology, some of the most basic questions regarding outbreaks remain unanswered. In our opinion, one reason for the lack of synthesis and predictive power is the inadequate treatment of spatial dynamics in outbreak theories. This paper explicitly addresses the role of spatial dynamics in the precipitation and propagation of MPB outbreaks. We first describe a spatially dynamic model of the MPB/forest interaction that includes chemical ecology, spatial redistribution of beetles, attack, and resulting host mortality. The model is a system of 6 coupled, partial differential equations with 7 state variables and 20 parameters. It represents an attempt to capture the relatively complex predator/prey interaction between MPB and host trees by including the minimum phenomenological descriptions necessary for ecological credibility. This system of equations describes the temporal dynamics of: beetle attraction as a function of pheromone concentration; the change in numbers of flying and nesting beetles; tree resistance/susceptibility; and tree recovery from attack. Spatial dynamics are modeled by fluxes due to gradients in pheromones and kairomones, and the random redistribution of beetles in absence of semiochemicals. We then use the parameterized model to explore three issues central to the ecology of MPB/forest interaction. The first of these is in response to the need for objective ways to compare patterns of successful beetle attacks as they evolve in space. Simulation results indicate that at endemic levels, the pattern of successful attacks are determined almost exclusively by the underlying distribution of susceptible host trees (environmental determinism). As an outbreak develops, the pattern of successfully attacked trees switches to one that is dynamically driven by the self-generated semiochemical landscape (dynamic determinism). This switch from an environmentally determined spatial pattern to a dynamically driven pattern is the hallmark of an outbreak. We discuss the application of a spatial correlation coefficient that can be used to differentiate between the spatial distribution of killed trees in endemic and outbreak phases. The second issue we address through simulation is synchrony in adult emergence. Synchronous adult emergence is critical for the mass attack strategy necessary for overcoming tree defenses. Results from these simulations indicate that the degree of synchrony in adult emergence can have important consequences for assessing the risk of an outbreak. The final issue we investigate through simulation is the effect of spatial pattern of nurse trees (those successfully attacked the previous year) on outbreak potential. Simulations indicated that the spatial proximity of nurse trees was an important determinant of subsequent successful attacks. We conclude with a discussion of the general implications of our simulation experiments.