Aboveground Net Primary Production and Leaf-Area Index in Early Postfire Vegetation in Yellowstone National Park

Aboveground net primary production (ANPP) and leaf-area index (LAI) of lodgepole pine (Pinus contorta var. latifolia Engelm. ex Wats.) saplings and aboveground productivity of herbaceous vegetation components were determined 9 years after the 1988 fires in Yellowstone National Park (YNP). Measurements were made in four sites representing a wide range of early postfire vegetation present in YNP, including high-density lodgepole pine, low-density lodgepole pine, and two nonforest stands. LAI of the pine saplings and total ANPP (trees plus herbs) generally increased with increasing sapling density, from 0.002 m² m^{− 2} and 0.25 Mg ha^{− 1} year^{− 1} in the infertile nonforest stand (100 pine saplings ha^{− 1}) to 1.8 m² m^{− 2} and 4.01 Mg ha^{− 1} year^{− 1} in the high-density pine stand (62,800 saplings ha^{− 1}). Aboveground herbaceous productivity was not strongly correlated with sapling density, but appeared to be influenced by soil fertility. In the high-density pine stand, tree ANPP and LAI were within the lower range of values reported for similar mature coniferous forests. This finding suggests that at least some ecosystem processes (related to ANPP and LAI) may have nearly recovered after only 9 years of postfire succession, in at least some of the young forests developing after the 1988 Yellowstone fires. Received 7 April 1998; accepted 1 December 1998.     

Spatial–temporal analysis of species range expansion: the case of the mountain pine beetle, Dendroctonus ponderosae

Aim The spatial extent of western Canada’s current epidemic of mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae, Scolytinae), is increasing. The roles of the various dispersal processes acting as drivers of range expansion are poorly understood for most species. The aim of this paper is to characterize the movement patterns of the mountain pine beetle in areas where range expansion is occurring, in order to describe the fine‐scale spatial dynamics of processes associated with mountain pine beetle range expansion. Location Three regions of Canada’s Rocky Mountains: Kicking Horse Pass, Yellowhead Pass and Pine Pass. Methods Data on locations of mountain pine beetle‐attacked trees of predominantly lodgepole pine (Pinus contorta var. latifolia) were obtained from annual fixed‐wing aircraft surveys of forest health and helicopter‐based GPS surveys of mountain pine beetle‐damaged areas in British Columbia and Alberta. The annual (1999–2005) spatial extents of outbreak ranges were delineated from these data. Spatial analysis was conducted using the spatial–temporal analysis of moving polygons (STAMP), a recently developed pattern‐based approach. Results We found that distant dispersal patterns (spot infestations) were most often associated with marginal increases in the areal size of mountain pine beetle range polygons. When the mountain pine beetle range size increased rapidly relative to the years examined, local dispersal patterns (adjacent infestation) were more common. In Pine Pass, long‐range dispersal (> 2 km) markedly extended the north‐east border of the mountain pine beetle range. In Yellowhead Pass and Kicking Horse Pass, the extension of the range occurred incrementally via ground‐based spread. Main conclusions Dispersal of mountain pine beetle varies with geography as well as with host and beetle population dynamics. Although colonization is mediated by habitat connectivity, during periods of low overall habitat expansion, dispersal to new distant locations is common, whereas during periods of rapid invasion, locally connected spread is the dominant mode of dispersal. The propensity for long‐range transport to establish new beetle populations, and thus to be considered a driver of range expansion, is likely to be determined by regional weather patterns, and influenced by local topography. We conclude that STAMP appears to be a useful approach for examining changes in biogeograpical ranges, with the potential to reveal both fine‐ and large‐scale patterns.     

Spatial Community Structure of Mountain Pine Beetle Fungal Symbionts Across a Latitudinal Gradient

Symbiont redundancy in obligate insect–fungal systems is thought to buffer the insect host against symbiont loss and to extend the environmental conditions under which the insect can persist. The mountain pine beetle is associated with at least three well-known and putatively obligate ophiostomatoid fungal symbionts that vary in their environmental tolerances. To better understand the spatial variation in beetle–fungal symbiotic associations, we examined the community composition of ophiostomatoid fungi associated with the mountain pine beetle as a function of latitude and elevation. The region investigated represents the leading edge of a recent outbreak of mountain pine beetle in western Canada. Using regression and principal components analysis, we identified significant spatial patterns in fungal species abundances that indicate symmetrical replacement between two of the three fungi along a latitudinal gradient and little variation in response to elevation. We also identified significant variation in the prevalence of pair-wise species combinations that occur within beetle galleries. Frequencies of pair-wise combinations were significantly different from what was expected given overall species abundances. These results suggest that complex processes of competitive exclusion and coexistence help determine fungal community composition and that the consequences of these processes vary spatially. The presence of three fungal symbionts in different proportions and combinations across a wide range of environmental conditions may help explain the success of mountain pine beetle attacks across a broad geographic range.     

Landscape Patterns and Legacies Resulting from Large, Infrequent Forest Disturbances

We review and compare well-studied examples of five large, infrequent disturbances (LIDs)—fire, hurricanes, tornadoes, volcanic eruptions, and floods—in terms of the physical processes involved, the damage patterns they create in forested landscapes, and the potential impacts of those patterns on subsequent forest development. Our examples include the 1988 Yellowstone fires, the 1938 New England hurricane, the 1985 Tionesta tornado, the 1980 eruption of Mount St. Helens, and the 1993 Mississippi floods. The resulting landscape patterns are strongly controlled by interactions between the specific disturbance, the abiotic environment (especially topography), and the composition and structure of the vegetation at the time of the disturbance. The very different natures of these interactions yield distinctive temporal and spatial patterns and demand that ecologists increase their knowledge of the physical characteristics of disturbance processes. Floods and fires can occur over a long period, whereas volcanic eruptions and wind-driven events often last for no more than a few hours or days. Tornadoes and floods produce linear patterns with sharp edges, but fires, volcanic eruptions, and hurricanes can affect broader areas, often with gradual transitions of disturbance intensity. In all cases, the evidence suggests that LIDs produce enduring legacies of physical and biological structure that influence ecosystem processes for decades or centuries. Received 14 July 1998; accepted 6 October 1998.     

Twenty Years After the 1988 Yellowstone Fires: Lessons About Disturbance and Ecosystems

The 1988 Yellowstone fires were among the first in what has proven to be an upsurge in large severe fires in the western USA during the past 20 years. At the time of the fires, little was known about the impacts of such a large severe disturbance because scientists had had few previous opportunities to study such an event. Ecologists predicted short- and long-term effects of the 1988 fires on vegetation, biogeochemistry, primary productivity, wildlife, and aquatic ecosystems based on scientific understanding of the time. Twenty-plus years of subsequent study allow these early predictions to be evaluated. Most of the original predictions were at least partially supported, but some predictions were refuted, others nuanced, and a few postfire phenomena were entirely unexpected. Post-1988 Yellowstone studies catalyzed advances in ecology focused on the importance of spatial and temporal heterogeneity, contingent influences, and multiple interacting drivers. Post-1988 research in Yellowstone also has changed public perceptions of fire as an ecological process and attitudes towards fire management. Looking ahead to projected climate change and more frequent large fires, the well-documented ecological responses to the 1988 Yellowstone fires provide a foundation for detecting and evaluating potential changes in fire regimes of temperate mountainous regions.     

Lodgepole pine forest age class dynamics and susceptibility to mountain pine beetle attack

Because mountain pine beetle attack mature pine stands, an understanding of forest age class dynamics is important to managing forests within the distribution of the beetle. The assumed theoretical negative exponential forest age distribution provides an estimate when ecosystem dynamics are in equilibrium. This study investigates the dynamics of forest age distribution for non-equilibrium ecosystem dynamics, which result primarily from large and irregular stand-replacement fire disturbances that alter the forest age distribution. A model experiment using the SEM-LAND model on a 1 million ha lodgepole pine forest landscape was conducted to estimate how the proportion of susceptible area could be influenced by different fire regimes. The results of the simulation suggest that the temporal dynamics of the area susceptible to mountain pine beetle attack are complex and depend on the fire history of the study area, if the area is experiencing large and irregular stand-replacement fires. The age range of the lodgepole pine forest stands susceptible to mountain pine beetle attack might significantly affect the estimate of the area susceptible to attack.     

Drought effects on above- and belowground production of a grazed temperate grassland ecosystem

The effect of climatic variation on terrestrial aboveground productivity (ANPP) has been well studied. However, little is known about how variable climate, including drought, may influence belowground productivity (BNPP), which constitutes most of the annual primary production of grasslands. The objectives of this study were to (1) examine how a 3-year period of declining moisture, which began as climatically wet to average across Yellowstone National Park (YNP) and ended in drought, affected ANPP and BNPP in grasslands of YNP and (2) how herds of grazing ungulates, which were shown previously to stimulate grassland shoot and root growth in YNP, may have interacted with climatic conditions to influence grassland production. ANPP and 0–20 cm BNPP, representing the bulk of the root dynamics, were measured in grazed and ungrazed (fenced) grassland at nine sites ranging widely in elevation, soil conditions and plant production during the 3-year study. Results revealed that 0–20 cm BNPP was strongly influenced by drought (P = 0.0005) and declined from 1999 to 2001 among ungrazed and grazed grasslands by 39 and 49%, respectively. The greater reduction in 0–20 cm BNPP among grazed grasslands was due, in part, to a decline (P = 0.07) in the stimulatory effect of grazing, i.e., the ratio g BNPP stimulated: g shoot consumed. In contrast, ANPP was unaffected by drought in either type of grassland. Thus, the effect of this drought in YNP was a large reduction in BNPP, which was a function of (1) a direct negative influence of increased moisture stress on root growth and (2) a weak interaction between drought and herbivory that led to a decline in the positive feedback from grazers to BNPP. These findings highlight the need to better understand factors that control root growth and to study the effects of climatic variation on grasslands within an ecosystem framework to include potentially important climate–consumer interactions.     

The effect of warmer winters on the demography of an outbreak insect is hidden by intraspecific competition

Warmer climates are predicted to increase bark beetle outbreak frequency, severity, and range. Even in favorable climates, however, outbreaks can decelerate due to resource limitation, which necessitates the inclusion of competition for limited resources in analyses of climatic effects on populations. We evaluated several hypotheses of how climate impacts mountain pine beetle reproduction using an extensive 9‐year dataset, in which nearly 10,000 trees were sampled across a region of approximately 90,000 km², that was recently invaded by the mountain pine beetle in Alberta, Canada. Our analysis supports the hypothesis of a positive effect of warmer winter temperatures on mountain pine beetle overwinter survival and provides evidence that the increasing trend in minimum winter temperatures over time in North America is an important driver of increased mountain pine beetle reproduction across the region. Although we demonstrate a consistent effect of warmer minimum winter temperatures on mountain pine beetle reproductive rates that is evident at the landscape and regional scales, this effect is overwhelmed by the effect of competition for resources within trees at the site level. Our results suggest that detection of the effects of a warming climate on bark beetle populations at small spatial scales may be difficult without accounting for negative density dependence due to competition for resources.     

What explains landscape patterns of tree mortality caused by bark beetle outbreaks in Greater Yellowstone?

Aim Bark beetle outbreaks have recently affected extensive areas of western North American forests, and factors explaining landscape patterns of tree mortality are poorly understood. The objective of this study was to determine the relative importance of stand structure, topography, soil characteristics, landscape context (the characteristics of the landscape surrounding the focal stand) and beetle pressure (the abundance of local beetle population eruptions around the focal stand a few years before the outbreak) to explain landscape patterns of tree mortality during outbreaks of three species: the mountain pine beetle, which attacks lodgepole pine and whitebark pine; the spruce beetle, which feeds on Engelmann spruce; and the Douglas‐fir beetle, which attacks Douglas‐fir. A second objective was to identify common variables that explain tree mortality among beetle–tree host pairings during outbreaks. Location Greater Yellowstone ecosystem, Wyoming, USA. Methods We used field surveys to quantify stand structure, soil characteristics and topography at the plot level in susceptible stands of each forest type showing different severities of infestation (0–98% mortality; n= 129 plots). We then used forest cover and beetle infestation maps derived from remote sensing to develop landscape context and beetle pressure metrics at different spatial scales. Plot‐level and landscape‐level variables were used to explain outbreak severity. Results Engelmann spruce and Douglas‐fir mortality were best predicted using landscape‐level variables alone. Lodgepole pine mortality was best predicted by both landscape‐level and plot‐level variables. Whitebark pine mortality was best – although poorly – predicted by plot‐level variables. Models including landscape context and beetle pressure were much better at predicting outbreak severity than models that only included plot‐level measures, except for whitebark pine. Main conclusions Landscape‐level variables, particularly beetle pressure, were the most consistent predictors of subsequent outbreak severity within susceptible stands of all four host species. These results may help forest managers identify vulnerable locations during ongoing outbreaks.     

Influence of bioclimatic variables on tree-line conifer distribution in the Greater Yellowstone Ecosystem: implications for species of conservation concern

Aim Tree‐line conifers are believed to be limited by temperature worldwide, and thus may serve as important indicators of climate change. The purpose of this study was to examine the potential shifts in spatial distribution of three tree‐line conifer species in the Greater Yellowstone Ecosystem under three future climate‐change scenarios and to assess their potential sensitivity to changes in both temperature and precipitation. Location This study was performed using data from 275 sites within the boundaries of Yellowstone and Grand Teton national parks, primarily located in Wyoming, USA. Methods We used data on tree‐line conifer presence from the US Forest Service Forest Inventory and Analysis Program. Climatic and edaphic variables were derived from spatially interpolated maps and approximated for each of the sites. We used the random‐forest prediction method to build a model of predicted current and future distributions of each of the species under various climate‐change scenarios. Results We had good success in predicting the distribution of tree‐line conifer species currently and under future climate scenarios. Temperature and temperature‐related variables appeared to be most influential in the distribution of whitebark pine (Pinus albicaulis), whereas precipitation and soil variables dominated the models for subalpine fir (Abies lasiocarpa) and Engelmann spruce (Picea engelmannii). The model for whitebark pine substantially overpredicted absences (as compared with the other models), which is probably a result of the importance of biological factors in the distribution of this species. Main conclusions These models demonstrate the complex response of conifer distributions to changing climate scenarios. Whitebark pine is considered a ‘keystone’ species in the subalpine forests of western North America; however, it is believed to be nearly extinct throughout a substantial portion of its range owing to the combined effects of an introduced pathogen, outbreaks of the native mountain pine beetle (Dendroctonus ponderosae), and changing fire regimes. Given predicted changes in climate, it is reasonable to predict an overall decrease in pine‐dominated subalpine forests in the Greater Yellowstone Ecosystem. In order to manage these forests effectively with respect to future climate, it may be important to focus attention on monitoring dry mid‐ and high‐elevation forests as harbingers of long‐term change.     

Successful colonization, reproduction, and new generation emergence in live interior hybrid spruce Picea engelmannii×glauca by mountain pine beetle Dendroctonus ponderosae

1 Although mountain pine beetle Dendroctonus ponderosae Hopkins are able to utilize most available Pinus spp. as hosts, successful colonization and reproduction in other hosts within the Pinaceae is rare.
2 We observed successful reproduction of mountain pine beetle and emergence of new generation adults from interior hybrid spruce Picea engelmannii × glauca and compared a number of parameters related to colonization and reproductive success in spruce with nearby lodgepole pine Pinus contorta infested by mountain pine beetle.
3 The results obtained indicate that reduced competition in spruce allowed mountain pine beetle parents that survived the colonization process to produce more offspring per pair than in more heavily-infested nearby pine.
4 We also conducted an experiment in which 20 spruce and 20 lodgepole pines were baited with the aggregation pheromone of mountain pine beetle. Nineteen pines (95%) and eight spruce (40%) were attacked by mountain pine beetle, with eight (40%) and three (15%) mass-attacked, respectively.
5 Successful attacks on nonhost trees during extreme epidemics may be one mechanism by which host shifts and subsequent speciation events have occurred in Dendroctonus spp. bark beetles.     

Landscape level analysis of mountain pine beetle in British Columbia, Canada: spatiotemporal development and spatial synchrony within the present outbreak

Eruptive herbivores can exert profound landscape level influences. For example, the ongoing mountain pine beetle outbreak in British Columbia, Canada, has resulted in mortality of mature lodgepole pine over >7 million ha. Analysis of the spatio‐temporal pattern of spread can lend insights into the processes initiating and/or sustaining such phenomena. We present a landscape level analysis of the development of the current outbreak. Aerial survey assessments of tree mortality, projected onto discrete 12×12 km cells, were used as a proxy for insect population density. We examined whether the outbreak potentially originated from an epicenter and spread, or whether multiple localized populations erupted simultaneously at spatially disjunct locations. An aspatial cluster analysis of time series from 1990 to 2003 revealed four distinct time series patterns. Each time series demonstrated a general progression of increasing mountain pine beetle populations. Plotting the geographical locations of each temporal pattern revealed that the outbreak occurred first in an area of west‐central British Columbia, and then in an area to the east. The plot further revealed many localized infestations erupted in geographically disjunct areas, especially in the southern portion of the province. Autologistic regression analyses indicated a significant, positive association between areas where the outbreak first occurred and conservation lands. For example, the delineated area of west‐central British Columbia is comprised of three conservation parks and adjacent working forest. We further examined how population synchrony declines with distance at different population levels. Examination of the spatial dependence of temporal synchrony in population fluctuations during early, incipient years (i.e. 1990–1996) suggested that outbreaking mountain pine beetle populations are largely independent at scales >200 km during non‐epidemic periods. However, during epidemic years (i.e. 1999–2003), populations were clearly synchronous across the entire province, even at distances of up to 900 km. The epicentral pattern of population development can be used to identify and prioritize adjacent landscape units for both reactive and proactive management strategies intended to minimize mountain pine beetle impacts.     

The impact of phloem nutrients on overwintering mountain pine beetles and their fungal symbionts

In the low nutrient environment of conifer bark, subcortical beetles often carry symbiotic fungi that concentrate nutrients in host tissues. Although bark beetles are known to benefit from these symbioses, whether this is because they survive better in nutrient-rich phloem is unknown. After manipulating phloem nutrition by fertilizing lodgepole pine trees (Pinus contorta Douglas var. latifolia), we found bolts from fertilized trees to contain more living individuals, and especially more pupae and teneral adults than bolts from unfertilized trees at our southern site. At our northern site, we found that a larger proportion of mountain pine beetle (Dendroctonus ponderosae Hopkins) larvae built pupal chambers in bolts from fertilized trees than in bolts from unfertilized trees. The symbiotic fungi of the mountain pine beetle also responded to fertilization. Two mutualistic fungi of bark beetles, Grosmannia clavigera (Rob.-Jeffr. & R. W. Davidson) Zipfel, Z. W. de Beer, & M. J. Wingf. and Leptographium longiclavatum Lee, S., J. J. Kim, & C. Breuil, doubled the nitrogen concentrations near the point of infection in the phloem of fertilized trees. These fungi were less capable of concentrating nitrogen in unfertilized trees. Thus, the fungal symbionts of mountain pine beetle enhance phloem nutrition and likely mediate the beneficial effects of fertilization on the survival and development of mountain pine beetle larvae.     

Effects of fire on landscape heterogeneity in Yellowstone National Park,Wyoming

Abstract. A map of burn severity resulting from the 1988 fires that occurred in Yellowstone National Park (YNP) was derived from Landsat Thematic Mapper (TM) imagery and used to assess the isolation of burned areas, the heterogeneity that resulted from fires burning under moderate and severe burning conditions, and the relationship between heterogeneity and fire size. The majority of severely burned areas were within close proximity (50 to 200 m) to unburned or lightly burned areas, suggesting that few burned sites are very far from potential sources of propagules for plant reestablishment. Fires that occurred under moderate burning conditions early during the 1988 fire season resulted in a lower proportion of crown fire than fires that occurred under severe burning conditions later in the season. Increased dominance and contagion of burn severity classes and a decrease in the edge: area ratio for later fires indicated a slightly more aggregated burn pattern compared to early fires. The proportion of burned area in different burn severity classes varied as a function of daily fire size. When daily area burned was relatively low, the proportion of burned area in each burn severity class varied widely. When daily burned area exceeded 1250 ha, the burned area contained about 50 % crown fire, 30 % severe surface burn, and 20 % light surface burn. Understanding the effect of fire on landscape heterogeneity is important because the kinds, amounts, and spatial distribution of burned and unburned areas may influence the reestablishment of plant species on burned sites.     

A spatiotemporal pattern analysis of historical mountain pine beetle outbreaks in British Columbia,Canada

This study documents the spatiotemporal patterns of mountain pine beetle infestations by applying a novel approach based on a landscape infestation dynamics conceptual model in combination with morphological spatial pattern analysis using the mountain pine beetle infested pine mortality data (1960–2010) collected by the annual British Columbia aerial overview survey. The pattern analysis at the provincial level reveals that the 1980s outbreak did not crash as originally thought. The current outbreak is most likely a result of the progressive buildup of the epidemic infestations during the transition period (1985–1995) under favourable weather conditions and substantially improved host resources. This is also true for the Northeast and Cariboo areas of the province specifically, even though the infestations in the Cariboo area remained at incipient‐epidemic levels during the transition period after the 1980s outbreak crashed in 1985. In the Southeast area, the current outbreak apparently continued from the outbreak that initiated in the late 1970s and early 1980s. The 1980s outbreak originated from multiple spatially separate locations whereas the current outbreak initiated from a single location in Tweedsmuir Provincial Park. The centralized and self‐amplifying buildup of the current outbreak implicates at least three substantial expansions that occurred in 2002, 2006, and 2008. This study suggests that at the provincial level, as well as for the Northeast and Southeast areas of the province, the current outbreak is declining but most likely will continue for many years given the ongoing and future warming climate and a large proportion of pines that remain in the habitats of mountain pine beetles. This study also suggests that dispersals, particularly long‐distance dispersal, may play a key role in driving the spread and expansion of the current outbreak although uncertainty remains due to the local dynamics of the beetle populations.     

Variability in Leaf Area and Stemwood Increment Along a 300-year Lodgepole Pine Chronosequence

Large disturbances such as the 1988 Yellowstone fires produce considerable spatial heterogeneity in ecosystem processes across landscapes, in part by affecting vegetation structure. However, the persistence of this heterogeneity with time since disturbance, and thus the role of large disturbances in shaping the heterogeneity of ecosystem processes over large spatial and temporal scales, remains unclear. Such an inquiry requires that variability as well as mean conditions of forest structure and growth be examined if changes are to be projected for heterogeneous postdisturbance landscapes. We studied a chronosequence of unburned, mature lodgepole pine stands (stand ages ranging from 50 to 300 or more years) to examine the variability in stand density, leaf-area index (LAI), and stem growth [basal area increment (BAI), a surrogate for aboveground net primary productivity (ANPP)] with stand age, the relationships between these factors, and how these factors were related to stand and site characteristics. Variation in LAI and BAI was explained primarily by differences in stand density and age (r²=0.51 for both LAI and BAI), and both LAI and BAI were most variable in the youngest age class [coefficient of variation (CV), 38% and 41% for LAI and BAI]. The relationship between LAI or BAI and stand density was significantly weaker (r² < 0.20) at stand ages characterized by canopy closure (50–175 years), suggesting that stand structure and production are closely linked. Thus, the spatial variability of stand production, which is initially very high following large fires in this landscape, is detectable for over a century before successional changes in forest structure greatly affect the initial postdisturbance landscape pattern of stand production. Given the recent focus on spatial heterogeneity of ecosystem processes across large landscapes, projecting changes in postdisturbance patterns of stand production has very strong significance for ecosystem science.     

Effect of Climate on Wildfire Size: A Cross-Scale Analysis

Theory predicts that wildfires will encounter spatial thresholds where different drivers may become the dominant influence on continued fire spread. Studying these thresholds, however, is limited by a lack of sufficiently detailed data sets. To address this problem, we searched for scale thresholds in data describing wildfire size at the Avon Park Air Force Range, south-central Florida. We used power-law statistics to describe the “heavy-tail” of the fire size distribution, and quantile regression to determine how the edges of data distributions of fire size were related to climate. Power-law statistics revealed a heavy-tail, a pattern consistent with scale threshold theory, which predicts that large fires will be rare because only fires that cross all thresholds will become large. Results from quantile regression suggested that different climate conditions served as critical thresholds, influencing wildfire size at different spatial scales. Modeling at higher quantiles (≥75th) implicated drought as driving the spread of larger fires, whereas modeling at lower quantiles (≤25th) implicated that wind governed the spread of smaller fires. Fires of intermediate size were negatively associated with relative humidity. Our results are consistent with the idea that fire spread involves scale thresholds, with the small-scale drivers allowing fires to spread after ignition, but with further spread only being possible when large-scale drivers are favorable. These results suggest that other data sets that have heavy-tailed distributions may contain patterns generated by scale thresholds, and that these patterns may be revealed using quantile regression.     

Hiding Without Cover? Defining Elk Security in a Beetle-Killed Forest

Recent mountain pine beetle (Dendroctonus ponderosae; pine beetle) outbreaks in the western United States have affected nearly 18 million ha of pine (Pinus spp.) forest and are unprecedented in spatial extent, severity, and duration, yet little is known about wildlife responses to large-scale insect outbreaks. Elk (Cervus canadensis) are important wildlife whose dominant management paradigm on public lands has focused on providing security habitat to increase survival during hunting seasons and to maintain elk presence on public lands to promote hunter opportunity. To assess the effect of pine beetles and associated changes in forest structure on elk security, we used a time series to characterize canopy cover pre- and post-pine beetle outbreak, characterized relative canopy cover among the dominant forest types in the study area post-pine beetle outbreak, and used global positioning system location data from male and female elk to define habitat relationships and security during the archery and rifle hunting seasons. Our study area was within the Elkhorn Mountains of southwest Montana, USA, 2015–2017, which experienced 80% mortality of lodgepole pine (Pinus contorta) forests during a pine beetle outbreak that peaked in 2008. We observed an 8.5% reduction in canopy cover within pine beetle-infested lodgepole pine forests, yet canopy cover remained relatively high among other forest types post-outbreak. The top-ranked habitat security models contained positive relationships with canopy cover, distance to motorized routes, terrain ruggedness, and slope with few notable differences among sexes and seasons. Across sexes and seasons, 75% and 50% of elk use was within areas with average canopy cover values ≥31 ± 6.65 (SD)% and ≥53 ± 5.7% that were an average of ≥2,072 ± 187.93 m and ≥3,496 ± 157.32 m from a motorized route, respectively. Therefore, we recommend fall elk security be defined as areas that meet these criteria for minimum canopy cover and distance from motorized routes in the Elkhorn Mountains and in other landscapes with similar forest characteristics and hunting pressures. Although we observed expected reductions in canopy cover within pine beetle-infested forests, defoliation alone did not appear to negatively affect elk security or reduce canopy cover below our management recommendations. Nonetheless, because of the prevalence of standing dead trees in our study area, we recommend future work that investigates the relationships with pine beetle-infested areas post-blowdown because changes in ground structure and costs of locomotion may affect elk habitat and security. © 2019 The Authors. Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

Coarse Woody Debris following Fire and Logging in Wyoming Lodgepole Pine Forests

The accumulation and decomposition of coarse woody debris (CWD) are processes that affect habitat, soil structure and organic matter inputs, and energy and nutrient flows in forest ecosystems. Natural disturbances such as fires typically produce large quantities of CWD as trees fall and break, whereas human disturbances such as timber harvesting remove much of the CWD. Our objective was to compare the amount of CWD removed and left behind after clear-cutting to the amount consumed and left behind after natural fires in Rocky Mountain lodgepole pine. The masses of fallen logs, dead-standing trees, stumps, and root crowns more than 7.5 cm in diameter were estimated in clear-cut and intact lodgepole pine forests in Wyoming and compared to estimates made in burned and unburned stands in Yellowstone National Park (YNP), where no timber harvesting has occurred. Estimates of downed CWD consumed or converted to charcoal during an intense crown fire were also made in YNP. No significant differences in biomass of downed CWD more than 7.5 cm in diameter were detected between burned stands and those following a single clear-cut. However, the total mass of downed CWD plus the mass of snags that will become CWD was nearly twice as high in burned stands than in clear-cuts. In YNP, approximately 8% of the downed CWD was consumed by fire and an additional 8% was converted to charcoal, for an estimated loss of about 16%. In contrast, approximately four times more wood (70%) was removed by clear-cutting. Considering all CWD more than 7.5 cm in diameter that was either still present in the stand or removed by harvesting, slash treatment, or burning, clear-cut stands lost an average of 80 Mg ha⁻¹ whereas stands that burned gained an average of 95 Mg ha⁻¹. Some CWD remains as slash and stumps left behind after harvesting, but stands subjected to repeated harvesting will have forest floor and surface soil characteristics that are beyond the historic range of variability of naturally developing stands. Received 16 November 1999; Accepted 31 May 2000.     

Breach of the northern Rocky Mountain geoclimatic barrier: initiation of range expansion by the mountain pine beetle

Aim Our aim is to examine the historical breach of the geoclimatic barrier of the Rocky Mountains by the mountain pine beetle (Dendroctonus ponderosae Hopkins). This recent range expansion from west of the North American continental divide into the eastern boreal forest threatens to provide a conduit to naïve pine hosts in eastern North America. We examine the initial expansion events and determine potential mechanism(s) of spread by comparing spread patterns in consecutive years to various dispersal hypotheses such as: (1) meso‐scale atmospheric dispersal of insects from source populations south‐west of the Rocky Mountains in British Columbia (i.e. their historical range), (2) anthropogenic transport of infested plant material, and (3) spread of insect populations across adjacent stands via corridors of suitable habitat. Location British Columbia, Canada. Methods We explore potential mechanism(s) of invasion of the mountain pine beetle using spatial point process models for the initial 3 years of landscape‐level data collection, 2004–2006. Specifically, we examine observed patterns of infestation relative to covariates reflecting various dispersal hypotheses. We select the most parsimonious models for each of the initial 3 years of invasion using information criteria statistics. Results The initial range expansion and invasion of the beetle was characterized by aerial deposition along a strong north‐west to south‐east gradient, with additional aerial deposition and localized dispersal from persisting populations in following years. Main conclusions Following deposition of a wave front of mountain pine beetles parallel to the Rocky Mountains via meso‐scale atmospheric dispersal, the areas of highest intensity of infestations advanced up to 25 km north‐east towards jack pine (Pinus banksiana) habitat in a single year. There appeared to be no association between putative anthropogenic movement of infested materials and initial range expansion of the mountain pine beetle across the continental divide.