The Relative Abundance of Mountain Pine Beetle Fungal Associates Through the Beetle Life Cycle in Pine Trees

The mountain pine beetle (MPB) is a native bark beetle of western North America that attacks pine tree species, particularly lodgepole pine. It is closely associated with the ophiostomatoid ascomycetes Grosmannia clavigera, Leptographium longiclavatum, Ophiostoma montium, and Ceratocystiopsis sp.1, with which it is symbiotically associated. To develop a better understanding of interactions between beetles, fungi, and host trees, we used target-specific DNA primers with qPCR to assess the changes in fungal associate abundance over the stages of the MPB life cycle that occur in galleries under the bark of pine trees. Multivariate analysis of covariance identified statistically significant changes in the relative abundance of the fungi over the life cycle of the MPB. Univariate analysis of covariance identified a statistically significant increase in the abundance of Ceratocystiopsis sp.1 through the beetle life cycle, and pair-wise analysis showed that this increase occurs after the larval stage. In contrast, the abundance of O. montium and Leptographium species (G. clavigera, L. longiclavatum) did not change significantly through the MPB life cycle. From these results, the only fungus showing a significant increase in relative abundance has not been formally described and has been largely ignored by other MPB studies. Although our results were from only one site, in previous studies we have shown that the fungi described were all present in at least ten sites in British Columbia. We suggest that the role of Ceratocystiopsis sp.1 in the MPB system should be explored, particularly its potential as a source of nutrients for teneral adults.     

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.     

The Influence of Previous Mountain Pine Beetle (Dendroctonus ponderosae) Activity on the 1988 Yellowstone Fires

We examined the historical record of mountain pine beetle (Dendroctonus ponderosae Hopkins) activity within Yellowstone National Park, Wyoming, for the 25-years period leading up to the 1988 Yellowstone fires (1963–86) to determine how prior beetle activity and the resulting tree mortality affected the spatial pattern of the 1988 Yellowstone fires. To obtain accurate estimates of our model parameters, we used a Markov chain Monte Carlo method to account for the high degree of spatial autocorrelation inherent to forest fires. Our final model included three statistically significant variables: drought, aspect, and sustained mountain pine beetle activity in the period 1972–75. Of the two major mountain pine beetle outbreaks that preceded the 1988 fires, the earlier outbreak (1972–75) was significantly correlated with the burn pattern, whereas the more recent one (1980–83) was not. Although regional drought and high winds were responsible for the large scale of this event, the analysis indicates that mountain pine beetle activity in the mid-1970s increased the odds of burning in 1988 by 11% over unaffected areas. Although relatively small in magnitude, this effect, combined with the effects of aspect and spatial variation in drought, had a dramatic impact on the spatial pattern of burned and unburned areas in 1988.     

The Role of Wildfire,Prescribed Fire,and Mountain Pine Beetle Infestations on the Population Dynamics of Black-Backed Woodpeckers in the Black Hills,South Dakota

Wildfire and mountain pine beetle infestations are naturally occurring disturbances in western North American forests. Black-backed woodpeckers (Picoides arcticus) are emblematic of the role these disturbances play in creating wildlife habitat, since they are strongly associated with recently-killed forests. However, management practices aimed at reducing the economic impact of natural disturbances can result in habitat loss for this species. Although black-backed woodpeckers occupy habitats created by wildfire, prescribed fire, and mountain pine beetle infestations, the relative value of these habitats remains unknown. We studied habitat-specific adult and juvenile survival probabilities and reproductive rates between April 2008 and August 2012 in the Black Hills, South Dakota. We estimated habitat-specific adult and juvenile survival probability with Bayesian multi-state models and habitat-specific reproductive success with Bayesian nest survival models. We calculated asymptotic population growth rates from estimated demographic rates with matrix projection models. Adult and juvenile survival and nest success were highest in habitat created by summer wildfire, intermediate in MPB infestations, and lowest in habitat created by fall prescribed fire. Mean posterior distributions of population growth rates indicated growing populations in habitat created by summer wildfire and declining populations in fall prescribed fire and mountain pine beetle infestations. Our finding that population growth rates were positive only in habitat created by summer wildfire underscores the need to maintain early post-wildfire habitat across the landscape. The lower growth rates in fall prescribed fire and MPB infestations may be attributed to differences in predator communities and food resources relative to summer wildfire.     

Rapid Increases in Forest Understory Diversity and Productivity following a Mountain Pine Beetle (Dendroctonus ponderosae) Outbreak in Pine Forests

The current unprecedented outbreak of mountain pine beetle (Dendroctonus ponderosae) in lodgepole pine (Pinus contorta) forests of western Canada has resulted in a landscape consisting of a mosaic of forest stands at different stages of mortality. Within forest stands, understory communities are the reservoir of the majority of plant species diversity and influence the composition of future forests in response to disturbance. Although changes to stand composition following beetle outbreaks are well documented, information on immediate responses of forest understory plant communities is limited. The objective of this study was to examine the effects of D. ponderosae-induced tree mortality on initial changes in diversity and productivity of understory plant communities. We established a total of 110 1-m² plots across eleven mature lodgepole pine forests to measure changes in understory diversity and productivity as a function of tree mortality and below ground resource availability across multiple years. Overall, understory community diversity and productivity increased across the gradient of increased tree mortality. Richness of herbaceous perennials increased with tree mortality as well as soil moisture and nutrient levels. In contrast, the diversity of woody perennials did not change across the gradient of tree mortality. Understory vegetation, namely herbaceous perennials, showed an immediate response to improved growing conditions caused by increases in tree mortality. How this increased pulse in understory richness and productivity affects future forest trajectories in a novel system is unknown.     

In Vitro Interactions Between Yeasts and Bacteria and the Fungal Symbionts of the Mountain Pine Beetle (Dendroctonus ponderosae)

Multi-trophic interactions between prokaryotes, unicellular eukaryotes, and ecologically intertwined metazoans are presumably common in nature, yet rarely described. The mountain pine beetle, Dendroctonus ponderosae, is associated with two filamentous fungi, Grosmannia clavigera and Ophiostoma montium. Other microbes, including yeasts and bacteria, are also present in the phloem, but it is not known whether they interact with the symbiotic fungi or the host beetle. To test whether such interactions occur, we performed a suite of in vitro assays. Overall, relative yield of O. montium grown with microbes isolated from larval galleries was significantly greater than when the fungus was grown alone. Conversely, the yield of G. clavigera grown with these same microbes was less than or equal to when it was grown alone, suggesting that O. montium, and at least some microbes in larval galleries, have a mutualistic or commensal relationship, while G. clavigera and those same microbes have an antagonistic relationship. A bacterium isolated from phloem not colonized by beetles was found to inhibit growth of both G. clavigera and O. montium and appears to be an antagonist to both fungi. Our results suggest that bacteria and yeasts likely influence the distribution of mycangial fungi in the host tree, which, in turn, may affect the fitness of D. ponderosae.     

长白山白桦林中红松种群动态的研究

山地红松、云杉、冷杉林(简称红松云冷杉林),是山地云冷杉林与红松阔叶林相连接带上的垂直地带性植被。在长白山分布于海拔1100—1400m之间,构成窄狭的山地红松云冷杉林亚带(周以良、李景文,1964)。红松云冷杉林受干扰后,白桦林是最为主要的派生森林类型之一。所以,红松云冷杉林演替过程中。白桦林阶段以及林下红松的更新居于重要的位置。本世纪以来,由于东北山地森林的大规模开发,红松云冷杉林面积急剧减小;而作为红松云冷杉林演替先锋阶段的白桦林比重相     

Effects of Dwarf Mistletoe on Stand Structure of Lodgepole Pine Forests 21-28 Years Post-Mountain Pine Beetle Epidemic in Central Oregon

Lodgepole pine (Pinus contorta) forests are widely distributed throughout North America and are subject to mountain pine beetle (Dendroctonus ponderosae) epidemics, which have caused mortality over millions of hectares of mature trees in recent decades. Mountain pine beetle is known to influence stand structure, and has the ability to impact many forest processes. Dwarf mistletoe (Arceuthobium americanum) also influences stand structure and occurs frequently in post-mountain pine beetle epidemic lodgepole pine forests. Few studies have incorporated both disturbances simultaneously although they co-occur frequently on the landscape. The aim of this study is to investigate the stand structure of lodgepole pine forests 21–28 years after a mountain pine beetle epidemic with varying levels of dwarf mistletoe infection in the Deschutes National Forest in central Oregon. We compared stand density, stand basal area, canopy volume, proportion of the stand in dominant/codominant, intermediate, and suppressed cohorts, average height and average diameter of each cohort, across the range of dwarf mistletoe ratings to address differences in stand structure. We found strong evidence of a decrease in canopy volume, suppressed cohort height, and dominant/codominant cohort diameter with increasing stand-level dwarf mistletoe rating. There was strong evidence that as dwarf mistletoe rating increases, proportion of the stand in the dominant/codominant cohort decreases while proportion of the stand in the suppressed cohort increases. Structural differences associated with variable dwarf mistletoe severity create heterogeneity in this forest type and may have a significant influence on stand productivity and the resistance and resilience of these stands to future biotic and abiotic disturbances. Our findings show that it is imperative to incorporate dwarf mistletoe when studying stand productivity and ecosystem recovery processes in lodgepole pine forests because of its potential to influence stand structure.     

Spatial Distribution of Mountain Pine Beetle Outbreaks in Relation to Climate and Stand Characteristics:A Dendroecological Analysis

Principal components analysis, followed by K-means cluster analysis, was used to detect variations in the timingand magnitude of Pinus contorfa Dougl. ex Loud. growth releases attributed to mountain pine beetle outbreaks in31 stands of central British Columbia. Four major growth release patterns were identified from 1970 to 2000.Variations in the timing of growth releases among clustered stands corresponded well to aerial survey dataindicating the timing of beetle outbreaks in the study area. Redundancy analysis was used to determine howvariations in the timing and magnitude of growth releases attributed to beetle outbreaks changed with variationsin climate or stand conditions over the study area. The first RDA axis, which accounted for 39% of the variations ingrowth patterns among stands, was significantly (P<0.05) correlated with gradients in the percentage of pine instands killed by mountain pine beetle, summer aridity, variation in summer precipitation, distance from initialinfestation site, average pine age, and maximum August temperatures. The second RDA axis explained 6% of thevariations and was significantly correlated with gradients in the beetle climate suitability index, extreme coldmonth temperatures, and site index. Comparisons of growth release patterns with aerial survey data and redun-dancy analyses indicated that dendrochronological techniques are useful for identifying mountain pine beetleoutbreaks in central British Columbia, particularly among stands that had a density high enough to produce agrowth release signal. Provided future studies account for interannual weather fluctuations, identification ofgrowth increases due to stand thinning caused by beetle outbreaks will be useful for reconstructing the history ofbeetle outbreaks over much longer time periods.     

Spatial Distibution of Mountain Pine Beetle Outbreaks in Relation to Climate and Stand Characteristics:A Dendroecological Analysis

Principal components analysis, followed by K-means cluster analysis, was used to detect variations in the timing and magnitude of Pinus contorta Dougl. ex Loud. growth releases attributed to mountain pine beetle outbreaks in 31 stands of central British Columbia. Four major growth release patterns were identified from 1970 to 2000.Variations in the timing of growth releases among clustered stands corresponded well to aerial survey data indicating the timing of beetle outbreaks in the study area. Redundancy analysis was used to determine how variations in the timing and magnitude of growth releases attributed to beetle outbreaks changed with variations in climate or stand conditions over the study area. The first RDA axis, which accounted for 39% of the variations in growth patterns among stands, was significantly (P＜0.05) correlated with gradients in the percentage of pine in stands killed by mountain pine beetle, summer aridity, variation in summer precipitation, distance from initial infestation site, average pine age, and maximum August temperatures. The second RDA axis explained 6% of the variations and was significantly correlated with gradients in the beetle climate suitability index, extreme cold month temperatures, and site index. Comparisons of growth release patterns with aerial survey data and redundancy analyses indicated that dendrochronological techniques are useful for identifying mountain pine beetle outbreaks in central British Columbia, particularly among stands that had a density high enough to produce a growth release signal. Provided future studies account for interannual weather fluctuations, identification of growth increases due to stand thinning caused by beetle outbreaks will be useful for reconstructing the history of beetle outbreaks over much longer time periods.     

Competitive Interactions among Symbiotic Fungi of the Southern Pine Beetle

The southern pine beetle, a damaging pest of conifers, is intimately linked to three symbiotic fungi. Two fungi, Ceratocystiopsis ranaculosus and Entomocorticium sp. A, are transported within specialized structures (mycangia) in the beetle exoskeleton and are mutualists of the beetle. A third fungus, Ophiostoma minus, is transported externally on the beetle exoskeleton (phoretically) and is an antagonist of the beetle. This study examined competitive interactions among these three fungi. The results of de Wit replacement series and primary and secondary resource capture assays with these fungi provide strong evidence for differential competition between the phoretic and mycangial fungi. O. minus was the most able to capture both uncolonized and colonized resources. Entomocorticium sp. A and C. ranaculosus, although equal to one another in competitive abilities, differed in their ability to compete with O. minus. Entomocorticium sp. A was able to maintain space free of O. minus to a much greater degree than was C. ranaculosus. The outcome of such competitive interactions may have significant impacts on the biology of this ecologically and economically important beetle.     

Competition and Coexistence In a Multi-partner Mutualism: Interactions Between two Fungal Symbionts of the Mountain Pine Beetle In Beetle-attacked Trees

Despite overlap in niches, two fungal symbionts of the mountain pine beetle (Dendroctonus ponderosae), Grosmannia clavigera and Ophiostoma montium, appear to coexist with one another and their bark beetle host in the phloem of trees. We sampled the percent of phloem colonized by fungi four times over 1 year to investigate the nature of the interaction between these two fungi and to determine how changing conditions in the tree (e.g., moisture) affect the interaction. Both fungi colonized phloem at similar rates; however, G. clavigera colonized a disproportionately larger amount of phloem than O. montium considering their relative prevalence in the beetle population. High phloem moisture appeared to inhibit fungal growth shortly after beetle attack; however, by 1 year, low phloem moisture likely inhibited fungal growth and survival. There was no inverse relationship between the percent of phloem colonized by G. clavigera only and O. montium only, which would indicate competition between the species. However, the percent of phloem colonized by G. clavigera and O. montium together decreased after 1 year, while the percent of phloem from which no fungi were isolated increased. A reduction in living fungi in the phloem at this time may have significant impacts on both beetles and fungi. These results indicate that exploitation competition occurred after a year when the two fungi colonized the phloem together, but we found no evidence of strong interference competition. Each species also maintained an exclusive area, which may promote coexistence of species with similar resource use.     

A Decade of Streamwater Nitrogen and Forest Dynamics after a Mountain Pine Beetle Outbreak at the Fraser Experimental Forest,Colorado

Forests of western North America are currently experiencing extensive tree mortality from a variety of bark beetle species, and insect outbreaks are projected to increase under warmer, drier climates. Unlike the abrupt biogeochemical changes typical after wildfire and timber harvesting, the outcomes of insect outbreaks are poorly understood. The mountain pine bark beetle (Dendroctonus ponderosae) began to attack lodgepole pine (Pinus contorta) at the Fraser Experimental Forest in 2002 and spread throughout the research area by 2007. We compared streamwater nitrogen (N) from 2003 through 2012 with data from the previous two decades in four watersheds with distinct forest management histories, stand structures, and responses to the beetle outbreak. Watersheds dominated by old-growth had larger trees and lost 85% of overstory pine and 44% of total basal area to bark beetles. In contrast, managed watersheds containing a mixture of second-growth (30–60 year old) and old-growth (250- to 350-year old) had higher density of subcanopy trees, smaller mean tree diameter, and lower bark beetle-induced mortality (~26% of total basal area). Streamwater nitrate concentrations were significantly higher in old-growth watersheds during the outbreak than pre-outbreak levels during snowmelt and base flow seasons. In mixed-age stands, streamwater nitrate concentrations were unaffected by the outbreak. Beetle outbreak elevated inorganic N export 43 and 74% in two old-growth watersheds though the amounts of N released in streamwater were low (0.04 and 0.15 kg N ha^?1) relative to atmospheric inputs (<2% of annual N deposition). Increased height, diameter, and foliar N of measured in residual live trees augmented demand for N, far in excess of the change in N export during the outbreak. Reallocation of soil resources released after pine mortality to overstory and understory vegetation helps explain high nutrient retention in watersheds affected by bark beetle outbreaks.     

灵空山油松种群年龄结构与动态分析

对山西灵空山自然保护区的油松（Pinus tabulaeformis）种群进行了调查,利用生存分析理论及谱分析理论,分析了油松种群径级结构和静态生命表,绘制了存活曲线,同时分析了种群数量的周期性。主要结果如下：（1）油松种群幼苗幼树个体数较多,中径级的个体数较稳定,而高径级由于种间竞争及环境因子的影响,个体数明显下降。（2）油松种群的存活曲线呈现Deevey-Ⅱ型。动态表现为前期增长,中期稳定,后期衰退。（3）油松种群波动存在小周期,但变化不大,说明油松种群发展较为稳定。