exo-Brevicomin biosynthetic pathway enzymes from the Mountain Pine Beetle,Dendroctonus ponderosae

exoBrevicomin (exo-7-ethyl-5-methyl-6,8-dioxabicyclo[3.2.1]octane) is an important semiochemical for a number of beetle species, including the highly destructive Mountain Pine Beetle (Dendroctonus ponderosae). It is also found in other insects and the African elephant. Despite its significance, very little is known about its biosynthesis. A recent microarray analysis implicated a small cluster of three D. ponderosae genes in exo-brevicomin biosynthesis, two of which had identifiable open reading frames (Aw et al., 2010; BMC Genomics 11:215). Here we report further expression profiling of two genes in that cluster and functional analysis of their recombinantly-produced enzymes. One encodes a short-chain dehydrogenase that used NAD(P)⁺ as a co-factor to catalyze the oxidation of (Z)-6-nonen-2-ol to (Z)-6-nonen-2-one. We therefore named the enzyme (Z)-6-nonen-2-ol dehydrogenase (ZnoDH). The other encodes the cytochrome P450, CYP6CR1, which epoxidized (Z)-6-nonen-2-one to 6,7-epoxynonan-2-one with very high specificity and substrate selectivity. Both the substrates and products of the two enzymes are intermediates in the exo-brevicomin biosynthetic pathway. Thus, ZnoDH and CYP6CR1 are enzymes that apparently catalyze the antepenultimate and penultimate steps in the exo-brevicomin biosynthetic pathway, respectively.     

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.     

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.     

Proteomics Indicators of the Rapidly Shifting Physiology from Whole Mountain Pine Beetle,Dendroctonus ponderosae (Coleoptera: Curculionidae), Adults during Early Host Colonization

We developed proteome profiles for host colonizing mountain pine beetle adults, Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae). Adult insects were fed in pairs on fresh host lodgepole pine, Pinus contorta Dougl. ex Loud, phloem tissue. The proteomes of fed individuals were monitored using iTRAQ and compared to those of starved beetles, revealing 757 and 739 expressed proteins in females and males, respectively, for which quantitative information was obtained. Overall functional category distributions were similar for males and females, with the majority of proteins falling under carbohydrate metabolism (glycolysis, gluconeogenesis, citric acid cycle), structure (cuticle, muscle, cytoskeleton), and protein and amino acid metabolism. Females had 23 proteins with levels that changed significantly with feeding (p<0.05, FDR<0.20), including chaperones and enzymes required for vitellogenesis. In males, levels of 29 proteins changed significantly with feeding (p<0.05, FDR<0.20), including chaperones as well as motor proteins. Only two proteins, both chaperones, exhibited a significant change in both females and males with feeding. Proteins with differential accumulation patterns in females exhibited higher fold changes with feeding than did those in males. This difference may be due to major and rapid physiological changes occurring in females upon finding a host tree during the physiological shift from dispersal to reproduction. The significant accumulation of chaperone proteins, a cytochrome P450, and a glutathione S-transferase, indicate secondary metabolite-induced stress physiology related to chemical detoxification during early host colonization. The females'' activation of vitellogenin only after encountering a host indicates deliberate partitioning of resources and a balancing of the needs of dispersal and reproduction.     

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.     

Determinants of Lichen Diversity in a Rain Forest Understory

Change in lichen diversity is often used as a bioindicator to estimate effects of atmospheric pollution, but natural variation in lichen cover and species richness can be very high. We examined the top-down effects of spore-consuming ants and the bottom-up effects of nutrient and light availability on lichen diversity associated with the leaf surface of the rain forest understory plant, Piper cenocladum. Plots containing P. cenocladum were randomly assigned to treatments in factorial experiments that included high and low light levels, nutrient enrichment, and presence and absence of the ant mutualist, Pheidole bicornis . At the conclusion of the experiments, plants were harvested and size of leaves, secondary metabolite content (amides), epiphyll cover, and the species richness of the lichens (which comprised 85% of the epiphyll community) were quantified. Epiphyll cover (mosses, liverworts, and lichens) was greater on plants that had ant-mutualists and balanced resources. Lichen species richness was greater for plants with balanced resources, particularly for those with high light availability. Relationships between toxins and lichen cover and richness were weak and unclear. In this system, natural sources of variation were reliable determinants of lichen diversity and both biotic and abiotic influences were important.     

Nitrogen Cycling Responses to Mountain Pine Beetle Disturbance in a High Elevation Whitebark Pine Ecosystem

Ecological disturbances can significantly affect biogeochemical cycles in terrestrial ecosystems, but the biogeochemical consequences of the extensive mountain pine beetle outbreak in high elevation whitebark pine (WbP) (Pinus albicaulis) ecosystems of western North America have not been previously investigated. Mountain pine beetle attack has driven widespread WbP mortality, which could drive shifts in both the pools and fluxes of nitrogen (N) within these ecosystems. Because N availability can limit forest regrowth, understanding how beetle-induced mortality affects N cycling in WbP stands may be critical to understanding the trajectory of ecosystem recovery. Thus, we measured above- and belowground N pools and fluxes for trees representing three different times since beetle attack, including unattacked trees. Litterfall N inputs were more than ten times higher under recently attacked trees compared to unattacked trees. Soil inorganic N concentrations also increased following beetle attack, potentially driven by a more than two-fold increase in ammonium (NH₄⁺) concentrations in the surface soil organic horizon. However, there were no significant differences in mineral soil inorganic N or soil microbial biomass N concentrations between attacked and unattacked trees, implying that short-term changes in N cycling in response to the initial stages of WbP attack were restricted to the organic horizon. Our results suggest that while mountain pine beetle attack drives a pulse of N from the canopy to the forest floor, changes in litterfall quality and quantity do not have profound effects on soil biogeochemical cycling, at least in the short-term. However, continuous observation of these important ecosystems will be crucial to determining the long-term biogeochemical effects of mountain pine beetle outbreaks.     

小兴安岭阔叶红松林不同演替系列森林细根生物量的研究

以小兴安岭地区阔叶红松林旱生演替系列的蒙古栎次生林和蒙古栎红松林、中生演替系列的枫桦次生林和枫桦红松林、湿生演替系列的白桦次生林和云冷杉红松林这6种林型为研究对象,采用土钻取样法对不同演替系列细根生物量的空间变化、垂直分布规律及其影响因子进行系统地研究。结果表明：（1）3种演替系列总细根生物量存在显著差异（P<0.05）。中生演替系列生物量最高,平均达8.56 t·hm^-2,其次是旱生演替系列8.18 t·hm^-2,湿生演替系列生物量最低,为5.79 t·hm^-2;（2）对于不同演替系列细根生物量垂直变化,在阔叶红松林的正向演替过程中,森林细根生物量逐渐增加,且随着土层的增加而降低,拟合效果均达显著水平（P<0.05）,中根和粗根的垂直分布差异较大;（3）在小兴安岭地区,6块林地不同径级细根生物量与土壤有机C和全N、土壤容重、土壤含水率呈显著负相关（P>0.05）,而土壤pH值与细根生物量呈显著正相关性（P>0.05）。     

Cryptic postzygotic isolation in an eruptive species of bark beetle (Dendroctonus ponderosae)

Studies of postzygotic isolation often involve well-differentiated taxa that show a consistent level of incompatibility, thereby limiting our understanding of the initial stages and development of reproductive barriers. Dendroctonus ponderosae provides an informative system because recent evidence suggests that distant populations produce hybrids with reproductive incompatibilities. Dendroctonus ponderosae shows an isolation-by-distance gene flow pattern allowing us to characterize the evolution of postzygotic isolation (e.g., hybrid inviability, hybrid sterility) by crossing populations along a continuum of geographic/genetic divergence. We found little evidence of hybrid inviability among these crosses. However, crosses between geographically distant populations produced sterile males (consistent with Haldane's rule). This effect was not consistent with the fixation of mutations in an isolation-by-distance pattern, but instead is spatially localized. These reproductive barriers are uncorrelated with a reduction in gene flow suggesting their recent development. Crosses between geographically proximal populations bounding the transition from compatibility to hybrid male sterility showed evidence of unidirectional reduction in hybrid male fecundity. Our study describes significant postzygotic isolation occurring across a narrow and molecularly cryptic geographic zone between the states of Oregon and Idaho. This study provides a view of the early stages of postzygotic isolation in a geographically widespread species.     

Landscape-scale genetic variation in a forest outbreak species, the mountain pine beetle (Dendroctonus ponderosae)

The mountain pine beetle Dendroctonus ponderosae is a native species currently experiencing large-scale outbreaks in western North American pine forests. We sought to describe the pattern of genetic variation across the range of this species, to determine whether there were detectable genetic differences between D. ponderosae occupying different host trees in common localities, and to determine whether there was molecular evidence for a past demographic expansion. Using a combination of amplified fragment length polymorphism (AFLP) and mitochondrial sequencing analyses, we found evidence of genetic structuring among populations that followed a broad isolation-by-distance pattern. Our results suggest that the geographical pattern of gene flow follows the core distribution of the principal D. ponderosae host species, around rather than across the Great Basin and Mojave Deserts. Patterns of haplotype diversity and divergence were consistent with a range-wide population expansion. This signal was particularly pronounced in the northern part of the species' range, where outbreak activity is currently increasing. Using AFLP markers, we were unable to detect significant differences among groups of insects sampled from different host trees in common locations. Incidentally, we found that a large proportion of the polymorphic AFLP markers were gender-specific, occurring only in males. While we did not include these markers in our analyses, this finding warrants further investigation.     

Abundance and Movement of Understory Birds in a Maulino Forest Fragmented by Pine Plantations

Understory birds are especially vulnerable to habitat fragmentation because of the reduction in habitat quality and bird movement. We study the separate effects of understory, overstory and landscape on four understory birds (tapaculos), in Central Chile, comprising a landscape mosaic of pine (Pinus radiata D. Don) plantations and native Maulino forest fragments. We also determined whether habitats with poor understory could be barrier to tapaculos movements. Abundance was measured using stationary playbacks and habitat barrier through playbacks. Understory structure was the main factor that predicted tapaculos presence and abundance. Two species, the Andean Tapaculo (Scytalopus magellanicus fuscus Gmelin) and the Ochre-flanked Tapaculo (Eugralla paradoxa Kittlitz), were positively associated with dead pine branches and negatively to forest fragment size. Tapaculos were less abundant in mature native forest, but appeared willing to cross between different habitat types. However, the Chestnut-throated Huet-huet (Pteroptochos castaneus Philippi and Landbeck), did not move from forest fragments to pine with poor understory. Overall, tapaculos species varied in their response to fragmentation depending on their habitat selection and movement capacities.     

长白山原始阔叶红松林林下草本植物多样性格局及其影响因素

于2009年生长季(5～9月份)对布置在1.0hm2的原始阔叶红松林样地内的100个草本样方(1m×1m)进行物种及其植株高度、盖度以及土壤因子、光照环境进行调查,分析不同层片与土壤因子、光照环境的相关性以及乔木层、灌木层、土壤因子、光照环境对草本层多样性格局的影响,以阐明长白山阔叶红松林林下草本层多样性格局及其影响因素。结果表明:(1)阔叶红松林下的草本植物存在明显的季节动态变化,其中,5月份黑水银莲花(Anemone amurensis)、延胡索(Corydalis ambigua)等早春短命植物占优势,6月份部分早春短命植物和早夏植物、晚夏植物同时出现,物种丰富度最高,7月份是夏季植物的生长旺季,物种多样性最高,8～9月份草本植物逐渐枯萎。(2)灌木层的多样性与草本层的多样性呈极显著正相关,灌木层的株数密度与不同月份的草本层盖度呈正相关、与草本层多样性呈负相关,而乔木层对草本层盖度和多样性的影响均不显著。(3)土壤速效钾、光照和土壤水分是关联乔木层和草本层的重要环境因子。研究认为,阔叶红松林下的乔木层通过影响光照、降水及枯落物的含量而间接影响草本层的多样性格局。     

Verbenone-releasing flakes protect individual Pinus contorta trees from attack by Dendroctonus ponderosae and Dendroctonus valens (Coleoptera: Curculionidae, Scolytinae)

1 In a study site in interior northern California, twenty individual lodgepole pines Pinus contorta were sprayed with a suspension of DISRUPT Micro‐Flake^® Verbenone (4,6,6‐trimethylbicyclo(3.1)hept‐3‐en‐2‐one) Bark Beetle Anti‐Aggregant flakes (Hercon Environmental, Emigsville, Pennsylvania) in water, with sticker and thickener, from ground level to a height of 7 m. Twenty trees sprayed with just water, sticker and thickener served as controls. All trees were baited immediately after spraying with mountain pine beetle Dendroctonus ponderosae aggregation pheromone lures, and lures were refreshed after 4 weeks. 2 Trees treated with verbenone had significantly lower attack density by D. ponderosae than controls at 2, 4, 6 and 8 weeks after application of flakes. 3 None of the treated trees was attacked by red turpentine beetle Dendroctonus valens, whereas control trees averaged nearly two D. valens attacks per tree, 8 weeks after treatment. 4 A dry frass index, used to predict ultimate tree mortality, was significantly higher in control trees than treated trees for all four sampling intervals. This index proved to be a significant predictor of ultimate tree mortality. 5 Ten months after application, treated trees showed significantly lower mortality than control trees.     

Reproductive isolation and environmental adaptation shape the phylogeography of mountain pine beetle (Dendroctonus ponderosae)

Chromosomal rearrangement can be an important mechanism driving population differentiation and incipient speciation. In the mountain pine beetle (MPB, Dendroctonus ponderosae), deletions on the Y chromosome that are polymorphic among populations are associated with reproductive incompatibility. Here, we used RAD sequencing across the entire MPB range in western North America to reveal the extent of the phylogeographic differences between Y haplotypes compared to autosomal and X‐linked loci. Clustering and geneflow analyses revealed three distinct Y haplogroups geographically positioned within and on either side of the Great Basin Desert. Despite close geographic proximity between populations on the boundaries of each Y haplogroup, there was extremely low Y haplogroup mixing among populations, and gene flow on the autosomes was reduced across Y haplogroup boundaries. These results are consistent with a previous study suggesting that independent degradation of a recently evolved neo‐Y chromosome in previously isolated populations causes male sterility or inviability among Y haplotype lineages. Phylogeographic results supported historic contraction of MPB into three separate Pleistocene glacial refugia followed by postglacial range expansion and secondary contact. Distinct sets of SNPs were statistically associated with environmental data among the most genetically distinct sets of geographic populations. This finding suggests that the process of adaptation to local climatic conditions is influenced by population genetic structure, with evidence for largely independent evolution in the most genetically isolated Y haplogroup.     

Redescription and Life History of Contortylenchus brevicomi,a Parasite of the Southern Pine Beetle Dendroctonus frontalis

Larval and adult life stages are described for Contortylenchus brevicomi (Massey) Rühm parasitizing a Mississippi population of Dendroctonus frontalis, the southern pine beetle. Fourth-stage larvae and free-living adult females of this species are identified and described for the first time. The life cycle of C. brevicomi can be reconstructed from this study. The adult female nematode lays eggs in a mature beetle. Larval development progresses within the hemocoel until fourth-stage larvae exit the host. Mating occurs in beetle galleries and only females enter an immature beetle host.     

Understory Plant Community Composition Is Associated with Fine-Scale Above- and Below-Ground Resource Heterogeneity in Mature Lodgepole Pine (Pinus contorta) Forests

Understory plant communities play critical ecological roles in forest ecosystems. Both above- and below-ground ecosystem properties and processes influence these communities but relatively little is known about such effects at fine (i.e., one to several meters within-stand) scales, particularly for forests in which the canopy is dominated by a single species. An improved understanding of these effects is critical for understanding how understory biodiversity is regulated in such forests and for anticipating impacts of changing disturbance regimes. Our primary objective was to examine the patterns of fine-scale variation in understory plant communities and their relationships to above- and below-ground resource and environmental heterogeneity within mature lodgepole pine forests. We assessed composition and diversity of understory vegetation in relation to heterogeneity of both the above-ground (canopy tree density, canopy and tall shrub basal area and cover, downed wood biomass, litter cover) and below-ground (soil nutrient availability, decomposition, forest floor thickness, pH, and phospholipid fatty acids (PLFAs) and multiple carbon-source substrate-induced respiration (MSIR) of the forest floor microbial community) environment. There was notable variation in fine-scale plant community composition; cluster and indicator species analyses of the 24 most commonly occurring understory species distinguished four assemblages, one for which a pioneer forb species had the highest cover levels, and three others that were characterized by different bryophyte species having the highest cover. Constrained ordination (distance-based redundancy analysis) showed that two above-ground (mean tree diameter, litter cover) and eight below-ground (forest floor pH, plant available boron, microbial community composition and function as indicated by MSIR and PLFAs) properties were associated with variation in understory plant community composition. These results provide novel insights into the important ecological associations between understory plant community composition and heterogeneity in ecosystem properties and processes within forests dominated by a single canopy species.     

林下层植物在退化马尾松林恢复初期养分循环中的作用

以鼎湖山退化马尾松 (Pinusmassoniana)林恢复过程中林下层植物凋落物、分解和养分动态为对象 ,研究了林下层植物在退化马尾松林恢复初期养分循环中的作用。结果表明 ,林下层年凋落物量除在第 5年有所下降外均随时间逐年上升 ,但其增加速率随年份不同而异 ,总平均年增长速率为 3 8%。第 4年凋落物量为 0 .2 0 t· hm- 2 · a- 1,第 1 1年为 1 .1 7t·hm- 2·a- 1。凋落物养分元素平均浓度为 (% ) :N0 .95 ,P0 .0 4,K0 .5 7,Ca0 .1 3和 Mg0 .0 8,基本上以夏季和秋季最高冬春交替月份最低。第 1 1年凋落物各元素养分归还量为 (kg· hm- 2·a- 1) :N1 1 .1 0 ,P0 .47,K6.65 ,Ca1 .48和 Mg 0 .91。凋落物在分解过程中失重率呈直线模型变化 ,第 1年的分解速率为 3 1 % ,至试验结束时凋落物的残存量占起始量的 66%。在凋落物分解过程中 ,N和 P浓度随时间逐渐上升 ,但 N增加的速度较 P快 ,其余元素浓度均下降 ,但 K下降的速度最快。在凋落物分解过程中 ,N是唯一表现残留量呈先上升然后下降变化的元素。P的残留量变化与凋落物的失重率变化几乎一致。各元素在分解试验结束时残留量占起始量的百分比分别为 :N 90 % ,P 67% ,K 9% ,Ca 3 0 %和Mg 1 4%。可见 ,林下层凋落物在退化马尾松林恢复初期碳及其它营养元素循     

Evolving Management Strategies for a Recently Discovered Exotic Forest Pest: The Pine Shoot Beetle, Tomicus Piniperda (Coleoptera)

Established populations of the Eurasian pine shoot beetle (Tomicus piniperda (L.); Coleoptera: Scolytidae) were first discovered in North America in Ohio in 1992. As of 31 December 2000, T. piniperda was found in 303 counties in 12 US states (Illinois, Indiana, Maine, Maryland, Michigan, New Hampshire, New York, Ohio, Pennsylvania, Vermont, West Virginia, and Wisconsin) and in 43 counties in 2 Canadian provinces (Ontario and Quebec). A federal quarantine imposed in November 1992 regulates movement of pine (Pinus) trees, logs, and certain pine products from infested to uninfested areas within US. The forest products, Christmas tree, and nursery industries are affected by the quarantine. This paper summarizes information on the discovery and spread of T. piniperda in North America, survey efforts, recent interception history, development and changes in the federal quarantine, development of a national compliance management program, and extension and research efforts.     

凉水原始阔叶红松林与白桦次生林植物多样性的比较研究

采用典型抽样法和生态群落学调查法对黑龙江凉水国家级自然保护区紫椴（Tilia amurensis）红松（Pinus koraiensis）林、蒙古栎（Quercus mongolica）红松林及白桦（Betula platyphylla）次生林的植物群落结构特征和物种多样性进行了研究。结果表明,紫椴红松林有维管束植物31种,隶属21科,25属;蒙古栎红松林有维管束植物32种,隶属25科,28属;白桦次生林有维管束植物31种,隶属20科,28属。白桦次生林乔木层和草本层的物种丰富度S指数显著高于2种阔叶红松林,而其灌木层的物种丰富度S指数最低;3种林型乔木层的Simpson物种多样性指数无显著差异,紫椴红松林的灌木层Simpson指数显著高于其他2种林型,3种林型草本层之间的Simpson指数差异显著;紫椴红松林乔木层的Shannon-Wiener物种多样性指数显著低于其他2种林型,白桦次生林灌木层的Shannon-Wiener指数显著低于2种阔叶红松林,蒙古栎红松林草本层的Shannon-Wiener指数最高;此外,蒙古栎红松林乔木层的Pielou均匀度指数显著高于其他林型,而3种林型灌木层的Pielou指数无显著差异,紫椴红松林草本层的Pielou指数最低。