Features of the life cycles of <Emphasis Type="Italic">Pterostichus montanus</Emphasis> (Motschulsky, 1844) and <Emphasis Type="Italic">Carabus loschnikovi</Emphasis> (Fischer-Waldheim, 1822) (Coleoptera,Carabidae) in conditions of the mountain taiga belt in the Eastern Sayan

We studied the seasonal dynamics and demographic structure of abundant ground beetle species from the mountain taiga belt in the Eastern Sayan. Data on the dynamics of the sexual and age structure of the populations as well as on the reproductive capacity of females in the biotopes on the slopes with different exposure and height were obtained. Life cycles with one-and two-year development were revealed for the ground beetles typical for the mountain taiga belt. As an example, data on the life cycles of Pterostichus montanus (Motschulsky, 1844) with one-year spring development and Carabus loschnikovi (Fischer-Waldheim, 1822) with two-year polyvariant multiseasonal development are given for the first time. Two strategies were revealed in the life cycles of ground beetles under alpine conditions: an accelerated population development in spring one-year species and a two-year development with pronounced polyvariance in two intrapopulation groups of ground beetles of the Carabus genus. The seasonal dynamics of the activity and reproduction periods proved to vary for these species on the slopes along the height gradient.Translated from Izvestiya Akademii Nauk, Seriya Biologicheskaya, No. 1, 2005, pp. 36–46.Original Russian Text Copyright © 2005 by Sharova, Khobrakova.     

Long-term dynamics and structure of shrews association (Soricidiae) in the mountain taiga of the Eastern Sayan

The dynamics of shrews association in the mountain taiga of the Eastern Sayan was considered during the years 1981 to 2010. The work was carried out in the territory of “Stolby” reserve. The structure of the association, the duration of cyclic changes of different species, effect of succession and climatic processes on the composition and structure of the community have been studied.     

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.     

A Structured Threshold Model for Mountain Pine Beetle Outbreak

A vigor-structured model for mountain pine beetle outbreak dynamics within a forest stand is proposed and analyzed. This model explicitly tracks the changing vigor structure in the stand. All model parameters, other than beetle vigor preference, were determined by fitting model components to empirical data. An abrupt threshold for tree mortality to beetle densities allows for model simplification. Based on initial beetle density, model outcomes vary from decimation of the entire stand in a single year, to inability of the beetles to infect any trees. An intermediate outcome involves an initial infestation which subsequently dies out before the entire stand is killed. A model extension is proposed for dynamics of beetle aggregation. This involves a stochastic formulation.     

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.     

贺兰山冲积扇荒漠草地拟步甲群落小尺度空间格局动态

为探讨贺兰山冲积扇荒漠草地拟步甲群落小尺度空间格局动态特征,本研究于2019年5—10月每月采用陷阱法调查200 m×200 m空间尺度的拟步甲昆虫群落,基于地统计分析方法,研究拟步甲昆虫群落的空间格局,并分析空间格局与地形因子的关系。6次调查共捕获拟步甲科成虫7属10种1086只,群落组成具有较明显的空间和时间变异性。其中,拟步甲群落多样性以5月最大,7月最低。Moran I指数表明,拟步甲优势种空间自相关性具有明显的季节波动性,在5月、9月和10月表现为显著空间正相关。拟步甲群落和优势种具有较强的空间异质性,主要由结构性因素决定。普通克里金插值表明,不同季节群落空间格局梯度分布存在明显差异,夏季群落空间格局比较简单。交叉方差函数分析显示,优势种物种间大多表现为正空间关联性,且主要受结构性因素单一调控。典范对应分析(CCA)结果表明,坡度和海拔显著影响昆虫分布。本研究揭示了拟步甲昆虫群落小尺度空间异质性具有明显的季节变异,为研究贺兰山冲积扇生境地表甲虫物种多样性空间格局及维持机制奠定基础。     

伏牛山森林生态系统灌木植物功能群分类

伏牛山国家级自然保护区位于中国东部亚热带和暖温带的过渡区,植被优势种明显,以灌木优势种为主体进行植物功能群分类,可以对森林生态系统的功能、框架结构及类群分布有一个明确的认识.采用群落生态学的调查方法,在伏牛山南北坡设置66个典型样方.根据调查结果,通过计算重要值,选取优势度相对较大的灌木树种进行种间联结及相关性分析,以X2(卡方)检验为基础,结合联结系数AC和共同出现百分率PC来测定灌木优势种间的联结性,根据优势种间的联结性及其在海拔梯度上的变化异同来划分植物功能群,把灌木优势种划分为7组植物功能群.植物功能群间物种表现出显著正联结,一起在同一生境中出现的几率较大,在长期的生长演化过程中,能适应相似的资源环境和对干扰有相似的响应,所以将其划分同一组植物功能群.植物功能群内有一些重要的形态特征有相似之处,功能群间的形态特征有明显的区别,如叶片的大小、形状等,功能群的这些相似及区别还需要生理生态学的进一步研究.     

白皮松林林隙物种组成动态及更新规律

研究了白皮松林林隙物种组成和种间关系在林隙发育的各个阶段的动态变化,并对林隙内主要树种的更新反应进行了探讨。结果表明,在发育阶段为前期、中前期、中后期和后期的序列上,林隙中乔木层优势度最大的物种分别是白皮松（Pinus bungeana）、辽东栎（Quercus wutaishanica）、山桃（Prunus davidiana）和侧柏（Platy-cladus orientalis）;灌木层优势种包括白莲蒿（Artemisia gmelinii）、胡枝子（Lespedeza bicolor）、薄皮木（Lepto-dermis oblonga）和狼牙刺（Sophora viciifolia）;草本层优势种包括大披针苔草（Carex lanceolata）、糙隐子草（Cleistogenes squarrosa）、紫菀（Aster tataricus）和荩草（Arthraxon hispidus）;非林隙林分的优势种不明显。在林隙发育过程中乔木层、灌木层、草本层的平均相似性系数为0.32、0.38和0.27。可见林隙发育过程中群落各层次的相似性系数由小到大顺序为：草本〈乔木〈灌木;草本植物随着林隙环境条件的改变而出现较大的消亡和更新,显示出较大的波动性;乔木的稳定性和连续性介于灌木和草本之间。依据林隙中树种幼苗、幼树Ⅰ和幼树Ⅱ的数量差异,可将林隙更新反应分为单峰型、双峰型、差异不明显3种,并分别以白皮松、辽东栎和油松（Pinus tabu-laeformis）为代表树种。     

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.     

贺兰山甲虫物种丰富度分布格局及其环境解释

理解山地物种丰富度分布格局及其成因对于山地生物多样性保护具有重要意义。本文基于贺兰山地区甲虫31科252属469种的分布信息, 结合相关气候与生境异质性数据, 系统地探讨了贺兰山地区甲虫及6个优势科物种丰富度地理格局及其影响因素。结果表明, 甲虫物种丰富度及科属区系分化强度以贺兰山中段最高, 南段比北段高, 西坡比东坡高。基于183个栅格内物种分布的二元数据聚类分析, 贺兰山甲虫分布可分为北段强旱生景观甲虫地理群、中西段半湿生景观甲虫地理群、中东段及南段半旱生景观甲虫地理群3个地理群。冗余分析(RDA)表明年均温和年均降水量是影响最显著的因子。方差分解结果显示, 水分与能量因子共同解释了全部甲虫物种丰富度57.1%的空间变异, 单独解释率分别为5.9%和7.1%。生境异质性解释了全部甲虫物种丰富度35.2%的变异, 单独解释率仅为1.8%。气候因素与生境异质性对不同优势科物种丰富度的相对影响并不一致。在贺兰山的南段和北段, 生境异质性和水分因子对甲虫物种丰富度影响作用明显。水分和能量因子是贺兰山地区甲虫物种丰富度空间分布格局的主导因子, 生境异质性有助于提高甲虫物种丰富度。从未解释的比例来分析, 地形和土壤因素可能对贺兰山甲虫物种丰富度存在重要影响。     

Differences in ground beetles (Coleoptera: Carabidae) of original and reconstructed tallgrass prairies in northeastern Iowa, USA, and impact of 3-year spring burn cycles

Ground beetle assemblages were monitored at four tallgrass prairie sites burned on 3-year cycles in northeastern Iowa. The objectives of this study were to quantify differences in carabid communities between original and reconstructed tallgrass prairies, and to determine the responses of ground beetles to 3-year cycles of early spring fire commonly used to manage tallgrass prairies. Using pitfall traps, ground beetle assemblages in two original and two reconstructed tallgrass prairies were compared between 1994 and 1998, where beetles were sampled annually (0-, 1-, and 2-year post-fire conditions) from plots burned every 3 years. When burned, the greatest abundance, activity density, and species richness of carabid beetles occurred the year immediately following a spring burn, with abundance declining steadily with increased time since burning. Overall ground beetle diversity as determined by Shannon's diversity index was greatest in original tallgrass prairies several years after a fire. Some species of ground beetles were found only in original prairies, while others were found primarily in reconstructed prairie. Similarly, some species were more abundant the year immediately following a burn, while others were found in greater abundance with increased time since fire. NMS ordination and indicator species analysis clearly show differences in carabid species between original and reconstructed tallgrass prairies, but did not show differences among burn treatments.     

大兴安岭地区兴安落叶松林步甲群落多样性时间动态分析

兴安落叶松林是大兴安岭地区代表性的植被类型,其生物多样性具有独特性。步甲是森林生态系统环境和多样性的指示性物种,以及认识环境变化和生物多样性特征的关键物种。为研究大兴安岭地区兴安落叶松林步甲群落多样性的时间动态,分析步甲群落对时间变化的响应规律,于2019年5月下旬-8月下旬步甲活跃期,利用陷阱诱捕法在兴安落叶松林5个样地中采集步甲标本共15属34种1149头,其中大兴安岭地区地理新纪录物种7种,中国地理新纪录物种6种。研究结果表明,通缘步甲属（Pterostichus）和大步甲属（Carabus）物种丰富度最高;通缘步甲属未定种5（Pterostichus eximius）为极优势物种,对时间变化最敏感。兴安落叶松林小生境类型的多样化和步甲休眠期的选择是步甲群落个体数和物种数随时间变化呈双峰模式的主要因素,最高峰均出现在6月下旬;多样性与均匀度指数均在7月上旬达高峰期,8月下旬多样性下降而均匀度上升,各指数之间相关性较低。步甲群落在6月下旬到7月下旬对环境具有较高的适应度;群落结构在环境条件相对稳定的6月下旬到7月下旬和8月上旬到8月下旬均表现为极相似（I>0.75）。物种取食特征和生活史策略的多样化使步甲群落各指数随时间变化具有显著差异,而物种取食特征和生活史策略受环境因子的综合影响较大。稳定的森林环境条件下,步甲活跃期更长,群落结构相似度更高。该结果为步甲群落时间动态研究奠定了一定的理论基础,为大兴安岭地区地下生物多样性的保护和管理策略制定提供了一定的理论和数据依据。     

Mites (Acari) phoretic on ground beetles (Coleoptera,Carabidae) in a southern taiga forest in the environs of Vologda

Ground beetles were captured in a mixed southern taiga forest near the city of Vologda, Russia, from March to September 2014 using different collection techniques. Of the 250 specimens of ground beetles belonging to 22 species, 109 beetles representing 11 species yielded 326 specimens of mites of the cohorts Gamasina (4 species), Astigmatina (5), and Heterostigmatina (1). Two mite species, Antennoseius pseudospinosus Eidelberg, 1990, a common species in the steppes of southeastern Europe, and Halodarcia incideta Karg, 1969, a polyzonal European hydrophile, are recorded in the taiga zone for the first time. Dorsipes dorsipes Regenfuss, 1968, a specialized parasite of beetles of the genus Carabus Linnaeus, 1758, is new to the fauna of Eastern Europe. An adult mite of the genus Stylochirus G. Canestrini et R. Canestrini, 1882 was found for the first time in a natural hibernating chamber in close contact with an overwintering ground beetle, in particular, a male of S. fimetarius (Müller, 1859) on Carabus granulatus Linnaeus, 1785. The most common mite to occur on ground beetles was Antennoseius bullitus Karg, 1969, which was found on 7 carabid species with the mean occurrence of 41% and comprised 68% of the total mite sample. Joint phoresy of 2–3 mite species was recorded on 12 specimens of ground beetles; in 5 cases the co-occurring mites were Antennoseius bullitus and Stylochirus fimetarius. Three dominant mite species (85% of the total mite sample) were mainly collected off three dominant carabid species (70% of the beetle sample), but individual mite species preferred different hosts.     

Diversity and seasonal variation of ground and understory spiders from a tropical mountain cloud forest

We made intensive samplings to study the seasonal response of spiders across different forest strata (ground and understory) in a tropical mountain cloud forest from Mexico. We sampled spiders from ten plots in six sampling events during the dry and rainy season, to analyze their abundance, structure (distribution of abundance among species), diversity and the response of the five dominant species at each stratum. Results demonstrated that seasonal patterns of spider communities differed among strata, revealing a complex spatiotemporal dynamic. Abundance, structure, diversity of ground spiders, as well as the responses of four dominant species at this stratum, showed low seasonal variations. In contrast, a strong seasonal variation was observed for the understory assemblage, with lowest abundance and highest diversity in the rainy season, and different assemblage structures for each season. Seasonal patterns of each assemblage seem linked to the responses of their dominant species. We found high co‐occurrence among most of the ground dominant species with similar habitat use and with multivoltine patterns, contrasting with low co‐occurrence among most of the understory dominant species with similar habitat use and univoltine patterns. Our results showed that the spiders’ assemblages of tropical mountain cloud forest (opposed to what is found in temperate and boreal forests) increase their species richness with the height, and that their responses to seasonal change differ between strata. Management programs of these habitats should consider the spatial and temporal variations found here, as a better understanding of their ecological dynamics is required to support their sustainable management.     

秦岭太白山北坡蝗虫的群落结构

在秦岭太白山北坡23个样地内采集到蝗虫2总科7科25属34种。应用数量分类、多样性指数及相似性分析方法对太白山北坡蝗虫群落垂直分布样方资料进行了分析。应用数量分类法可将蝗虫群落按植被类型分为8种,即农耕带、栓皮栎林带、锐齿栎林带、辽东栎林带、桦木林带、冷杉林带、落叶松林带、高山草饲带。太白山蝗虫群落垂直分布规律明显。其中高海拔（如高山草甸带）的蝗虫种类数和多样性明显低于中低海拔,中低海拔（锐齿栎林带）蝗虫种类和多样性指数最大。但低海拔的农耕带人为干扰大,植被较单一,蝗虫优势种的优势度大,蝗虫密度大,而种类数较少。     

Ground beetles (coleoptera,carabidae) of the Ola Plateau highlands,Kolyma Uplands

The ground beetle fauna was studied in 12 biotopes of the tundra belt on Ola Plateau, 130 km NW of Magadan. In all, 16 species were found above timber line (900–1220 m), which is comparable with the diversity in the mountain tundras of the upper reaches of the Kolyma (Berman et al., 1984; Bukhkalo, 1997), but the faunistic similarity of these regions is low (Jaccard’s coefficient 22–29%). Besides the widespread species, a group of ground beetles with limited ranges is present; it includes Carabus kolymensis Lafer, 1989 occurring in the East Siberian mountain tundras, the Siberian-Nearctic, mainly arctic Pterostichus agonus G. Horn, 1880, and the East Siberian Pterostichus eximius A. Mor., 1862. The use of the term “arctic species” in arealogical and ecological studies is discussed.     

Orchard and riparian habitats enhance ground dwelling beetle diversity in Mediterranean agro-forestry systems

The relative contribution of mixed orchard and riparian vegetation patches to local and regional diversity of Mediterranean landscapes dominated by cork-oak woodlands was tested in 2006–2007 using ground, rove and darkling beetles (Coleoptera: Carabidae, Staphylinidae, Tenebrionidae). Mixed orchard and riparian gallery habitats recorded higher values of abundance and species richness for overall beetle species, although most darkling beetle species were associated with the sclerophyllous cork-oak woodlands. Ground and rove beetle community structure changed from the orchard and riparian habitats to samples placed 100 m away into the surrounding cork-oak woodland, i.e., non-cork-oak patches enhanced beta diversity within the landscape mosaic. Analysis of ground beetle traits concerning moisture preferences revealed a higher proportion of hygrophilous species in mixed orchard and riparian gallery habitats while xerophilous species were dominant within the cork-oak woodlands. The results of this study suggested that land-use management promoting the maintenance of habitat heterogeneity enhances biodiversity conservation of important hygrophilous and xerophilous species, and subsequently the sustainable use of Mediterranean agro-forest mosaics.     

Altitudinal variation of the life cycle of Poecilus fortipes (Coleoptera,Carabidae) in Eastern Siberia

Altitudinal variation of the demographic structure of populations of Poecilus fortipes (Chaud.) was studied in four mountain localities in the south of Eastern Siberia, and the life cycle of the species was reconstructed for each locality. The size of adults of both sexes decreased along the altitudinal gradient: the beetles from the Selenga middle mountains (600–650 m above sea level) were much larger, while those at the upper boundary of the forest-steppe belt in the Eastern Sayan (1500–1700 m asl) were smaller than others. Their linear dimensions decreased more distinctly in the transitional zone between middle and high mountains. The duration of the reproductive period did not depend on the altitude. However, the onset of oviposition shifted along the altitudinal gradient to the beginning of the vegetation season, due to its shortening and a decrease in the sum of effective temperatures. As the altitude increased, the mean number of eggs per female declined significantly from 15.9 ± 6.1 to 8.6 ± 5.8, whereas the maximum number of eggs increased from 26–28 to 40–43. On the contrary, the dates of emergence shifted gradually from the beginning and middle of the vegetation period to its middle and end. Populations from the Selenga middle mountains, the Vitim Plateau, and the lower margin of the forest-steppe belt in the Eastern Sayan mountains were characterized by a relatively low catch index and the absence of rebreeding old individuals, both these features increasing the risk of spontaneous local extinction. The annual life cycle with summer breeding was replaced along the altitudinal gradient with an obligate biennial life cycle with summer and early-summer breeding. This transition was observed as the altitude increased from 600–650 to 900–950 m asl while the sum of effective temperatures dropped below 1400°C, i.e., the value needed for implementation of the annual life cycle.     

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.