The role of various meadow margin types in shaping carabid and staphylinid beetle assemblages (Coleoptera: Carabidae,Staphylinidae) in meadow dominated landscapes

Less intensively managed semi-natural habitats, e.g., field and meadow margins like hedgerows, are thought to be crucial landscape components for maintaining biodiversity in highly disturbed and intensively managed agricultural landscapes. In this study, we focused on the effects of three meadow margin types on activity-density, species richness and species composition of carabid and staphylinid beetles recorded by pitfall traps in Central European landscapes dominated by intensively managed meadows. Carabid activity-density was significantly higher in meadows than in meadow margins and within meadow margins their activity-density increased from grassy meadow margins via shrubby ones to woody meadow margins. We found that recorded species richness of both carabid and staphylinid beetles was not significantly affected by habitat identity (meadow margin or neighbouring meadow) and meadow margin type. Recorded species composition of both investigated taxa was significantly affected by habitat identity and interaction between habitat identity and meadow margin type (i.e. it differed between particular meadow margin types). Assemblages inhabiting various meadow margin types were more dissimilar between each other than assemblages from neighbouring meadows. Meadow margins within grassland dominated landscapes maintain local species richness by hosting different species from those living in surrounding meadows. Dissimilarity of carabid and staphylinid assemblages from meadows neighbouring both sides of particular meadow margin did not differ between meadow margin types. Our results indicate that semi-natural habitats play an important role in maintaining biodiversity not only in agricultural landscapes dominated by arable fields, but also in those dominated by meadows.     

北京密云农业景观步甲群落空间分布格局

采用陷阱法对北京密云县西田各庄南部农业景观中玉米地、花生地、果园及半自然林地4种典型生境中的步甲群落进行调查分析.结果表明: 果园步甲群落α多样性最高,花生地最低,林地与玉米地居中,且二者无显著差异;林地、花生地及果园的步甲群落结构明显不同,但均与玉米地的步甲群落结构有不同程度的相似性.果园维持着较多的捕食性步甲和兼食性步甲个体数;且不同生境间捕食性步甲的物种周转率与兼食性步甲的差异更明显.低集约化的果园生境可能较半自然生境林地维持更高的步甲群落α多样性,但多样化的景观组成有利于维持步甲群落及捕食性步甲较高的β多样性.重视景观多样性和低集约化农田生境的保护对保护农业景观步甲群落多样性和实现其害虫控制功能具有重要的意义.     

Diversity and rarity hotspots and conservation of butterfly communities in and around the Aokigahara woodland of Mount Fuji,central Japan

In central Japan, Aokigahara woodland is considered to be one of the most natural areas around Mount Fuji and a core area in the conservation of the biodiversity of Mount Fuji. We chose butterflies as an indicator species of biodiversity and examined six communities in and around the woodland in 2000 using transect counts to examine and search for diversity and rarity hotspots and their associated landscapes. The results showed that butterfly species richness and species diversities H 1/ were significantly higher in forest-edge sites than in forest-interior and/or open-land sites, and variation in the total number of species among these three landscape types was well accounted for by ecologically specialist species, such as landscape specifics, oligovoltines, narrow diet feeders and low-density species. Thus, the species regarded as vulnerable to extinction, including Red List species, were observed more often in forest-edge sites than in forest-interior and/or open-land sites. As a result, in the study area, diversity and rarity hotspots were found in forest-edge landscapes. The reasons why butterfly diversity and rarity hotspots were established in forest-edge landscapes were analyzed and interpreted from several points of view, including disturbance level, landscape elements and plant species richness. From these results, and the fact that some species were confined to forest-interior sites, we conclude that it is very important to conserve and manage forest-edge habitats (considered to be semi-natural) as well as forest-interior habitats (considered to be the most natural) to maintain the diversity of butterfly communities and preserve the various types of threatened species in and around the Aokigahara woodland.     

Landscape- and time-dependent benefits of wildflower areas to ground-dwelling arthropods

Wildflower areas are a popular agri-environment scheme to counteract agro-biodiversity loss. Yet, their benefits are controversially discussed. Since inconsistent benefits may be owed to landscape context and temporal dynamics, we applied a multi-year study to unravel effects of permanent and transient habitats on ground-dwelling arthropods in wildflower areas and cereal fields.Across three consecutive years, we studied activity density, species richness and community composition of rove beetles, carabid beetles and spiders in ten pairs of wildflower areas and cereal fields along independent gradients of proportions of permanent semi-natural habitats or transient wildflower areas.Arthropod responses to the proportions of permanent semi-natural habitats often followed a hump-shaped pattern, whereas transient wildflower areas seemed to drive linear responses. An interactive effect on rove beetle richness in wildflower areas implies that benefits were highest either at intermediate proportions of permanent semi-natural habitats or at lower proportions, but with additional availability of transient wildflower areas or in landscapes with low proportions of permanent semi-natural habitats, but high proportions of wildflower areas. However, ground-dwelling arthropod activity density or species richness did not systematically increase over the three study years.Our results suggest that both permanent and transient habitats differ in how they affect biodiversity, possibly due to different temporal continuity and resource diversity. Benefits of permanent semi-natural habitats seemed highest at an equilibrium between an increasing resource-related species pool and an increasing diversity dilution, whereas benefits of transient wildflower areas seemed to increase with resource complementarity and connectivity at the landscape scale. Nevertheless, both habitats seem to complement each other and considered in concert, seem to be most effective in promoting benefits of wildflower areas to ground-dwelling arthropods at intermediate landscape complexity. The substantial variation in diversity patterns among years with weather extremes, suggests that optimizing benefits of wildflower areas requires further multi-year studies.     

Landscape effects on the thermotolerance of carabid beetles and the role of behavioral thermoregulation

Physiological thermotolerance and behavioral thermoregulation are central to seasonal cold adaptation in ectothermic organisms. For species with enhanced mobility, behavioral responses may be of greater importance in the cold stress response. Employing the carabid beetles as a study organism, the current study compared physiological thermotolerance and behavioral thermoregulation in carabid species inhabiting cereal fields in different landscape contexts, from fine grain heterogeneous “complex” landscapes to homogenous “simple” landscapes. Physiological thermotolerance was determined via measurement of the CT_min and chill coma temperature. Behavioral responses to cold temperature exposure were determined employing a purpose built arena, and thoracic temperature measured to estimate the efficacy of the behavior as a form of behavioral thermoregulation. Results revealed an influence of landscape composition on the cold tolerance of carabid beetles, although species differed in their sensitivity to landscape intensification. A reduced effect of landscape on the thermotolerance of larger carabid beetles was observed, thought to be the consequence of greater mobility preventing local acclimation to microclimatic variation along the landscape intensification gradient. Investigation into behavioral thermoregulation of the 3 largest species revealed burrowing behavior to be the main behavioral response to cold stress, acting to significantly raise carabid body temperature. This finding highlights the importance of behavioral thermoregulation as a strategy to evade cold stress. The use of behavioral thermoregulation may negate the need to invest in physiological thermotolerance, further offering explanation for the lack of landscape effect on the physiological thermotolerance of larger carabids.     

Introduced and native ground beetle assemblages (Coleoptera: Carabidae) along a successional gradient in an urban landscape

According to the intermediate disturbance hypothesis (IDH), species diversity should be higher at sites with intermediate levels of disturbance. We tested this hypothesis using ground beetles (Coleoptera: Carabidae) collected in pitfall traps from sites that varied in time since last disturbance. This successional gradient was embedded in an urban landscape near Montreal, Quebec. We predicted that diversity in young forests and old fields would be higher than in agricultural fields and old forests. Fifty-five species (2932 individuals) were found in 2003 and 46 species (2207 individuals) in 2004. In both years, species richness was highest from traps placed in agricultural fields. We collected nine introduced species; these had higher catch rates than the native species in both years (64.8% of total catch). When introduced species were removed from the Nonmetric Multidimensional Scaling ordination analysis, the assemblages from agricultural fields were less distinct compared to those of the other habitats, suggesting the introduced fauna is important in structuring carabid assemblages from the agricultural fields. Introduced species may play a significant role in the community composition of ground beetles in urban landscapes, and their influence may be the cause of the lack of support found for the IDH.     

Woodland remnants as an urban wildlife refuge: a cross-taxonomic assessment

Urban nature is crucial for the quality of human life both within cities and beyond. In many developed cities, the numbers of restoration-through-revegetation projects have rapidly increased over the decades. However, the extent to which revegetated habitats perform compensatory roles for remnant habitats is poorly understood. We compared butterfly and ground beetle assemblages among three park types (five remnant parks, four newly established parks and five old established parks) and seven built-up sites in Tokyo, central Japan. Butterflies were classified into woodland or open-land and into patch-dependent or matrix-dwelling species. For both taxa, remnant parks and built-up sites had the highest and lowest species richness and abundance, respectively. Although the richness and abundance of open-land and matrix-dwelling butterflies did not differ among the three park types, those of woodland and patch-dependent species were significantly highest in remnant parks. In short, after 50 years, established parks did not attain the same insect assemblages as those in remnant parks. These results illustrate that whist revegetation is an effective and fast-acting conservation measure for generalist species (i.e., widely distributed species), this value is limited for specialists. In highly urbanised landscapes, therefore, even small remnant woodlands provide important refuges for urban wildlife. Remnant protection programs at the early stage of city development would decide the fate of urban biodiversity.     

Cork-oak woodlands as key-habitats for biodiversity conservation in Mediterranean landscapes: a case study using rove and ground beetles (Coleoptera: Staphylinidae, Carabidae)

Land-use intensification in Mediterranean agro-forest systems became a pressure on biodiversity, concerning particularly the woodland sensitive species. In 2001, the effects of a land-use gradient from old-growth cork-oak forest to a homogeneous agricultural area were assessed using rove beetles as indicators in a Mediterranean landscape. The aim was to find which species were negatively affected by land-use intensification at the landscape level and whether they benefited from cork-oak patches occurring along the land-use gradient. A total of 3,196 rove beetles from 88 taxa were sampled from all landscape types. Agricultural area recorded significantly higher numbers of abundance and species richness in relation to the cork-oak mosaics, i.e. the old-growth forest and the managed agro-forest landscapes (montados). Moreover, 70% of rove beetle indicator species common enough to be tested by IndVal displayed their highest indicator value for agriculture, showing a lower number of woodland indicators in comparison to ground beetles. Nevertheless, one rove beetle taxon was considered a specialist of closed woodland mosaics while no specialist ground beetle was found for that landscape typology. Some rare rove beetle species were also important in typifying diversity patterns of old-growth cork-oak forests. Hence, future management in Mediterranean landscapes should take into account not only indicator species common enough to be tested by IndVal, but also rare and endemic species. Considering the added value of cork-oak woodland cover for sensitive rove and ground beetle diversity, the strengthening of cork-oak woodland connectivity seems to be a crucial management that is required in agricultural Mediterranean landscapes.     

First fungus gnats of genus Manota Williston (Diptera: Mycetophilidae) from Japan

The genus Manota is recorded from Japan for the first time. Three new species, Manota satoyamanis, Manota indahae and Manota tunoae spp. nov., are described, based on specimens collected in an ecological sampling program of arthropods in the “satoyama” landscape of Ishikawa Prefecture. “Satoyama” represents the traditional rural landscape of Japan, which is characterized by a mosaic of secondary forests, plantations, ponds and rice paddy fields. The new species raise the number of Palearctic Manota species from five to eight.     

Interactive effects of local and landscape factors on farmland carabids

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Plants, insects and birds in semi-natural pastures in relation to local habitat and landscape factors

The preservation of remaining semi-natural grasslands in Europe has a high conservation priority. Previously, the effects of artificial fertilisation and grazing intensity on grassland animal and plant taxa have been extensively investigated. In contrast, little is known of the effects of tree and shrub cover within semi-natural grasslands and composition of habitats in the surrounding landscape on grassland taxa. We evaluated the effect that each of these factors has on species richness and community structure of vascular plants, butterflies, bumble bees, ground beetles, dung beetles and birds surveyed simultaneously in 31 semi-natural pastures in a farmland landscape in south-central Sweden. Partial correlation analyses showed that increasing proportion of the pasture area covered by shrubs and trees had a positive effect on species richness on most taxa. Furthermore, species richness of nectar seeking butterflies and bumble bees were negatively associated with grazing intensity as reflected by grass height. At the landscape level, species richness of all taxa decreased (butterflies and birds significantly so) with increasing proportion of urban elements in a 1-km² landscape area centred on each pasture, while the number of plant and bird species were lower in landscapes with large proportion of arable fields. Our results differed markedly depending on whether the focus was on species richness or community structure. Canonical correspondence analyses (CCA) showed that the abundance of most taxa was ordered along a gradient describing tree cover within pastures and proportion of arable fields in the landscape. However, subsets of grassland birds and vascular plants, respectively, showed markedly different distribution patterns along axis one of the CCA. In contrast to current conservation policy of semi-natural pastures in Sweden, our results strongly advise against using a single-taxon approach (i.e., grassland vascular plants) to design management and conservation actions in semi-natural pastures. Careful consideration of conservation values linked to the tree and shrub layers in grasslands should always precede decisions to remove trees and shrubs on the grounds of promoting richness of vascular plants confined to semi-natural grasslands. Finally, the importance of landscape composition for mobile organisms such as birds entails that management activities should focus on the wider countryside and not exclusively on single pastures.     

Forest–farm edge effects on communities of ground beetles (Coleoptera: Carabidae) under different landscape structures

In fragmented landscapes, ecological processes may be significantly influenced by edge effects, but few data are available for edge effects across forest–farmland edges. We investigated patterns of species richness, abundance, and species composition in ground beetles across forest–farm edges in two different agro-forest landscapes in Korea. Nine and five sites were selected from Hwaseong, a fragmented landscape, in 2011 and 2012, respectively, while eight sites were selected from Hoengseong, a relatively well-protected landscape, in 2012. Ground beetles were collected by pitfall trapping. Species richness was higher in the surrounding habitat than in the forest interior or edge in both Hwaseong and Hoengseong. However, in Hwaseong, species richness of the forest edge was similar to that of the forest interior, while in Hoengseong forest edge species richness was intermediate between that of the forest interior and surrounding areas. In addition, non-metric multidimensional scaling based on the combined data of both locations showed that the species composition of ground beetles in the forest edge was more similar to that of the forest interior than the surrounding areas, although some open-habitat species occurred at the forest edges. Three characteristic groups (forest specialists, edge-associated species, and open-habitat species) of ground beetle species were detected by indicator value analysis. In our study, ground beetle assemblages differed in the forest edges of two agro-forest landscapes, suggesting that the edge effect on biota can be influenced by landscape structure.     

Interacting effects of landscape context and habitat quality on flower visiting insects in agricultural landscapes

Landscape context and habitat quality may have pronounced effects on the diversity of flower visiting insects. We investigated whether the effects of landscape context and habitat quality on flower visiting insects interact in agricultural landscapes in the Netherlands. Landscape context was expressed as the area of semi-natural habitats or the density of linear landscape features, and was quantified at spatial scales ranging from 250 to 2000 m. Habitat quality was determined as flower abundance. Species richness and abundance of hoverflies and bees were determined along 16 stream banks experiencing similar environmental conditions but situated in areas with contrasting landscape context. Only flower abundance and the area of semi-natural habitats within 500–1000 m were significantly related to species richness of hoverflies and bees and these factors had interacting effects on both species groups. Our results suggest that the regional area of semi-natural habitats had a positive effect on hoverfly species richness when flower abundance was relatively high, but not when flower abundance was low. Moreover, flower abundance had positive effects on hoverfly species richness only in areas with relatively many semi-natural habitats. Contrastingly, flower abundance had a more positive effect on bee species richness in landscapes with few semi-natural habitats compared to landscapes with more semi-natural habitats. Our results suggest that the importance of landscape context for the species richness of flower visiting insects depends upon the quality of the habitat patches.     

Biodiversity in agricultural landscapes: the ground beetle communities of woody uncultivated habitats

Agricultural landscapes can be defined as mosaics of landscape elements which are affected by farming practices. Woodland habitats, even though they are managed, are amongst the most stable elements of agricultural landscapes and can play a key role in the maintenance of biodiversity. This study of the ground beetle (carabid) communities of woodlands and woody linear features in a Scottish agricultural landscape shows that these habitats contribute significantly to the overall landscape diversity of these beetles. Communities in woods and hedgerows display the same species diversity and are both characterized by the presence of forest species. The main factors constraining carabid communities in both environments are the grazing intensity and, to a lesser extent, the type of soil. Heavily grazed locations are characterized by the occurrence of grassland species while forest species are restricted to ungrazed locations. At the landscape scale, the distribution of the forest species is limited by spatial isolation, indicating that there are insufficient functional links between woodland habitats in the study area. Isolation could be compensated for either by a better control of grazing so that linear features can be used as dispersal corridors for forest carabids or by planting more linear features and woods in the area.     

The spatial distribution of birds and carabid beetles in pine plantation forests: the role of landscape composition and structure

Aim To evaluate the joint and independent effects of spatial location, landscape composition and landscape structure on the distribution patterns of bird and carabid beetle assemblages in a mosaic landscape dominated by pine plantation forests. Location A continuous 3000‐ha landscape mosaic with native maritime pine Pinus pinaster plantations of different ages, deciduous woodlands and open habitats, located in the Landes de Gascogne forest of south‐western France. Methods We sampled breeding birds by 20‐min point counts and carabid beetles by pitfall trapping using a systematic grid sampling of 200 points every 400 m over the whole landscape. Explanatory variables were composed of three data sets derived from GIS habitat mapping: (1) spatial variables (polynomial terms of geographical coordinates of samples), (2) landscape composition as the percentage cover of the six main habitats, and (3) landscape structure metrics including indices of fragmentation and spatial heterogeneity. We used canonical correspondence analysis with variance partitioning to evaluate the joint and independent effects of the three sets of variables on the ordination of species assemblages. Moran's I correlograms and Mantel tests were used to assess for spatial structure in species distribution and relationships with separate landscape attributes. Results Landscape composition was the main factor explaining the distribution patterns of birds and carabids at the mesoscale of 400 × 400 m. Independent effects of spatial variables and landscape structure were still significant for bird assemblages once landscape composition was controlled for, but not for carabid assemblages. Spatial distributions of birds and carabids were primarily influenced by the amount of heathlands, young pine plantations, herbaceous firebreaks and deciduous woodlands. Deciduous woodland species had positive responses to edge density, while open habitat species were positively associated with mean patch area. Main conclusions Forest birds were favoured by an increase in deciduous woodland cover and landscape heterogeneity, but there was no evidence for a similar effect on carabid beetles. Fragmentation of open habitats negatively affected both early‐successional birds and carabids, specialist species being restricted to large heathlands and young plantations. Several birds of conservation concern were associated with mosaics of woodlands and grasslands, especially meadows and firebreaks. Conserving biodiversity in mosaic plantation landscapes could be achieved by the maintenance of a significant amount of early‐successional habitats and deciduous woodland patches within a conifer plantation matrix.     

Influence of within‐field position and adjoining habitat on carabid beetle assemblages in winter wheat

• 1 The influence of within‐field position and adjoining habitat on carabid beetles was studied in 20 winter wheat fields in ten different Swiss agricultural landscapes. In each landscape, two winter wheat fields (one with adjoining sown wildflower area and one with adjoining grassy margin) were investigated.
• 2 Carabid beetles were caught in pitfall traps 3 and 30 m from the edge in each of the 20 wheat fields. Significantly more individuals were found in the centres (30‐m position) than at the edges (3‐m position). Conversely, species richness was significantly higher at the field edges than in the centres.
• 3 Of the ten most abundant species, Poecilus cupreus, Agonum muelleri and Pterostichus melanarius were significantly more abundant in the field centres than at the edges. Harpalus rufipes was significantly more abundant in the fields adjoining sown wildflower areas than in the fields adjoining grassy margins.
• 4 In conclusion, the response of carabid beetles to within‐field position and adjoining habitats was species specific. This needs to be taken into account in habitat management for biodiversity conservation and pest control.

     

Temporal variability of aphid biological control in contrasting landscape contexts

Landscape complexity may provide ecosystem services to agriculture through the provision of natural enemies of agricultural pests. Strong positive effect of adjacent semi-natural habitats on natural enemies in croplands has been evidenced, but the resulting impact on biological control remains unclear. Taking into account the temporal dynamics of pest and natural enemies in agricultural landscapes provides better resolution to the studies and better understanding of the biological control service.In this study, the population dynamics of aphids and two groups of predators (coccinellid and carabid beetles) were examined. Insects were sampled in 20 wheat fields, surrounded by structurally simple and complex landscapes in Chilean central valley. Considering the whole sampling period, the diversity of aphids and natural enemies were similar in wheat crops surrounded by both types of landscapes, and the abundance of ladybirds was higher in crops in the complex landscapes. The dynamics of predators was more advanced in complex landscapes than in the simple ones, whereas the dynamics of aphids were similar in both types of landscape. Negative correlation between abundance of predators and aphid population growth rate in both landscape contexts were observed suggesting a control of the pest population by the predators. Different temporal patterns were observed in these correlations in the two landscape contexts, which suggests differences in the biological control related to the landscape composition.The present study shows that colonization of crops by natural enemies occurs sooner in structurally complex landscapes and suggests that this early colonization may facilitate an early and efficient control of aphid populations, nevertheless the biological control efficiency seems to be higher in structurally simple landscapes later in the season.     

植被边界带对相邻麦田地表步甲和蜘蛛分布及蚜虫发生的影响

通过管理半自然生境提高农田中天敌多样性及农田害虫的生物控制效率是当前生态农业研究的基本问题之一。为探讨不同类型半自然植被边界带对相邻麦田地表天敌的发生以及对小麦蚜虫生物控制潜力的影响,在北京顺义区赵全营镇采用陷阱法调查了不同类型植被边界带（人工林地、人工种植草带、自然演替草带）相邻的小麦田中,距离边界0,15 m和30 m处步甲和蜘蛛两类重要地表天敌的多度分布及小麦田蚜虫发生情况,同时分析了边界带植被群落结构对步甲和蜘蛛群落分布的影响。结果表明：人工林地相邻的小麦田中维持了显著较高的步甲、蜘蛛和蚜虫发生密度;不同类型植被边界带相邻小麦田中步甲和蜘蛛群落组成显著不同,植被边界带宽度以及乔木层盖度是影响相邻麦田步甲和蜘蛛群落分布的主要因子;距离植被边界带不同取样界面处,不同迁移方向的步甲和蜘蛛的活动密度无显著性差异;与林地相邻的麦田中,蜘蛛活动密度从农田边界向农田内部处依次降低,与人工种植草带以及自然演替草带相邻的麦田中,农田边界处蜘蛛的活动密度显著高于农田内部。研究显示,半自然植被边界带与麦田交界处维持了较高的蜘蛛活动密度,并具有向农田内部扩散的趋势,然而,并非高天敌密度就意味着低害虫发生率,深入研究天敌群落与害虫发生关系及其影响因素是未来天敌保护和害虫生物防治的重要方面。同时,因为不同类型植被边界带维持的天敌群落具有高度异质性,未来的研究需要充分考虑不同类型植被边界带对天敌维持的作用,以及不同类型生境界面地表天敌的活动规律。     

Comparative analysis of two edaphic zoocoenoses (Oribatid mites and Carabid beetles) in five habitats of the ‘Pietraporciana’ and ‘Lucciolabella’ Nature Reserves (Orcia Valley,central Italy)

Oribatid mite and Carabid beetle communities were investigated at five sites in the ‘Pietraporciana’ and ‘Lucciolabella’ Nature Reserves (central Italy). In this part of southern Tuscany many attempts have been made to encourage the regeneration of native habitats and to preserve existing ones. Human-induced changes in the original forest landscape have had a direct impact on mite and carabid populations. Significant differences in species diversity and abundance among different sites were revealed throughout the sampling period. Species richness, abundance and diversity of oribatid mites decrease from woodland sites to open habitats where evenness was high. There is an inverse trend between the number of species and richness of carabid beetles and those of oribatid mites. Canonical correspondence analysis (CCA) of oribatid and carabid beetle compositions discriminated the sites, demonstrating how even small areas with different vegetation, composition, structure, environment and microclimate were characterised by distinct edaphic populations.     

北京东北旺农田景观步甲群落结构的时空动态比较

2000年5～10月在北京海淀区东北旺实验基地,采用陷阱法对林地、农田边界和农田3类生境共14个处理的步甲进行了取样。并在此基础上分析了不同生境和不同季节步甲科昆虫群落的动态分布特征。结果表明,林地和农田边界类生境相对于农田类生境拥有更多的步甲个体数和物种数;无论是优势种还是稀有种都趋向于在林地和边界处生活;边界对相邻农田步甲群落的多样性有积极影响;对边界进行适度的干扰(秋季翻耕)有利于提高步甲群落的个体数量;农田生境中灌溉、施肥和秸秆还田对农田中步甲群落的个体数量和物种数量的空间分布均无显著影响。此外,还结合步甲的时空分布特性对步甲受威胁状况进行了初步的分析。