Life-history trait and functional diversity patterns of ground beetles and spiders along a coastal heathland successional gradient

We investigated patterns in species richness and diversity, life-history traits and functional diversity of carabid beetles and spiders along a coastal heathland successional gradient. We sampled in five successional stages on the island of Hiddensee, Germany. Species richness of carabids and spiders and Simpson diversity for spiders did not differ among stages. Functional dispersion (FDis), a measure of functional richness, was lowest in the youngest stage for both carabids and spiders. Older successional stages represent more heterogeneous habitats and offer a broader range of niches, which might enhance functional dispersion. Functional evenness (FEve) differed among successional stages for spiders, indicating an uneven distribution of species abundances in the functional trait space. Functional divergence (FDiv) did not differ among stages for either taxon. Shifts in life-history traits were noted in both arthropod groups, but not always in the same direction: Body size of carabid beetles increased towards older successional stages, but decreased for spiders. The number of flightless carabid species increased towards older habitats. Each successional stage seemingly favors species with distinct life-history traits (species sorting). We conclude that a trait-based approach including the use of functional diversity measurements (FDis) could contribute valuable information for biodiversity conservation.     

Even the Smallest Non-Crop Habitat Islands Could Be Beneficial: Distribution of Carabid Beetles and Spiders in Agricultural Landscape

Carabid beetles and ground-dwelling spiders inhabiting agroecosystems are beneficial organisms with a potential to control pest species. Intensification of agricultural management and reduction of areas covered by non-crop vegetation during recent decades in some areas has led to many potentially serious environmental problems including a decline in the diversity and abundance of beneficial arthropods in agricultural landscapes. This study investigated carabid beetle and spider assemblages in non-crop habitat islands of various sizes (50 to 18,000 square metres) within one large field, as well as the arable land within the field, using pitfall traps in two consecutive sampling periods (spring to early summer and peak summer). The non-crop habitat islands situated inside arable land hosted many unique ground-dwelling arthropod species that were not present within the surrounding arable land. Even the smallest non-crop habitat islands with areas of tens of square metres were inhabited by assemblages substantially different from these inhabiting arable land and thus enhanced the biodiversity of agricultural landscapes. The non-crop habitat area substantially affected the activity density, recorded species richness and recorded species composition of carabid and ground-dwelling spider assemblages; however, the effects were weakened when species specialised to non-crop habitats species were analysed separately. Interestingly, recorded species richness of spiders increased with non-crop habitat area, whereas recorded species richness of carabid beetles exhibited an opposite trend. There was substantial temporal variation in the spatial distribution of ground-dwelling arthropods, and contrasting patterns were observed for particular taxa (carabid beetles and spiders). In general, local environmental conditions (i.e., non-crop habitat island tree cover, shrub cover, grass cover and litter depth) were better determinants of arthropod assemblages than non-crop habitat island size, indicating that the creation of quite small but diversified (e.g., differing in vegetation cover) non-crop habitat islands could be the most efficient tool for the maintenance and enhancement of diversity of ground-dwelling carabids and spiders in agricultural landscapes.     

不同土地利用方式对河北坝上步甲物种多样性的影响

为了探讨土地利用方式对步甲物种多样性的影响及其与环境因子的关系, 于2009年5-10月利用巴氏罐法, 对河北坝上塞北管理区内的灌草丛、马铃薯地、苜蓿地、围封草地、中度放牧草地和重度放牧草地6种生境的步甲进行了系统调查。结果表明: （1）共捕获步甲标本10 804号, 计10属18种, 其中强足通缘步甲Pterostichus fortipes、侧步甲Carabus latreillei和双斑猛步甲Cymindis binotata占优势, 个体数分别占总数的67.5%, 7.53%和7.06%。（2）苜蓿地中的步甲密度、物种丰富度、Margalef丰富度指数、Shannon-Wiener多样性指数和均匀度指数均较高, 其他样地较低。（3）聚类分析结果显示, 不同生境的步甲群落相似性与受干扰强度和土地利用方式一致。（4）步甲主要在6-9月活动, 高峰期为7-8月。各物种在不同土地利用方式生境内的季节变化趋势与不同生境植被生长季节相关, 步甲个体数量随着植物生物量的增加而增加。强足通缘步甲Pterostichus fortipes高峰期大多出现在植物生长旺盛的7月, 仅围封草地内的高峰出现在8月; 双斑猛步甲Cymindis binotata的高峰期出现在8-9月。（5）典范对应分析表明, 土壤含水量、土壤温度、植物盖度、植物生物量及植株高度是影响步甲群落组成的重要环境因子。结论认为, 不同土地利用方式对环境的干扰影响了步甲赖以生存的生境, 进而影响了其物种的组成和数量分布格局。     

Environmental and historical effects on richness and endemism patterns of carabid beetles in the western Palaearctic

Identifying determinants of spatial diversity gradients is in the focus of biodiversity-related research and has gained considerable importance regarding global change and conservation strategies. Despite their overwhelming diversity and the crucial role of invertebrates in ecosystem functioning, our understanding of factors driving broad-scale invertebrate diversity is limited. Tackling this issue, our study analyzes macroecological patterns of a highly diverse insect taxon across large parts of the western Palaearctic. We used regression modelling to assess the influence of environmental factors on overall, widespread and restricted-range (endemic) carabid beetle diversity. Single-term regressions and variation partitioning among climatic, topographic and spatial variables showed that total carabid diversity as well as richness patterns of widespread species were most strongly correlated with spatially structured variables related to current climate (measures of ambient energy and, to a lesser degree, precipitation and AET). In contrast, restricted-range (endemic) species were most notably related to range in elevation. We discuss the possible role of this factor as a surrogate measure of historical processes and the impact of history on contemporary diversity distributions. Our results indicate that while overall diversity patterns of carabids strongly reflect current climate conditions, this primarily reflects the more widespread species, whereas the spatial distribution of restricted-range species is still significantly affected by historical processes. Thus, for a general understanding of determinants and mechanisms of broad-scale diversity, taking into account dispersal abilities and range sizes of species is essential, especially as large parts of global biodiversity are represented by invertebrates with low dispersal powers.     

Ground beetle (Coleoptera: Carabidae) assemblages in conventional and diversified crop rotation systems

Ground beetles (Coleoptera: Carabidae) are important in agro-ecosystems as generalist predators of invertebrate pests and weed seeds and as prey for larger animals. However, it is not well understood how cropping systems affect ground beetles. Over a 2-yr period, carabids were monitored two times per month using pitfall traps in a conventional chemical input, 2-yr, corn/soybean rotation system and a low input, 4-yr, corn/soybean/triticale-alfalfa/alfalfa rotation system. Carabid assemblages were largely dominated by a few species across all cropping treatments with Poecilus chalcites Say comprising >70% of pitfall catches in both years of study. Overall carabid activity density and species richness were higher in the low input, 4-yr rotation compared with the conventionally managed, 2-yr rotation. There were greater differences in the temporal activity density and species richness of carabids among crops than within corn and soybean treatments managed with different agrichemical inputs and soil disturbance regimes. Detrended correspondence analysis showed strong yearly variation in carabid assemblages in all cropping treatments. The increase in carabid activity density and species richness observed in the 4-yr crop rotation highlights the potential benefits of diverse crop habitats for carabids and the possibility for managing natural enemies by manipulating crop rotations.     

Effects of urbanization on ground beetle assemblages (Coleoptera, Carabidae) of grassland habitats in western Canada

We pitfall-trapped carabid beetles across an urban-rural gradient in central Alberta, Canada to assess the relative contributions of landscape structure and habitat quality in explaining variation in composition of assemblages. The gradient ranged from the "urban" zone in the city of Edmonton, through a "suburban" zone just outside the city, into a "rural" zone 25–60 km east of the city. We trapped beetles at three replicated sites for each of two habitats (unmanaged grasslands and highly managed graveyards) in each of the three zones. Most of the 24 species (3162 individuals) captured were commonly associated with open habitats. Pterostichus melanarius , an introduced species, made up 80% of the total catch. Carabid assemblages of grasslands and graveyards were distinct, with lower species richness and lower catch rates of native, introduced and brachypterous carabids in graveyards. Graveyard assemblages also had proportionally fewer brachypterous carabid individuals (32 vs 52%) and species (20 vs 36%), suggesting that populations recruit locally from elsewhere after disturbances. Native carabids were least abundant in the urban areas. Brachypterous carabids were similarly abundant across the gradient, but more occurred in the urban zone than elsewhere (32 vs 5–10%). Samples from graveyards were characterized by lower beta diversity (variation among samples) than those from grasslands, but beta diversity was more-or-less equally variable across the gradient within habitats. The amount of unmanaged grassland habitat and the degree of urbanization significantly influence carabid assemblages, and therefore are relevant to management and conservation of urban grassland areas.     

Useful model organisms,indicators, or both? Ground beetles (Coleoptera,Carabidae) reflecting environmental conditions

Classic studies have successfully linked single-species abundances, life-history traits, assemblage structures and biomass of carabid beetles to past and present, human-caused environmental impacts and variation in ‘natural’ conditions. This evidence has led many to suggest carabids to function as ‘indicators’ − a term that bears multiple meanings. Here, a conservation-oriented definition for an indicator is used, carabid indicator potential from seven views is evaluated, and ways to proceed in indicator research are discussed. (1) Carabid species richness poorly indicates the richness and abundance of other taxa, which underlines the importance of using multiple taxa in environmental assessments. The ability of assemblage indices and specialist or functional-group abundances to reflect rare species and habitats should be examined in detail. (2) Experimental evidence suggests that carabids may potentially serve as keystone indicators. (3) Carabids are sensitive to human-altered abiotic conditions, such as pesticide use in agro-ecosystems and heavy metal contamination of soils. Carabids might thus reflect ecological sustainability and ‘ecosystem health’. (4) Carabid assemblages host abundant species characteristic of particular habitat types or successional stages, which makes them promising dominance indicators. (5) Carabids reflect variation in ‘natural’ conditions, but vegetation and structural features are more commonly adopted as condition indicators. Carabids nevertheless provide yet another, equally accurate, view on the structure of the environment. (6) Carabids may function as early-warning signalers, as suggested by recent studies linking climate and carabid distributions. (7) Carabids reflect natural and human-caused disturbances and management, but the usefulness of these responses for conservation purposes requires further research. In summary, European carabids appear useful model organisms and possibly indicators because they are diverse, taxonomically and ecologically well-known, efficiently reflect biotic and abiotic conditions, are relevant at multiple spatial scales, and are easy to collect in sufficiently large numbers to allow statistical analyses. The assumption that carabid responses would reflect rare environmental conditions or the responses of rare and threatened species ‒ crucial information for conservationists and managers ‒ has not yet been critically evaluated. Even if it holds, the usefulness will be context dependent: species and their populations vary, conditions vary, questions put forward vary, and assessment goals vary.     

Spatial distribution of insect indicator taxa as a basis for peat bog conservation planning

Peat bogs are highly endangered ecosystems worldwide. In the Western Balkans, they represent fragmented habitat patches, where insect diversity and implications for the habitat loss are poorly known. Carabid beetles and ants, widely used environmental indicators, were sampled by pitfall trapping in the largest and oldest Croatian peat bog and at its edges. We found that tyrphobiontic—bog specialist—species were absent in both taxonomic groups, which is likely due to biogeographical limitation for dispersal of these species into this region. With this study, tyrphophilous—typical of bogs—species have been recognised as environmental indicators in the Western Balkans peat bogs. Assemblages of both taxa were strongly affected by specific conditions in the peat bog – high soil moisture and Sphagnum moss dominance, showing that edges are more favourable habitats for both taxa. Overall, carabid beetle activity density was significantly higher at the edge than at the peat bog, while species richness increased for both taxa at the edge, but not significantly. Spatial distribution of tyrphophilous species differed between studied taxa, with ants being more abundant in the peat bog, and carabid beetles at the edge. Additionally, some stenotopic and globally endangered species preferred edges. Occurrence of tyrphophilous and globally endangered species highlights the great conservation value of peat bog habitats in the Western Balkans. However, successional changes in vegetation are changing hydrological conditions of the peat bog. Here we suggest active conservation measures to preserve these unique soil invertebrate zoocoenoses.     

山西庞泉沟自然保护区步甲物种组成及多样性格局

利用巴氏罐诱法,对山西庞泉沟国家自然保护区的不同功能区——核心区（绿色长廊）、缓冲区（八道沟）、远离农田实验区（神尾沟）及靠近农田实验区（八水沟）——林地地表甲虫步甲物种多样性进行了调查和比较,目的在于探明该保护区有代表性林地步甲物种的组成及受不同程度人为干扰下的多样性格局。结果表明：（1）本研究共捕获步甲物种42种,38种是该保护区的新记录种类,其中粗皱步甲Carabus crassesculptus和沟步甲C. canaliculatus 为优势种,与尖角通缘步甲Pterostichus eschscholtzii,强足通缘步甲P. fortipes,罕丽步甲C. manifestus和长叶步甲C. vladimirskyi一起构成研究地的常见种。（2）不同功能区物种组成和数量分布各不相同,多样性各项指标均是靠近农田的实验区最高,核心区最低。（3）各功能区具有特有的指示性种类,其中黄条春步甲Notiophilus reitteri为核心区指示种; 凹唇春步甲N. impressifrons 为缓冲区指示种; 粒步甲C．granulatus,刻翅步甲C．sculptipennis 和中华通缘步甲P．chinensis为实验区指示种。（4）群落相似性分析表明缓冲区与核心区及远离农田实验区之间具有较高的相似性,靠近农田实验区与其他3个地点相似性较低。结果提示适度干扰可提高物种丰富度,但降低人为干扰（如旅游和交通压力）有利于步甲多样性的保护。     

Measuring the efficiency of fen restoration on carabid beetles and vascular plants: a case study from north‐eastern Germany

The aim of the study was to assess the effects of fen rewetting on carabid beetle and vascular plant assemblages within riverine fens along the river Peene in north‐eastern Germany. Drained (silage grassland), rewetted (restored formerly drained silage grassland), and near‐natural (fairly pristine) stands were compared. Eighty‐four beetle species (7,267 individuals) and 135 plant species were recorded. The richness of vascular plant species and the number of endangered species were highest on near‐natural fens. Fourteen years of rewetting did not increase plant species numbers compared with drained fens. For carabid beetles, however, species richness and the number of stenotopic species were highest on rewetted fens. Rewetting caused the replacement of generalist carabids by wetland specialists, but did not provide suitable habitat for specialist fen carabids or for plant species of oligo‐ or mesotrophic fen communities. Therefore, raising the water table on fens with nutrient‐rich, degraded peat was not sufficient for restoring species assemblages of intact fens, although water level was the most important environmental factor separating species assemblages. Our study illustrated that insects and plants may respond differentially to restoration, stressing the need to consider different taxa when assessing the efficiency of fen restoration. Furthermore, species assemblages of intact fens could not be restored within 14 years, highlighting the importance of conserving pristine habitat.     

Distribution of carabid beetles (Coleoptera: Carabidae) across a forest-grassland ecotone in southwestern China

This paper studied the occurrence of carabid beetles (Coleoptera: Carabidae) in the forest edge, the adjacent forest interior, and the surrounding grassland in southwestern China. Beetles were collected with pitfall traps along five replicated transects. Forest species rarely penetrated into the grassland from the forest interior, and the grassland specialists were not found in the forest interior. The forest edge hosted additional species from the adjacent grassland that increased its overall species richness. Nearly all forest species (23 of 24 species) and grassland species (13 of 15 species) can be found in the forest edge. Carabids of the forest edge were more similar to those of the forest interior than to those of the grassland by ordination and cluster analysis. Based on the specificity and fidelity, carabids can be distinguished into five species groups: habitat generalists, grassland-associated species, forest generalists, forest specialists, and edge-associated species. Multiple linear regression analysis showed that canopy cover and/or shrub cover were the most important factors in determining the richness, abundance, and diversity of carabids. The forest edge may serve as a transition zone for dispersal and re-colonization of carabid beetles from adjacent habitats and therefore is important for natural conservation.     

人工固沙植被恢复对地表节肢动物群落组成及多样性的影响

人工绿洲是干旱区主要的自然景观之一,为了维护绿洲稳定而营建的防护林强烈改变地表生态水文过程,进而影响着地上和地下生物多样性。然而,以往的研究对地上植被关注较多,而对土壤动物研究较少。以张掖绿洲外围人工固沙植被群落和天然固沙植被群落为研究对象,探讨人工固沙植被恢复对荒漠地表节肢动物群落组成及多样性的影响及不同动物类群对植被变化的响应模式。研究表明,天然固沙植被群落转变为人工固沙植被群落显著降低了地表节肢动物数量,但提高了地表节肢动物类群丰富度和多样性,这在5月份表现尤为明显。植被类型对地表节肢动物群落的影响不同,8月人工柽柳林群落地表节肢动物活动密度、类群丰富度和多样性均显著高于人工梭梭林群落。人工固沙植被恢复显著降低了适应荒漠环境的拟步甲科甲虫,而蚁科和部分蜘蛛的数量显著增加,它们对人工固沙植被恢复的响应模式不同决定了群落结构及多样性的变化规律。此外,研究还发现一些地表节肢动物类群对不同生境具有强烈的指示作用,如拟步甲科等可指示荒漠生境,潮虫科等可指示梭梭林生境,而蠼螋科和狼蛛科等可指示柽柳林生境。综上所述,人工栽植固沙灌木形成的固沙植被群落导致了一些适应荒漠环境的地表节肢动物类群的数量降低,但也为更多的地表节肢动物类群提供了适应栖居环境和充足的食物资源等,从而提高了地表节肢动物的多样性。     

Impact of the invasive moss Campylopus introflexus on carabid beetles (Coleoptera: Carabidae) and spiders (Araneae) in acidic coastal dunes at the southern Baltic Sea

Campylopus introflexus is an invasive moss in Europe and North America that is adapted to acidic and nutrient-poor sandy soils with sparse vegetation. In habitats like acidic coastal dunes (grey dunes) it can reach high densities, build dense carpets and modify habitat conditions. While the impact of the moss invasion on the vegetation is well analyzed, there is a lack of knowledge regarding possible effects on arthropods. In the present study we analyzed the impact of Campylopus introflexus on the ground-dwelling arthropods carabid beetles and spiders, as both taxa are known to be useful indicator taxa even on a small-scale level. In 2009 we compared species composition in a) invaded, moss-rich (C. introflexus) and b) native, lichen-rich (Cladonia spp.) acidic coastal dunes by using pitfall traps. A total of 1,846 carabid beetles (39 species) and 2,682 spiders (66 species) were caught. Species richness of both taxa and activity densities of spiders were lower in invaded sites. Species assemblages of carabids and spiders differed clearly between the two habitat types and single species were displaced by the moss encroachment. Phytophagous carabid beetles, web-building spiders and wolf spiders were more abundant in native, lichen-rich sites. Shifts in species composition can be explained by differences in the vegetation structure, microclimate conditions and most likely a reduced food supply in invaded sites. By forming dense carpets and covering large areas, the moss invasion strongly alters typical arthropod assemblages of endangered and protected (EU-directive) acidic coastal dunes.     

Differentiation of spring carabid beetle assemblages between semi‐natural habitats and adjoining winter wheat

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How good are carabid beetles (Coleoptera,Carabidae) as indicators of invertebrate abundance and order richness?

Ground invertebrates were sampled by pitfall trapping on Brentmoor Heath, Surrey, UK during the summers of 2009 and 2010 to determine whether the abundance and species richness of carabids on bare ground patches are representative of invertebrate abundance and order richness on the same patches. Four patch sizes were investigated as well as the adjacent mown and unmown areas, and a range of environmental parameters were measured at each patch. Results show that increasing abundance and richness of carabids can give a broad indication of whether invertebrate abundance and order richness are increasing. Invertebrates in general are sensitive to a greater number of environmental variables and complex interactions than carabids are, due to the vast array of biological requirements of the large range of invertebrate species. All else being equal, smaller patches appear to benefit both carabid abundance and invertebrate abundance. Converse to the results of carabid species richness, larger patches appear to benefit invertebrate order richness. This study highlights the importance of testing the effect of habitat management strategies on a range of taxa rather than assuming the results for a bioindicator will suffice for management decisions. Whilst bioindicators may give a broad indication of effects on similar taxa, the precise nature of those effects will be determined by a number of species- and site-specific variables.     

Invasive moss alters patterns in life-history traits and functional diversity of spiders and carabids

Invasive plants can modify terrestrial habitats and affect the natural faunal composition. In acidic coastal dunes the invasive moss Campylopus introflexus can form dense carpets that largely replace native vegetation. As shown in a previous study, moss invasion affects habitat structure and ground-dwelling arthropod diversity. We suggested that including the functional diversity concept in the analysis of moss invasion impacts may offer further insights. We used pitfall trap data to compare trait composition and functional diversity of spiders and carabids in (a) invaded, moss-rich (C. introflexus) and (b) native, lichen-rich (Cladonia spp.) acidic coastal dunes. Moss invasion induced shifts in the trait values body size and feeding preference (carabids) and hunting mode (spiders): Species were smaller in native sites, and the percentages of web-building spiders and phytophagous beetles were reduced in invaded sites. Furthermore, moss invasion led to a more heterogeneous trait composition for spiders, and changed functional diversity of both arthropod groups, although with the opposite effects: While spiders were functionally more diverse in invaded sites, moss invasion reduced carabid beetles’ functional diversity. We also observed changes in the relationship between species richness and functional diversity that indicate a high functional similarity for spiders but a lower one for carabid beetles in native grey dunes. C. introflexus invasion not only alters the arthropod diversity and assemblage structure of endangered acidic coastal dunes but also interferes at a functional level. These results provide further insight into the way plant invasions might alter the structure and function of ecosystems.     

河西走廊天然固沙植被区地表甲虫多样性及其对沙漠化的指示作用

干旱、半干旱区沙漠化强烈影响动植物分布及多样性,地表甲虫是荒漠中主要的动物类群,它们对沙漠化引起的植被和土壤环境变化响应十分敏感。鉴于此,以河西走廊中部张掖绿洲外围的天然固沙植被区作为研究区,依据沙漠化发育程度选择流动沙丘（ASD）、丘间低地（IL）、半固定（SFSD）和固定沙丘（FSD）4种生境,调查了地表甲虫群落组成及影响甲虫分布的植被和土壤环境。研究发现,4种生境地表甲虫群落组成明显不同并存在季节变异,5月ASD与IL、SFSD和FSD生境地表甲虫群落的相异性大于8月。5月和8月SFSD生境地表甲虫活动密度均显著高于其他生境,8月FSD生境地表甲虫多样性指数显著高于其他生境。不同大小甲虫对沙漠化的响应模式不同,大中型甲虫对沙漠化的响应较小型甲虫敏感,这在5月表现尤为明显。地表甲虫与环境因子的RDA分析结果表明,12个植被和土壤环境因子解释了49.8%的地表甲虫群落变异,其中植被环境解释了甲虫群落变异的16.3%,土壤环境解释了甲虫群落变异的4.2%,植被和土壤环境相互作用解释了甲虫群落变异的29.3%。pRDA分析结果表明,草本物种丰富度、灌木盖度、土壤有机碳含量和粗砂含量是影响地表甲虫分布的主要环境因子,它们解释了43.7%的地表甲虫群落变异。Pearson相关分析表明,草本物种丰富度与地表甲虫活动密度呈显著正相关,而与地表甲虫均匀度呈显著负相关;灌木盖度与地表甲虫多样性呈显著正相关;地表甲虫物种丰富度与灌木盖度和草本物种丰富度均呈显著正相关。此外,研究还发现戈壁琵甲、克氏扁漠甲、中华砚甲和甘肃齿足象可以用于指示FSD生境,东鳖甲属昆虫可以用于指数SFSD生境,谢氏宽漠王可以用于指示IL及ASD生境。     

Taxon specific response of carabids (Coleoptera,Carabidae) and other soil invertebrate taxa on invasive plant Amorpha fruticosa in wetlands

False indigo (Amorpha fruticosa L.) is an invasive exotic plant introduced to Europe in the early eighteenth century. Its spread has been rapid, particularly in disturbed wetland habitats, where it forms dense impermeable monospecific stands and modifies habitat conditions. The impact of A. fruticosa on native plant communities has been well analyzed, however knowledge concerning the possible effects on soil invertebrates and particularly carabid beetles is completely lacking. This study analyzed the impact of an A. fruticosa invasion on carabid beetles and other soil invertebrates. Soil fauna was sampled by pitfall traps at natural habitats, initially colonized by A. fruticosa, and habitats largely invaded by A. fruticosa. In total 2,613 carabid beetles belonging to 50 species and 72,166 soil invertebrates were collected. The invasion of A. fruticosa strongly affected the carabid beetle species composition, which clearly differed between all studied sites. Widespread euritopic carabid beetle species showed positive responses to A. fruticosa invasion, while the activity density of open habitat species strongly declined. Mean individual biomass was significantly higher at invaded sites due to increased incidence of large carabids (genus Carabus Linné, 1758). Carabid beetle activity density and abundance of soil invertebrates were considerably higher at invaded sites than in natural sites. Conversely, the impact of A. fruticosa on carabid beetle species richness and diversity was less pronounced, most likely due to immigration from adjacent habitats. Changes in carabid beetle species composition and abundance of soil invertebrates were most likely due to changes in vegetation structure and microclimate. The results suggest that A. fruticosa invasion considerably affected carabid beetles, an insect group that is only indirectly related to plant composition. Therefore, severe future changes can be expected in invertebrate groups that are closely related to plant composition, since A. fruticosa cannot be completely removed from the habitat and covers relatively large areas.     

Habitat associations of carabid beetles in fields and forests on the Åland Islands, SW Finland

Carabid beetles (Coleoptera, Carabidae) were collected by using pitfall traps in 68 sites on the Åland Islands, SW Finland The sites were divided into four environmental types 1) cultivated fields and grazed pastures. 2) recently abandoned fields, 3) abandoned, overgrown cultivations, and 4) forests A total of 4901 carabids belonging to 77 species were caught Twenty of the most abundant species were classified into four distributional types according to their occurrence among the various environmental types 1) generalists (five species), 2) field species (five species), 3) species of open habitat (seven species), and 4) forest species (three species) Associations with environmental types were strictest among field species and forest species However, most of the species were found in low numbers outside each preferred environmental type The number of species and species diversity was highest in fields and pastures and lowest in forests, which agrees with previously detected patterns on the mainland Finland Pairwise similarity of the carabid assemblages among the environmental types was highest between the two types of abandoned cultivations The spatial distribution of two carabid species, Pterostichus melanarius and P niger , were examined in detail to ascertain whether any interspecific interaction could help explain their different island-mainland distribution observed in previous studies