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.     

紫胶林-农田复合生态系统地表甲虫多样性——以云南绿春为例

为揭示紫胶林-农田复合生态系统地表甲虫多样性, 于2006-2007年在云南省绿春县牛孔乡采用陷阱法调查了天然紫胶林、人工紫胶林和旱地的地表甲虫群落。共采集标本1 678头, 分别隶属于24科120种, 其中步甲科(Carabidae)和金龟科(Scarabaeidae)种类最丰富, 均占全部种类的12.50%。拟步甲科(Tenebrionidae)个体数量最丰富, 占个体总数的64.48%; 金龟科次之, 占个体总数的17.58%。大多数科的物种数和个体数在不同土地利用生境中的分布没有显著差异, 而步甲科、隐翅甲科(Staphylinidae)、叩甲科(Elateridae)、拟步甲科、瓢虫科(Coccinellidae)和小蠧科(Scolytidae)在不同土地利用生境中的分布有显著差异。天然紫胶林地表甲虫个体数量最少, 物种较丰富, 优势度最低, 多样性最高; 人工紫胶林个体数和物种数均丰富, 优势度和多样性居中; 旱地个体数量最多, 物种最贫乏, 优势度最高, 多样性最低。种级水平的聚类分析体现出人工紫胶林与旱地之间在种类组成上距离较近; 而科级水平的聚类反映出人工紫胶林和天然紫胶林更接近。结果提示, 紫胶林-农田复合生态系统具有区域内土地利用方式多样化的特点; 天然紫胶林在维持地表甲虫多样性水平上具有重要作用, 而人工紫胶林虽具有积极作用, 但仍需进一步恢复。     

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。     

Moth species richness,abundance and diversity in fragmented urban woodlands: implications for conservation and management strategies

Urban expansion threatens global biodiversity through the destruction of natural and semi-natural habitats and increased levels of disturbance. Whilst woodlands in urban areas may reduce the impact of urbanisation on biodiversity, they are often subject to under or over-management and consist of small, fragmented patches which may be isolated. Effective management strategies for urban woodland require an understanding of the ecology and habitat requirements of all relevant taxa. Yet, little is known of how invertebrate, and in particular moth, assemblages utilise urban woodland despite being commonly found within the urban landscape. Here we show that the abundance, species richness, and species diversity of moth assemblages found within urban woodlands are determined by woodland vegetation character, patch configuration and the surrounding landscape. In general, mature broadleaved woodlands supported the highest abundance and diversity of moths. Large compact woodlands with proportionally less edge exposed to the surrounding matrix were associated with higher moth abundance than small complex woodlands. Woodland vegetation characteristics were more important than the surrounding landscape, suggesting that management at a local scale to ensure provision of good quality habitat may be relatively more important for moth populations than improving habitat connectivity across the urban matrix. Our results show that the planting of broadleaved woodlands, retaining mature trees and minimising woodland fragmentation will be beneficial for moth assemblages.     

Distribution of ground-dwelling beetles （Coleoptera） across a forest-clearcut ecotone in Wolong Natural Reserve, southwestern China

Abstract The influence of edge effect on ground-dwelling beetles (Coleoptera) across a forest-clearcut ecotone was studied in Wolong Natural Reserve, southwestern China. During the field research, a total of 30 739 beetles were collected with pitfall traps along transects, which extending 100 m from the edge into the forest interior and 100 m into the clearcut. Of the collection, Carabidae comprised 92%, Staphylinidae 3%, Curculionidae 2%, and Tenebrionidae 2%, and these four families can be considered as abundant groups. Family richness, Shannon diversity and equitability display a significant decrease from forest interior, edge to clearcut. Based on the family composition and abundance, ground-dwelling beetles of the forest interior can be separated from those in the clearcut by Principal coordinate analysis ordination, and beetle assemblages in the forest edge were more similar to forest assemblages than to those found in the clearcut by cluster analysis. Seasonal dynamics of family richness showed a monotone peak in the middle season, with a highest value in the forest interior and a lowest value in the clearcut. Family abundance showed two peaks in the middle season, always with more individuals in the clearcut than in the forest interior or in the edge. Multiple linear regression analyses showed that the cover of shrub and litter were the two most important factors in determining family richness, Shannon diversity, equitability and abundance. Our results show that the forest edge and clearcut have obviously different composition and diversity of ground-dwelling beetles from forest interior at the family level. However, more edges have been formed due to increasing forest fragmentation (clearcutting or logging), so it is necessary to preserve large and intact forest to protect the diversity of ground-dwelling beetles in Wolong Natural Reserve.     

Contrasting diversity dynamics of phoretic mites and beetles associated with vertebrate carrion

Carrion is an ephemeral and nutrient-rich resource that attracts a diverse array of arthropods as it decomposes. Carrion-associated mites often disperse between animal carcasses using phoresy, the transport of one species by another. Yet few studies have contrasted the dynamics of mite assemblages with other insect taxa present at carrion. We examined and compared the changes in abundance, species richness and composition of mite and beetle assemblages sampled at kangaroo carcasses in a grassy eucalypt woodland at four different times over a 6-month period. We found that the majority of mites were phoretic, with the mesostigmatid genera Uroseius (Uropodidae), Macrocheles (Macrochelidae) and Parasitus (Parasitidae) the most abundant taxa (excluding astigmatid mites). Abundance and richness patterns of mites and beetles were very different, with mites reaching peak abundance and richness at weeks 6 and 12, and beetles at weeks 1 and 6. Both mites and beetles showed clear successional patterns via changes in species presence and relative abundance. Our study shows that mesostigmatid mite assemblages have a delay in peak abundance and richness relative to beetle assemblages. This suggests that differences in dispersal and reproductive traits of arthropods may contribute to the contrasting diversity dynamics of carrion arthropod communities, and further highlights the role of carrion as a driver of diversity and heterogeneity in ecosystems.     

宁夏黄土丘陵区不同生态恢复生境地表甲虫多样性

为了解宁夏黄土丘陵区不同生境地表甲虫群落多样性变化规律及与环境因子的关系, 并探讨不同生态恢复措施对维持地表甲虫群落多样性的影响, 2013年7-8月, 作者利用陷阱法调查了该区6种生境内的地表甲虫群落多样性。结果表明: 灌草混交林地、乔灌混交林地、生态薪炭林地中地表甲虫物种丰富度和个体数量均较高, 天然封育草地、生态经济林地和水平农田中物种丰富度较低, 生态经济林地和水平农田中甲虫个体数量显著高于天然封育草地。不同生境间, 灌草混交林地、乔灌混交林地与生态薪炭林地之间、生态经济林地与水平农田之间甲虫群落组成相似性较高。多元回归分析表明, 草本层生物量、灌木层盖度及土壤含水量是影响甲虫物种丰富度的决定因素, 林冠层盖度和枯落物厚度是决定地表甲虫个体数量的重要因素。CCA分析表明, 枯落物盖度、枯落物厚度、林冠层盖度及草本层盖度是影响地表甲虫群落组成的重要环境因子。研究表明, 灌草混交林地为地表甲虫群落多样性维持较好的生境类型, 是宁夏黄土丘陵区典型生态恢复的最优模式。     

Forest complexity drives dung beetle assemblages along an edge-interior gradient in the southwest Amazon rainforest

1. The habitat heterogeneity hypothesis predicts that heterogeneous habitats may provide more niches and diverse ways of exploiting environmental resources, thereby allowing more species to coexist, persist and diversify. 2. We aimed to investigate how an edge-interior gradient related to forest complexity influences species composition, abundance and richness of dung beetles in the western Amazon rainforest. We expected dung beetle abundance and richness to increase along the forest edge-interior gradient, in accordance with the habitat heterogeneity hypothesis. We also expected strong changes in species composition driven by species turnover in the forest interior and nestedness along the forest edges. We sampled dung beetles using baited pitfall traps across an edge-interior gradient. We also assessed the variation in forest features along the edge-interior gradient to identify changes in forest complexity. 3. Both species richness and abundance of dung beetles increased along the forest edge-interior, following the gradient of forest complexity. The Sorensen dissimilarity of dung beetle assemblages was higher among sampling units placed near the forest edge, although neither turnover, nor nestedness was different between the extremes of the forest edge-interior gradient. There was a clear compositional change along the edge-interior gradient mostly driven by species turnover. Individual indicator value analysis revealed that species were strongly associated with the forest interior conditions. 4. The simplification of the Amazon rainforest near clearings causes compositional changes in dung beetle assemblages. These changes are characterised by species-poor and low-abundance assemblages and may impair dung beetle ecological functions and therefore forest recovery.     

边缘效应对卧龙自然保护区森林-草地群落交错带地表甲虫多样性的影响

本文研究了卧龙国家自然保护区地表甲虫群落在天然落叶阔叶林森林边缘与森林内部和周围草地间多样性差异,在科级水平上探讨边缘效应对地表甲虫群落的影响。调查共设5个重复样带（间距大于500 m）;每个样带以距离梯度（25 m）的方式设置样点,分别由边缘深入到森林内部和草地中央100 m,共设45个样点,通过巴氏罐诱法调查地表甲虫群落组成和季节变化。本研究采集甲虫标本4 736 号,隶属于28个科,步甲科、隐翅虫科和叩甲科分别占总数的49.5%、23.5%和13.0%,共同构成本研究地区地表甲虫的优势类群。甲虫的个体数量从森林内部、边缘到周围草地依次降低,而科多样性和均匀度则依次增高,都达到了显著差异。主坐标分析（PCoA）排序表明,森林内部和周围草地间的地表甲虫群落组成差异较大;而森林边缘的群落组成与两者都有较高程度的相似性,反映了森林边缘的地表甲虫群落已经与森林内部的群落组成发生明显分化,除了森林物种成分外,还包含了若干从周围草地环境扩散来的物种成分。从季节动态看,森林边缘和森林内部在丰富度和个体数量的季节变化曲线方面相似性更高;捕食类和腐食类地表甲虫的季节变化在森林内部和边缘相似性更高,而植食类则在森林边缘和草地的相似性更高。多元回归分析表明,枯落物覆盖率是影响地表甲虫科多样性和均匀度的决定因素,枯落物厚度和林冠层覆盖率是决定甲虫个体数量分布的重要因素。以上结果表明,在科级水平上,地表甲虫群落在天然落叶阔叶林边缘已经与森林内部生境发生分化,而且正在加剧的森林片断化进程将会产生更多的森林边缘,因此,保留大面积的天然落叶阔叶林免受破坏和干扰是保护地表甲虫多样性的必要措施。     

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.     

Changes in Brazilian Drosophilidae (Diptera) assemblages across an urbanisation gradient

The present study aims to characterise the assemblages of Drosophilidae sampled from four sites subject to different levels of urbanisation, in the city of Florianópolis, South Brazil. Five samples were collected between August 2000 and August 2001 and five between August 2002 and August 2003 in each site. A gradual change in the dominant species was observed in the urbanisation gradient, with an increase in relative abundance of exotic species in urban areas. Among the native species, the relative abundance of Drosophila capricorni Dobzhansky & Pavan had an expressive decrease. In the other hand the richness in the low urbanisation site increased, probably due to the presence of urban gardens. The diversity index did not vary significantly among sites, but its correlation with evenness and richness changed.     

Heterogeneous patterns of abundance of epigeic arthropod taxa along a major elevation gradient

Species diversity is the variable most commonly studied in recent ecological research. Ecological processes, however, are driven by individuals and affected by their abundances. Understanding the variation in animal abundances along climatic gradients is important for predicting changes in ecosystem processes under global warming. High abundances make arthropods, despite their small body sizes, important actors in food webs, yet abundance distributions of major arthropod taxa along climatic gradients remain poorly documented. We sampled arthropod assemblages in disturbed and undisturbed vegetation types along an elevational gradient of 860–4550 m asl on the southern slopes of Mt. Kilimanjaro, Tanzania. In our analysis, we focused on 13 taxa of arthropods that represented three major functional groups: predators, herbivores, and decomposers. Abundance patterns were unimodal for most of the taxa and functional groups, including decomposer arthropods, and most of them peaked at low elevations in lower montane forest. When we assigned beetles to functional groups, however, decomposer beetle abundances declined almost linearly, and abundances of predator beetles (ca. 2400 m asl) and herbivore beetles (ca. 3000 m asl, undisturbed vegetation) peaked at higher elevations and exhibited unimodal patterns. Temperature, not primary productivity, was the best predictor of abundance for most of the taxa and groups. Disturbance was only of minor importance. Our results revealed different trends in the response of arthropod abundance along the elevational gradient that depended on the level of taxonomic and functional resolution. This highlights the need for more comparisons of different taxa along the same climatic gradients.     

Changes in arthropod diversity along a land use driven gradient of shrub cover in savanna rangelands: identification of suitable indicators

Shrub encroachment linked to heavy grazing has dramatically changed savanna landscapes, and is a major form of rangeland degradation. Our understanding of how shrub encroachment affects arthropod communities is poor, however. Here, we investigate the effects of shrub encroachment on abundance and diversity of ground-dwelling (wingless) arthropods at varying levels of shrub cover in the southern Kalahari. We also ascertain if invertebrate assemblage composition changes with habitat structure and identify which aspects of habitat structure (e.g., grass cover, herbaceous plant cover, shrub density) correlate most strongly with these changes. Ant, scorpion and dung beetle abundance increased with shrub cover, whereas grasshoppers and solifuges declined. Spider and beetle abundance exhibited hump-shaped relationships with shrub cover. RTU richness within orders either mirrored abundances, or exhibited no trend. Shrub density was the habitat component most correlated with similarities between invertebrate assemblages. Ground-dwelling arthropods showed clear shifts in species assemblage composition at a similarity level of 65% according to shrub density. Changes in indicator species showed that within the Tenebrionidae (darkling beetles), certain species respond positively to shrub thickening, replacing other species within the Family. Small-bodied, wingless Scarabaeidae (dung beetles) tended to increase with increased shrub density and three species emerged as significant indicators of more thickened habitats, although this might be a response to greater dung availability, rather than habitat structure itself. We conclude that because ground-dwelling invertebrates showed such clear responses in species assemblage composition, they present excellent candidates for use as indicator species in further studies into bush encroachment.     

Changes in carabid beetle assemblages across an urban-rural gradient in Japan

As part of the international Globenet project, carabid beetles (Coleoptera, Carabidae) were collected using pitfall traps from four urban, four suburban and four rural sites in Hiroshima City, Japan, during the 2001 summer season. In agreement with expectation, carabid abundance and species richness decreased significantly from rural to urban sites. Furthermore, no large, and only few individuals of medium-sized specialist species were collected from the urban environment, while many specimens of medium-sized and some large-sized specialist species were collected from the suburban and rural sites. Hiroshima city was characterised by medium-sized generalist carabids, while the suburbs and the rural environments were characterised by small-sized generalist beetles. These results did not apply at the species level. To summarise, we found a significant effect of urbanisation on the composition of carabid beetle assemblages in Hiroshima City. These changes were similar to those found in previous studies performed in Sofia (Bulgaria), Edmonton (Canada) and Helsinki (Finland). Thus, it appears that urbanisation has some similar and predictable effects on carabid assemblages in various parts of the world.     

Carabid Beetle and Spider Assemblages along a Forested Urban–Rural Gradient in Southern Finland

To investigate the effects of urbanization on carabid beetles (Carabidae) and ground dwelling spiders (Araneae) a study was completed along a 20km urban–rural forest gradient in the Helsinki–Espoo area of southern Finland. To study changes in assemblage structure, abundance and species richness, these taxa were collected in the year 2000 using pitfall traps, which had been placed in four forest sites within each of the urban, suburban and rural zones. We expected to find changes in the abundances and species richnesses in the two taxa across the urban–rural gradient, but did not find any. Our second and third hypotheses, stating that generalist species and small-bodied species should gain dominance along the gradient from rural to urban sites, were partly supported as carabid specialists were more characteristic of suburban and rural environments whereas generalists were more likely to be collected from rural areas compared to suburban or urban sites. Furthermore, medium to large-sized carabid individuals were more likely to be collected in the rural sites compared to urban forests. We found no evidence for significant changes in spider abundance or species richness across the urban–rural gradient in relation to body size or habitat specialization. We suggest that urbanization does not have significant effects on the total abundances and species richnesses in these two taxa. However, individual species responded differently to urbanization, and there were significant differences in the specialization and body sizes of carabids across the gradient.     

大巴山北坡地表甲虫的物种多样性

采用巴氏罐诱法取样,对大巴山北坡3种主要林型（林区人工林、农耕区天然次生林和农耕区人工林）地表甲虫的类群组成、数量分布进行了分析,并采用对应趋势分析和聚类分析法,研究了3种林型地表甲虫类群组成和数量分布的相似性.结果表明：2002年8月9—13日,研究区共捕获地表甲虫1094号,其中,隐翅虫科和叶甲科分别占40.59%和33.18%,步甲科、球蕈甲科、金龟科、象甲科、蚁甲科和缨甲科均各占1%以上,这8科地表甲虫是该地区常见的类群;地表甲虫的个体数量从林区人工林、农耕区天然次生林到农耕区人工林依次减少;农耕区人工林的科丰富度指数明显低于林区人工林和农耕区天然次生林.大巴山北坡农耕区天然次生林地表甲虫的类群组成和数量分布与林区人工林差异明显,农耕区人工林与林区人工林和农耕区天然次生林各有部分相似性.     

Arthropod responses to experimental fire regimes in an Australian tropical savannah: ordinal‐level analysis

Fire is widely used for conservation management in the savannah landscapes of northern Australia, yet there is considerable uncertainty over the ecological effects of different fire regimes. The responses of insects and other arthropods to fire are especially poorly known, despite their dominant roles in the functioning of savannah ecosystems. Fire often appears to have little long‐term effect on ordinal‐level abundance of arthropods in temperate woodlands and open forests of southern Australia, and this paper addresses the extent to which such ordinal‐level resilience also occurs in Australia’s tropical savannahs. The data are from a multidisciplinary, landscape‐scale fire experiment at Kapalga in Kakadu National Park. Arthropods were sampled in the two major savannah habitats (woodland and open forest) using pitfall traps and sweep nets, in 15–20 km² compartments subjected to one of three fire regimes, each with three replicates: ‘early’ (annual fires lit early in the dry season), ‘late’ (annual fires lit late in the dry season), and ‘unburnt’ (fires absent during the five‐year experimental period 1990–94). Floristic cover, richness and composition were also measured in each sampling plot, using point quadrats. There were substantial habitat differences in floristic composition, but fire had no measured effect on plant richness, overall composition, or cover of three of the four dominant species. Of the 11 ordinal arthropod taxa considered from pitfall traps, only four were significantly affected by fire according to repeated‐measures ANOVA . There was a marked reduction in ant abundance in the absence of fire, and declines in spiders, homopterans and silverfish under late fires. Similarly, the abundances of only four of the 10 ordinal taxa from sweep catches were affected by fire, with crickets and beetles declining in the absence of fire, and caterpillars declining under late fires. Therefore, most of the ordinal taxa from the ground and grass‐layer were unaffected by the fire treatments, despite the treatments representing the most extreme fire regimes possible in the region. This indicates that the considerable ordinal‐level resilience to fire of arthropod assemblages that has previously been demonstrated in temperate woodlands and open forests of southern Australia, also occurs in tropical savannah woodlands and open forests of northern Australia.     

Ground beetle (Coleoptera: Carabidae) assemblages associated with a satoyama landscape in Japan: the effects of soil moisture,weed height,and distance from woodlands

Agricultural landscapes generally include not only crop fields but also semi-natural habitats. In Japan, such a mixed rural landscape is called “satoyama.” Although ground beetles are potential predators of pests, the environmental factors that determine their distribution in Japanese rural landscapes have not been fully elucidated. To understand the effects of distance from woodland edges, soil moisture, and weed height on assemblages of carabid beetles, we examined the number of adult beetles in pitfall traps placed in a satoyama landscape in the lowlands of western Honshu, Japan. Our results show that the carabid species could be largely differentiated into woodland, intermediate, and open-land species. The “intermediate species” group includes species that depend on woodland or woodland edges for at least part of their life cycles. Paddy fields must have long provided semi-natural habitats that complement those in natural grasslands and wetlands for open-land beetles that prefer wet conditions. Weeds can also increase the abundance of some intermediate and woodland species; thus, the arrangement of such landscape elements as woodlands and paddies can determine the species richness and abundance of ground beetles in agricultural fields.     

Species Richness Responses to Structural or Compositional Habitat Diversity between and within Grassland Patches: A Multi-Taxon Approach

Habitat diversity (spatial heterogeneity within and between habitat patches in a landscape, HD) is often invoked as a driver of species diversity at small spatial scales. However, the effect of HD on species richness (SR) of multiple taxa is not well understood. We quantified HD and SR in a wet-dry gradient of open grassland habitats in Hortobágy National Park (E-Hungary) and tested the effect of compositional and structural factors of HD on SR of flowering plants, orthopterans, true bugs, spiders, ground beetles and birds. Our dataset on 434 grassland species (170 plants, 264 animals) showed that the wet-dry gradient (compositional HD at the between-patch scale) was primarily related to SR in orthopterans, ground-dwelling arthropods, and all animals combined. The patchiness, or plant association richness, of the vegetation (compositional HD at the within-patch scale) was related to SR of vegetation-dwelling arthropods, whereas vegetation height (structural HD at the within-patch scale) was related to SR of ground-dwelling arthropods and birds. Patch area was related to SR only in birds, whereas management (grazing, mowing, none) was related to SR of plants and true bugs. All relationships between HD and SR were positive, indicating increasing SR with increasing HD. However, total SR was not related to HD because different taxa showed similar positive responses to different HD variables. Our findings, therefore, show that even though HD positively influences SR in a wide range of grassland taxa, each taxon responds to different compositional or structural measures of HD, resulting in the lack of a consistent relationship between HD and SR when taxon responses are pooled. The idiosyncratic responses shown here exemplify the difficulties in detecting general HD-SR relationships over multiple taxa. Our results also suggest that management and restoration aimed specifically to sustain or increase the diversity of habitats are required to conserve biodiversity in complex landscapes.