Changes in carabid beetle diversity within a fragmented agricultural landscape

The distribution of carabid and cicindelid (Coleoptera: Carabidae) beetles in five distinct habitats (riparian, mature orchard, pine windbreak, young orchard, natural veld), within Tambuti Citrus Estate (Swaziland) was examined by pitfall trapping over 18 months. Habitats with high vegetation and litter cover had the highest species diversity and larger specimens, e.g. riparian border and pine windbreak, while the lowest diversity was observed in intensively managed mature citrus orchards. While species such as Tefflus delagorguei Guérin occurred in all the habitats sampled, certain species illustrated habitat specificity; e.g. Dromica ambitiosa Péringuey was observed only in the pine windbreaks while Haplotrachelus sp. Chaudoir occurred mainly in the vegetated riparian and natural veld habitats. Four unidentified carabid beetles were exclusive to the riparian border habitat. This habitat was the only one with a distinct assemblage of species in the agricultural mosaic studied. Multivariate analyses were used to assess the role of soil and environmental variables in relation to the ground beetle diversity within the agricultural mosaic studied.     

Biodiversity in agricultural landscapes: different non-crop habitats increase diversity of ground-dwelling beetles (Coleoptera) but support different communities

Agricultural intensification poses a major threat to the conservation of biodiversity and associated ecosystem services. Since non-crop habitats are regarded as important refuges for farmland biodiversity, various greening measures have been proposed to halt biodiversity loss. However, the effectiveness of these measures for biodiversity conservation is still under debate. Therefore, we here compared ground-dwelling beetle (Coleoptera) assemblages of different non-crop habitats (field margins, set-aside fields sown with wildflowers, and permanent grassland fallows) and wheat fields within an intensively used agricultural landscape in western Germany. Taxonomic diversity of Carabidae, Staphylinidae and other coleopteran families and their conservation value were higher in all non-crop habitats than on wheat fields. Surprisingly, though, different types of non-crop habitats did not differ in species richness or the number of threatened species. Thus, field margins and sown wildflower fields were as effective in promoting beetle diversity as grassland fallows. However, different non-crop habitats supported different species assemblages, and several species, in particular especially large ones, were restricted to grassland fallows. These results suggest that different greening measures are effective in promoting the biodiversity of beetles, and that permanent grassland fallows are essential for nature conservation. The fact that habitat types harbored different assemblages stresses the need to combine a variety of greening measures to yield the highest benefit for biodiversity.

     

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.     

Carrion beetles succession in three different habitats in Riyadh,Saudi Arabia

A main objective of the study is the establishment of a forensic entomological database for Riyadh, Saudi Arabia. Decomposition processes and beetle succession were analysed on rabbit carcasses in three different habitats (agricultural, desert and urban) in the period from May to July 2014. Due to the effects of the high temperature at the study sites, carrion reached the dry stage within 12 days in the agricultural habitat, and 6 days in the desert and urban habitats. A total of 125 beetles belonging to eight species and five families were collected during the decaying process, with their abundances increasing from the fresh to decay stages. The prevailing species belonged to the families of Dermestidae and Histeridae. It was not possible to confirm any definitive relationship between the occurrence of a single species and a particular stage of decomposition. The beetle communities were also not distinctively different between desert and urban habitats, but a distinct community was evident in the agriculture habitat. In addition, there were distinct beetle communities between the decay stage and the other stages. The dry stage recorded the lowest number of beetles. This study indicated that, the habitat type had an effect on the decay process and the abundance rate of the beetles.     

Distribution of ground-dwelling beetles in fragmented tropical habitats

The conservation of biodiversity is increasingly dependent on human-altered habitats. In a fragmented forest landscape in northern Costa Rica within the Mesoamerican Biological Corridor, an area of great conservation importance, we compared the diversity and composition of ground-dwelling beetle communities in five habitat types along a gradient of increasing disturbance: primary forest, logged forest, secondary forest, plantation (Gmelina arborea) and pasture. Using pitfall trapping we captured a total of 1,877 beetles (Coleoptera), comprising 422 morphospecies in 26 families. The plantation sites had the lowest number of species followed by secondary forest and pasture. Multivariate analysis separated the beetle fauna according to land use, and suggested that only the logged forest maintains a similar species assemblage to primary forest. However, each habitat harboured a number of unique species indicating the conservation value of the mosaic of habitats found in fragmented landscapes. Our results suggest that to maintain forest beetle species diversity it is essential that areas of natural forest are conserved. However, other land uses also provide beetle habitats and in fragmented areas active management to maintain a mosaic of land uses will contribute to the conservation of beetle diversity.     

小兴安岭森林地表鞘翅目成虫对不同干扰梯度的响应

为探讨小兴安岭凉水自然保护区森林生态系统中地表鞘翅目成虫群落对不同人为干扰梯度的响应,于2015年7月、8月和10月分别对轻度干扰[原始阔叶红松林(KY)和谷地云冷杉林(YL)]、中度干扰[阔叶红松择伐林(ZF)和次生白桦林(BH)]及重度干扰[落叶松人工林(RL)和红松人工林(RHS)]的6个林型进行取样调查。结果表明:(1)整个采样周期共捕获地表鞘翅目成虫879只,隶属9科44物种;其中轻度干扰生境共捕获6科29种251只(KY捕获5科21种150只,YL捕获4科20种101只),中度干扰生境捕获6科27种276只(ZF捕获3科20种144只,BH捕获6科23种132只),重度干扰生境捕获6科29种352只(RL捕获4科22种232只,RHS捕获5科17种120只)。(2)7月和8月步甲科和葬甲科占据数量优势,10月步甲科和葬甲科成虫数量明显减少而隐翅虫科数量占优势;不同林型及不同干扰梯度地表鞘翅目成虫物种总数和总个体数于7、8、10月均表现为下降趋势,且群落多样性也呈不同程度下降。(3)林型和月份对地表鞘翅目成虫群落结构具有显著影响,干扰梯度对群落结构无显著影响;6个林型之间地表鞘翅目成虫个体数量具有显著差异,但在物种组成上无显著差异;不同干扰梯度间地表鞘翅目成虫个体数、物种数无显著差异,且随干扰梯度变化没有明显的梯度性规律;相似性系数和聚类分析表明,属于同一干扰梯度的两个林型没有表现出高度的相似性。本研究表明干扰梯度不是决定各林型间地表鞘翅目成虫群落多样性存在差异的主要原因,凉水森林生态系统地表鞘翅目成虫对不同干扰梯度的响应不符合中度干扰假说,林型和时间则是影响地表鞘翅目成虫群落组成的显著因素,本实验为地表生物多样性保护和森林生态系统管理提供数据支撑。     

Composition and distribution of ground-dwelling beetles among oak fragments and surrounding pine plantations in a temperate forest of North China

In this study, we compared ground-dwelling beetle assemblages （Coleoptera） from a range of different oak fragments and surrounding conifer plantations to evaluate effects of forest size and surrounding matrix habitat in a temperate forest of north China. During 2000, beetles were sampled via pitfall traps within two large oak fragments （ca. 2.0-4.0 ha）, two small oak fragments （ca. 0.2-0.4 ha） and two surrounding matrices dom- inated by pine plantations （〉4 ha） in two sites of different aspects. Overall, no significantly negative effects from forest patch size and the surrounding matrix habitat were detected in total species number and abundance of ground-dwelling beetles. However, compared with small oak patches or pine plantations, more species were associated with an affinity for at least one large oak patch of the two aspects. Multivariate regression trees showed that the habitat type better determined the beetle assemblage structure than patch size and aspect, indicating a strong impact of the surrounding matrix. Linear mixed models indicated that species richness and abundance of all ground-dwelling beetles or beetle families showed different responses to the selected environmental variables. Our results suggest that more disturbed sites are significantly poorer in oak forest specialists, which are usually more abundant in large oak fragments and decrease in abundance or disappear in small fragments and surrounding matrix habitats. Thus, it is necessary to preserve a minimum size of forest patch to create conditions characteristic for forest interior, rather than the more difficult task of increasing habitat connectivity.     

The maintenance of extensively exploited pastures within the Alpine mountain belt: implications for dung beetle conservation (Coleoptera: Scarabaeoidea)

The abandonment of marginal areas and the decline of traditional small-scale cattle-breeding is bringing about substantial changes in many areas of the Alpine mountain belt. Extensively exploited pastures, which were colonized by typical ecological communities, are now being replaced by shrublands and forests. In this paper, we evaluate the effectiveness of the maintenance of small pastures for dung beetle (Scarabaeoidea: Geotrupidae, Aphodiidae, Scarabaeidae) conservation in a protected area of the north-western Italian Alps. From July until September 2007, we studied the dung beetle coenosis in the main habitats of the mountain belt: beech forest, scots pine forest, mountain pine forest and pasture. Twenty-three species were sampled. We compared abundance, species richness, α-diversity, evenness and species turnover between the four habitat assemblages and performed a correspondence analysis based on dominance patterns of the three families. Two main ensembles were identified: a pasture ensemble, highly diverse and dominated by Scarabaeidae, and a closed habitats ensemble, less structured and dominated by Aphodiidae and Geotrupidae. Combining habitat specificity and fidelity, the IndVal method showed that half of the species collected made a strong habitat selection toward pasture, while only C. granarius (Aphodiidae) was a reliable indicator of woody areas. We therefore conclude that the loss of extensively exploited pastures may bring about fundamental changes in dung insect communities. In protected areas, a management policy intended to preserve the traditional, sustainable human activities within the mountain belt is highly recommended for the conservation of these insects.     

Deforestation and biodiversity: effects of bushland cultivation on dung beetles in semi-arid Tanzania

Dung beetles provide important ecosystem services in the habitats where they occur. The activity of dung beetles enhances soil nutrient cycling and increases the soil’s ability to absorb and hold water. Consequently, these beetles are particularly important in semi-arid environments. This study analyses the importance of remaining wooded habitat patches (bushland) for the survival of a diverse dung beetle fauna in an otherwise cultivated landscape in semi-arid Tanzania. Dung beetles were sampled by pitfall trapping in maize fields and bushland habitats. In total, 6037 dung beetles (Scarabaeidae: Scarabaeinae), representing 77 species from 25 genera, were collected. Many species, particularly amongst the ball-rollers, showed a clear preference for wooded patches, species richness being significantly higher in the bushland patches than in the cultivated sites. The number of trapped specimens in bushland was also considerably higher than that found in maize fields, although the differences were not significant. In conclusion, bushland fragments appear to have an important conservation value as to maintaining a high diversity and abundance of dung beetles, thereby enhancing the ecosystem services provided by these beneficial insects.     

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.     

Relevance of exotic pine plantations as a surrogate habitat for ground beetles (Carabidae) where native forest is rare

Plantation forests are of increasing importance worldwide for wood and fibre production, and in some areas they are the only forest cover. Here we investigate the potential role of exotic plantations in supporting native forest-dwelling carabid beetles in regions that have experienced extensive deforestation. On the Canterbury Plains of New Zealand, more than 99% of the previous native forest cover has been lost, and today exotic pine (Pinus radiata) plantations are the only forest habitat of substantial area. Carabids were caught with pitfall traps in native kanuka (Kunzea ericoides) forest remnants and in a neighbouring pine plantation, grassland and gorse (Ulex europaeus) shrubland. A total of 2,700 individuals were caught, with significantly greater abundance in traps in young pine, grassland and gorse habitats than in kanuka and older pine. Rarefied species richness was greatest in kanuka, a habitat that supported two forest specialist species not present in other habitat types. A critically endangered species was found only in the exotic plantation forest, which also acts as a surrogate habitat for most carabids associated with kanuka forest. The few remaining native forest patches are of critical importance to conservation on the Canterbury Plains, but in the absence of larger native forest areas plantation forests are more valuable for carabid conservation than the exotic grassland that dominates the region.     

Community science data suggests that urbanization and forest habitat loss threaten aphidophagous native lady beetles

1. Community scientists have illustrated rapid declines of several aphidophagous lady beetle (Coccinellidae) species. These declines coincide with the establishment of alien coccinellids. We established the Buckeye Lady Beetle Blitz program to measure the seasonal occupancy of coccinellids within gardens across a wide range of landscape contexts. Following the Habitat Compression Hypothesis, we predicted that gardens within agricultural landscapes would be alien‐dominated, whereas captures of natives would be higher within landscapes encompassing a high concentration of natural habitat.
2. Within the state of Ohio, USA, community scientists collected lady beetles for a 7‐day period across 4 years in June and August using yellow sticky card traps. All identifications were verified by professional scientists and beetles were classified by three traits: status (alien or native), mean body length, and primary diet. We compared the relative abundance and diversity of coccinellids seasonally and determined if the distribution of beetles by size, status, and diet was related to landscape features.
3. Alien species dominated the aphidophagous fauna. Native aphidophagous coccinellid abundance was positively correlated with forest habitat while alien species were more common when gardens were embedded within agricultural landscapes. Urbanization was negatively associated with both aphidophagous alien and native coccinellids.
4. Synthesis and Applications: Our census of native coccinellid species within residential gardens—a widespread and understudied habitat—was enabled by volunteers. These data will serve as an important baseline to track future changes within coccinellid communities within this region. We found that native coccinellid species richness and native aphidophagous coccinellid abundance in gardens were positively associated with forest habitat at a landscape scale of 2 km. However, our understanding of when and why (overwintering, summer foraging, or both) forest habitats are important remains unclear. Our findings highlight the need to understand how declining aphidophagous native species utilize forest habitats as a conservation priority.

     

Carabids of differently aged reforested pinewoods and a natural pine forest in a historically modified landscape

We studied the responses of carabid beetles to Pinus sylvestris ageing in four plantations aged 2–3, 10–12, 40–45 and 80 years, and one mature natural pine forest – in NW Spain. In 2003, a total of 16,866 carabid individuals (50 species) were collected using pitfall traps. Overall carabid abundance and species richness differed significantly among the five age classes with the highest values occurring in the youngest age class. The carabid assemblages of the initial stages of the ageing sequence differed considerably from the later stages that were relatively similar to each other – a consequence of differences in species dominance at specific age classes. Yet, the carabid assemblage of the 80-year-old stage did not approach that of the natural pine forest, but, unexpectedly, the 40–45-year-old age class did. Canopy cover and soil pH were the main environmental variables that affected the carabid distribution. We also detected species-level responses to the ageing process, with open habitat species more abundantly collected at the youngest stages and forest specialists at the older stages. We conclude that carabid beetles responded predictably to the pine plantation ageing process, and that although pine forests in NW Spain do not seem to provide habitat for unique species, they act as secondary habitats for forest specialist species.     

The Rove Beetles (Coleoptera, Staphylinidae) of Exposed Riverine Sediments in Scotland and Northern England: Habitat Classification and Conservation Aspects

Exposed riverine sediments (ERS) by four rivers in Scotland and northern England were sampled for beetles in 1996 and 1997. One hundred and sixty rove beetle (Staphylinidae) species lists were analysed using ordination and classification techniques in order to identify habitat groups within and between catchments and to assess which factors were affecting species assemblage distribution. There were major differences between the species assemblages of ERS by rivers of highland and lowland catchments. Within catchments, assemblage distribution was mainly influenced by the position of sites within the catchment; vegetation cover and sediment composition had less influence. The number of rove beetle habitats was not the same as those for ground and phytophagous beetle groups, indicating that conservation considerations should take into account variations in ERS habitat diversity. A considerable number of records of nationally rare and scarce rove beetle species were recorded, most on ERS by rivers and tributaries unaffected by river management or engineering.     

Beetles succession on different microhabitats of small mammals in Riyadh,Kingdom of Saudi Arabia

The decay of small carcasses is considerably neglected although it may provide important forensic data. This study differentiates the beetle species correlated with rat, Rattus norvegicus and mouse, Mus musculus (L. 1758) carcasses placed in two different habitats at King Saud University in western Riyadh. Decay processes and beetle succession were analyzed from January to February 2018. We collected an average of 268.67 beetles belonging to nine species from eight families, including Histeridae, Dermestidae, Cleridae, Tenebrionidae, Staphylinidae, Coccinellidae, Melolonthinae, and Pyrrhocoridae. Collected specimens were identified morphologically and by molecular techniques based on the mitochondrial cytochrome oxidase subunit I (COI) gene. Their presence rose in the bloated and decaying stages of decomposition. The beetle communities differed between the two habitats. In addition, a distinctive beetle community was found in the decaying stage compared to the other stages. The lowest number of beetles was reported in the dry stage. The abundance of beetles was associated with carcass size (rat or mouse) and habitat type.     

Conserving butterflies in fragmented plantation forests: are edge and interior habitats equally important?

Edge effects are increasing in forest-dominated landscapes worldwide, due to increased fragmentation by other land uses. Understanding how species respond to edges is therefore critical to define adequate conservation measures. We compared the relative importance of interior and edge habitats for butterflies in a landscape composed of even-aged pine plantations interspersed with semi-natural habitats. Butterfly assemblages were surveyed simultaneously at the edge and the interior of 68 patches belonging to four main habitat types: herbaceous firebreaks, clearcuts and young pine stands, older pine stands, and deciduous woodlands. Butterfly species richness was higher at edges than in interior habitats, especially for pine stands. Assemblage composition differed significantly between edge and interior habitats, except for firebreaks. Of the 23 most abundant butterfly species, seven were significantly more abundant in one or all edge habitat types, five in interior habitats, and 11 species showed no edge-interior preference. Modelling the presence of individual species in edge habitats revealed the importance of habitat variables such as the abundance of nectar and host-plants, but also of the abundance of the same species in the adjacent interior habitat. Moreover, our results suggest that most species use several, different habitat types to find supplementary or complementary resources, including micro-climatic refuges to escape hot temperatures during summer. The use of adjacent edge and interior habitats by butterflies is probably a key process in such mosaic landscapes and underlines the importance of landscape heterogeneity for butterfly conservation.     

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.     

Diversity of surface dwelling beetle assemblages in open-cast lignite mines in Central Germany

We investigated species richness of ground dwelling beetle assemblages in two non-reclaimed lignite mines and a dump in Central Germany by means of pitfall trapping. During a period of five months, a total of 203 beetle species within 27 families represented by 4099 individuals were trapped. This included 75 species of ground beetles represented in a sample of 957 individuals from which 10 species are regionally endangered. The number of individuals, species richness, as well as the proportions of endangered species did not differ between successional stages whereas species composition of sites could be related well to a set of environmental variables. High values of beta-diversity between sites indicated that the total number of species recorded is caused by habitat diversity. From the viewpoint of nature conservation, we conclude that postmining areas can play a key role in conservation of beetle diversity in agricultural areas since they harbour threatened species whose original habitats are now rare due to human impact. An important task for future management of postmining areas is to maintain successional processes and to prevent loss of habitat diversity through afforestation. Areas with extreme soil conditions should also be preserved for long-term availability of bare soil and pioneer vegetation and associated fauna.     

Habitat loss decreases predator–prey ratios in a pine-bark beetle system

Increasing intensity of land use by humans has led to loss of natural habitats, resulting in isolation of remaining habitat fragments. Using a pine-bark beetle ecosystem as a model, we tested the hypothesis that the ratio of abundance of predators to prey should decrease with increasing habitat loss at the landscape scale. We selected ten red pine ( Pinus resinosa ) sites, representing extremes of available habitat within a 2 km radius surrounding each stand. The bark beetle, Ips pini , and its coleopteran predators were sampled using baited multiple funnel traps. Effects of stand isolation were considerable; ratios of predators to prey (mean number of predators/number of prey±SE) were significantly reduced in isolated stands (0.38±0.09) as compared to those with large amounts of surrounding conifer habitat (1.63±0.41). The decline in ratio occurred both because there was: a) a lower abundance of predators (ca 0.5–0.8×) captured in isolated stands; and b) a significantly higher number of prey (ca 2.2×) captured in isolated stands. Isolation or loss of habitat, therefore, differentially affected the two trophic levels, supporting theoretical predictions. Reductions in predator abundance and, presumably, enemy-caused mortality may lead to changes in the population dynamics of their prey species, possibly leading to increased outbreaks as habitat becomes increasingly isolated.     

Habitat selection and grouping of beetles (Coleoptera)

Beetles were collected by pitfall trapping for a two-year period in seven adjacent habitats in an upland site in North Wales. Positive correlations were demonstrated between number of beetle species and number and diversity of plant species. Similar correlations were shown between beetle numbers and plant species. However, only 15% of the beetle species were herbivores requiring host plants. The degree of habitat selection by individual beetle species was demonstrated, ranging from habitat specialist, being found in one habitat, lo habitat generalist, being found in most habitats. Herbivores were significantly more habitat specialist than predators or scavengers. The grouping of beetles, demonstrated by ordination analysis, was similar to, but less precise than, the grouping of plant species. The beetle groups reflect habitat selection preferences by individual species rather than a functional relationship between beetle species. They provide an example of the centrifugal structure of habitat selection theory.