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.     

Macroalgal Composition Determines the Structure of Benthic Assemblages Colonizing Fragmented Habitats

Understanding the consequences of fragmentation of coastal habitats is an important topic of discussion in marine ecology. Research on the effects of fragmentation has revealed complex and context-dependent biotic responses, which prevent generalizations across different habitats or study organisms. The effects of fragmentation in marine environments have been rarely investigated across heterogeneous habitats, since most studies have focused on a single type of habitat or patch. In this study, we assessed the effects of different levels of fragmentation (i.e. decreasing size of patches without overall habitat loss). We measured these effects using assemblages of macro-invertebrates colonizing representative morphological groups of intertidal macroalgae (e.g. encrusting, turf and canopy-forming algae). For this purpose, we constructed artificial assemblages with different combinations of morphological groups and increasing levels of fragmentation by manipulating the amount of bare rock or the spatial arrangement of different species in mixed assemblages. In general, our results showed that 1) fragmentation did not significantly affect the assemblages of macroinvertebrates; 2) at greater levels of fragmentation, there were greater numbers of species in mixed algal assemblages, suggesting that higher habitat complexity promotes species colonization. Our results suggest that predicting the consequences of fragmentation in heterogeneous habitats is dependent on the type and diversity of morphological groups making up those habitats.     

Leaving uncut refuges during meadow harvesting increases the functional diversity of Orthoptera

In the past, insect species richness was high in Central European seminatural grasslands, which were characterized by low‐intensity land use. Currently, however, the hay in most of these grasslands is mechanically harvested, which negatively impacts insect biodiversity. One way to reduce this negative effect is to leave unmown patches as refuges. In the current research we evaluated the short‐term effects of leaving an unmown patch on the taxonomic and functional diversity of the Orthoptera assemblage in a meadow. We found that orthopteran species richness and abundance were significantly reduced by mowing, whether or not a patch was left uncut. In contrast, functional evenness, indicating distribution of species abundances in a niche space, was reduced by mowing only if the plot lacked an uncut refuge. Functional richness, indicating the amount of niche space occupied by species, was elevated if the plot had an uncut refuge. Larger species were negatively affected by mowing, while habitat specialists, mobile species and soil‐ovipositing species benefitted from it. We infer that the presence of an uncut patch increased the diversity of habitats available to orthopterans and maintained even distribution of species among niche space. In summary, leaving an unmown refuge in grasslands could increase the functional diversity of orthopterans, even if it does not preserve taxonomic diversity.     

Interactions of components of habitats alter composition and variability of assemblages

1. The nature and resources supplied by different components of habitats influence species, creating variability from place to place within a habitat. 2. Experiments were done to investigate the effects of altering components of habitats on the variability of assemblages of numerous species of intertidal gastropods. 3. Artificial habitats with three levels of structure, combining different types of turf (i.e. different densities and height of fronds) were sampled 8 weeks after deployment in the intertidal. They were rapidly colonized by up to 66 species of gastropods. 4. Independently of the types of turf combined to form different habitats, there were differences in assemblages where there was more than one type of component present. Multivariate dissimilarities among units making up each habitat were also greater where there were more than one type of unit, but there was no such difference in the variance of numbers of species per unit. 5. Altering the relative abundances of different types of components made little change to the assemblages, nor their multivariate variability among units of habitat and the variance in number of species per unit in each habitat. 6. Differences in assemblages due to the different structure of habitat are complex to interpret and simple characterizations of structure of habitat are inadequate. Comparing different habitats requires appropriate experimentation to ensure that variability within habitats is also investigated.     

The diversity of insectivorous bat assemblages among habitats within a subtropical urban landscape

We investigated the bat (Microchiroptera) diversity of four major habitat types within a large Australian subtropical city (Brisbane, Australia) to determine whether species richness was affected by habitat changes associated with urbanization, as suggested from studies elsewhere. Forty sites, ten in each habitat type (remnant bushland, parkland, low‐density residential and high‐density residential) were surveyed using acoustic bat detectors on six non‐consecutive occasions. Fourteen bat species were recorded. The species accumulation curve of the entire Brisbane bat assemblage reached a plateau at 14 species. The total numbers of species in bushland, parkland, low‐density residential and high‐density residential habitats were 14, 13, 14 and 11 species, respectively. Asymptotic estimates of species richness for each habitat were close or equal to these totals. Mean asymptotic estimated species richness differed significantly among habitats, being lowest in high‐density residential sites and highest in low‐density residential sites. Evenness profiles were similar across habitats, and were not strongly dominated by a few species. Partitioning of diversity components showed that landscape (γ) diversity was mainly determined by the high species richness of low‐density residential and bushland habitats (α diversity), rather than high beta (β) diversity among habitats. These findings contradict those of other studies on bat diversity in which species richness was highest within ‘natural’ areas of the urban landscape and assemblages were dominated by one or two species. This highlights the need for caution in making generalizations based on existing information, which is dominated by studies in temperate regions.     

Declining diversity and abundance of High Arctic fly assemblages over two decades of rapid climate warming

Insects are particularly vulnerable to rapid environmental changes, which are disproportionally affecting high latitudes. Increased temperature could influence insect species differentially and reshape assemblages over time. We quantified temporal assemblage turnover of Arctic Diptera (flies) in the Muscidae, one of the most diverse and abundant families of Arctic insects, using time series data from Zackenberg, north‐east Greenland. We measured temporal patterns of abundance, diversity, and composition of muscid assemblages in wet fen, mesic and arid heath habitats from yearly collections spanning 1996–2014 and tested their relationship to climate. A total of 18 385 individuals representing 16 species of muscid flies were identified. A significant decrease of 80% of total muscid abundance was observed during the study period. Species richness declined in each habitat type but this trend was not significant across habitats. The number of common and abundant species also decreased significantly over time across habitats revealing a temporal modification of species evenness. Significant temporal changes in composition observed in the wet fen and across habitats were mainly driven by a change in relative abundance of certain species rather than by species replacement. Shift in composition in each habitat and decline in muscid abundance across habitats were associated with summer temperature, which has significantly increased over the study period. However, relationships between temperature and muscid abundance at the species level were noticeable for a few species only. Significant directional change in composition was documented in the wet fen but no biotic homogenization across habitats was observed. As one of the few studies of species‐level changes in abundance, diversity and composition of an insect taxon in the Arctic over the past two decades, our study shows that habitat types may modulate insect species responses to recent climate change and that contrasting species responses can alter species assemblages within a few decades.     

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.     

Conserving a variety of ancient forest patches maintains historic arthropod diversity

Forests are naturally extensive tracts. However, in South Africa natural fires over many millennia have reduced forested areas into small remnants spread throughout a grassland matrix. Small patches, especially distant patches, are generally considered to be adverse for forest specialists, owing to decreased forest interior and increased edge. Here we test this assumption by determining the impact of forest interpatch distance and patch size on epigaeic arthropod diversity in this globally rare vegetation type. Forty patches were selected: ten large (100–435 m diameter) that are distant (500–645 m) from other patches, ten large that are close to other patches (38–97 m), ten small (30–42 m) that are distant, and ten small-close patches. Each patch had two plots: edge and interior. Arthropods were sampled using pitfall traps, Berlese-Tullgren funnels and active searches. Interiors and edges had similar species richness and composition, excluding spiders, which were richer in interiors. Patch size significantly influenced species richness of predatory beetles and arthropod assemblages, excluding spiders. Effect of the interaction between patch size and interpatch distance on species richness and composition varied among taxa. Furthermore, large patches supported similar assemblages regardless of interpatch distance. Arthropod response, particularly ants to patch size and interpatch distance, was partly shaped by the matrix type. The percentage of surrounding grassland had little effect on arthropod diversity. We can conclude that large and close patches are important for arthropod conservation. Nevertheless, it is also important to conserve a variety of patch sizes at various distances to maximize overall arthropod composition.     

Pervasive effects of dispersal limitation on within- and among-community species richness in agricultural landscapes

Aim To determine whether the effect of habitat fragmentation and habitat heterogeneity on species richness at different spatial scales depends on the dispersal ability of the species assemblages and if this results in nested species assemblages. Location Agricultural landscapes distributed over seven temperate Europe countries covering a range from France to Estonia. Methods We sampled 16 local communities in each of 24 agricultural landscapes (16 km²) that differ in the amount and heterogeneity of semi‐natural habitat patches. Carabid beetles were used as model organisms as dispersal ability can easily be assessed on morphological traits. The proximity and heterogeneity of semi‐natural patches within the landscape were related to average local (alpha), between local (beta) and landscape (gamma) species richness and compared among four guilds that differ in dispersal ability. Results For species assemblages with low dispersal ability, local diversity increased as the proximity of semi‐natural habitat increased, while mobile species showed an opposite trend. Beta diversity decreased equally for all dispersal classes in relation to proximity, suggesting a homogenizing effect of increased patch isolation. In contrast, habitat diversity of the semi‐natural patches affected beta diversity positively only for less mobile species, probably due to the low dispersal ability of specialist species. Species with low mobility that persisted in highly fragmented landscapes were consistently present in less fragmented ones, resulting in nested assemblages for this mobility class only. Main conclusions The incorporation of dispersal ability reveals that only local species assemblages with low dispersal ability show a decrease of richness as a result of fragmentation. This local species loss is compensated at least in part by an increase in species with high dispersal ability, which obscures the effect of fragmentation when investigated across dispersal groups. Conversely, fragmentation homogenizes the landscape fauna for all dispersal groups, which indicates the invasion of non‐crop habitats by similar good dispersers across the whole landscape. Given that recolonization of low dispersers is unlikely, depletion of these species in modern agricultural landscapes appears temporally pervasive.     

Partitioning beta‐diversity through different pond hydroperiod lengths reveals predominance of nestedness in assemblages of immature odonates

Patterns of freshwater invertebrate assemblage structure in the transition from permanent to non‐permanent lentic habitats are well described in the literature. However, the effects of small changes in the hydroperiod of non‐permanent ponds on invertebrate assemblage structure remain less studied, especially on β‐diversity. Thus, we tested the effects of different pond hydroperiod lengths on the assemblage structure of immature odonates, in terms of both α‐ and β‐diversity. Small high‐altitude ponds with different hydroperiod lengths (assigned to ‘short’, ‘medium’ and ‘long’ hydroperiods) were sampled in southern Brazil between 2013 and 2014. Based on the hypothesis that shorter hydroperiods filter constituents of lentic fauna, i.e. that long‐living species cannot inhabit shorter‐hydroperiod ponds, we expected to find higher α‐ and β‐diversity in longer hydroperiods, as well as predominance of the nestedness component in β‐diversity. Restricted occurrence of some genera and higher α‐diversity of immature odonate assemblages was detected in long‐hydroperiod ponds. Within‐hydroperiod β‐diversity values did not vary among hydroperiods, because the occasional occurrence of some genera with high dispersal ability of adults in short‐hydroperiod ponds yielded similar values of the β‐diversity among hydroperiods. Partitioning of β‐diversity among hydroperiods revealed a significant higher contribution of the nestedness component rather than turnover. This pattern is explained by the occurrence of some generalist genera across the whole gradient of hydroperiod, as a subset of fauna in longer‐hydroperiod ponds. Thus, our results suggest that reduction in hydroperiod length, if occurring in the future climate change, would favor habitat‐generalist taxa in lentic ecosystems.     

Landscape mosaic of Araucaria forest and forest monocultures influencing understorey spider assemblages in southern Brazil

Abstract This study investigates how abundance, diversity and composition of understorey spiders were influenced by four different forest habitats in a southern Brazilian Araucaria forest. The study area encompasses a landscape mosaic comprised of Araucaria forest, Araucaria plantation, Pinus plantation, and Eucalyptus plantation. Understorey spiders were collected by beating the vegetation inside three patches of each forest habitat. To assess possible predictors of spider assemblage structure, several patch features were analysed: potential prey abundances, estimation of vegetation cover, diversity index of vegetation types, patch ages, patch areas, and geographical distance between patches. To assess the influence of high‐level taxa approaches on spider assemblage patterns, analyses were carried out individually for family, genera and species levels. Additionally, Mantel tests were carried out in underlying similarity matrices between each taxon. Significant differences in spider abundances among forest habitats were found. Pinus plantations showed the highest abundance of spiders and Eucalyptus plantations showed the lowest abundance. Spider abundance was significantly influenced by patch ages, geographical distance and vegetation cover. Expected numbers of families, genera and species did not vary among forest habitats. Spider composition of two Eucalyptus patches differed from the other forest patches, probably due to their low vegetation cover and isolation. Genera composition was the best correlate of species composition, showing that a higher‐level surrogate can be an alternative to the species approach. The understorey spider diversity in this managed area could be maintained when suitable habitat structures are provided, thus ensuring the connectivity between different habitat types. Further studies should focus on individual species responses to the conversion of native forest to monocultures.     

Role of colonization in spatio-temporal patchiness of microgastropods in coralline turf habitat

The ability of benthic macrofauna to disperse and colonize new habitats throughout their life may contribute substantially to small-scale patchiness in abundances in different habitats. Microgastropods in coralline turf on rocky shores in Australia are very patchy in abundance at different spatio-temporal scales. They therefore represent an ideal assemblage for testing hypotheses about processes of colonization. Patterns and rates of colonization of 10 species of microgastropods were investigated in one intertidal habitat (coralline turf) in Botany Bay, New South Wales, Australia, using artificial substrata, which are considered to be a good mimic of natural coralline turfs. The experiment was designed to test the hypotheses that (1) patterns (abundance of colonizing individuals) and mode (juveniles vs. adults) of colonization depends on the proximity of a patch to a potential source of dispersing colonists (i.e. patch of natural coralline turf), (2) different species show different rates of colonization, and (3) patterns of succession are not repeatable among different patches of natural algae. Seven different plots (natural patches of coralline turf) were randomly chosen in the area of study. Artificial units (called patches) were placed at different distances around each plot (within the plot and 0, 50 or 100 cm away from the edge). Samples were collected 6, 13, and 27 days after the experiment started. Colonization was rapid (i.e. within 6 days) for most species. Pattern (number of individuals) and mode of colonization (adults vs. juveniles) varied among algal plots. Most species responded differently across patches causing no consistent patterns of colonization. Furthermore, patterns of colonization of artificial units were not always synchronous with, or in the same direction as, changes in abundances in the nearest algal plots, which themselves showed no consistent spatio-temporal pattern.     

Guinean biodiversity at the edge: Rodents in forest patches of southern Mali

Southern Mali mainly belongs to the Sudanian savanna bioclimatic zone, but forest patches showing botanical affinities with Guinean humid forest remain as gallery forests or ravine forests. To characterize the rodent diversity of this area and check for the presence of some species of Guinean affinities in this group, rodent assemblages were sampled in four regions of southern Mali, using trapping and observational data in forest and surrounding habitats. Twenty-four species were recorded, comprising a representative sample of the expected overall diversity in this group according to rarefaction curves. Praomys rostratus was the dominant species in the most humid, closed lowland forest. Praomys daltoni was also present in this habitat type, being all the more abundant as habitat degradation was apparent. It became the dominant species in ravine forest on rocky substrate where P. rostratus was completely absent. In Sudanian savanna habitats and in herbaceous and cultivated areas, Mastomys erythroleucus dominated a diverse rodent community. A few species were found that testified for Guinean affinities of the most humid forest patches, especially in the extreme southeast of the country (region of Sikasso). Rodent assemblages of the Bafing and Mts Mandingues areas, in the western part of the study area, showed the highest similarity, in relation with environmental characteristics of this region representing an extension of the Fouta Djallon plateau in Guinea. The results obtained highlight the high biodiversity value of this forest-savanna mosaic, and provide new arguments in favour of the preservation of West African forest patches and their surrounding habitats.     

Habitat associations of epigeal spiders in upland calcareous grassland landscapes: the importance for conservation

Upland calcareous grassland landscapes are typically comprised of a matrix of calcareous grassland, acid grassland and limestone heath plant communities. This matrix of habitats is produced by a combination of underlying geology, climate and management. These landscapes are typically managed through grazing, with management targeted to maintain particular plant communities in the calcareous grassland habitat, whilst patches of acid grassland and limestone heath are not targeted by conservation management. The biodiversity value of acid grassland and limestone heath patches within the calcareous grassland matrix are unknown. This study provides the first assessment of their biodiversity value by examining aspects of epigeal spider diversity supported by these non-target habitat patches in comparison to calcareous grassland. Spiders were sampled in each habitat from April to August 2014 using pitfall traps across three upland regions in Great Britain. Spider species assemblages were distinct between limestone heath and both grassland types. Distinction in species assemblages are likely due to differences in vegetation structure and microclimate, e.g., humidity, degree of shade. Each habitat type supported several rare species (e.g., Jacksonella falconeri, Agyneta subtilis) revealing the contribution to spider fauna. The distinct spider species assemblage and presence of rare species in limestone heath patches demonstrate their importance in the upland calcareous grassland matrix. This study highlights the value of monitoring biodiversity in non-target habitats within a habitat matrix alongside those that are actively targeted by management.     

Effects of anthropogenic habitat disturbance on local pollinator diversity and species turnover across a precipitation gradient

Anthropogenic habitat disturbance can have profound effects on multiple components of forest biotas including pollinator assemblages. We assessed the effect of small-scale disturbance on local richness, abundance, diversity and evenness of insect pollinator fauna; and how habitat disturbance affected species turnover across the landscape and overall diversity along a precipitation gradient in NW Patagonia (Argentina). We evaluated the effect of disturbance on overall pollinator fauna and then separately for bees (i.e. Apoidea) and non-bee pollinators. Locally, disturbed habitats had significantly higher pollinator species richness and abundances than undisturbed habitats for the whole pollinator assemblage, but not for bees or non-bees separately. However, significant differences in species richness between habitats vanished after accounting for differences in abundance between habitat types. At a local scale Shannon–Weaver diversity and evenness did not vary with disturbance. A β diversity index indicated that, across forest types, species turnover was lower between disturbed habitats than between undisturbed habitats. In addition, rarefaction curves showed that disturbed habitats as a whole accumulated fewer species than undisturbed habitats at equivalent sample sizes. We concluded that small patches of disturbed habitat have a negligible effect on local pollinator diversity; however, habitat disturbance reduced β diversity through a homogenization of the pollinator fauna (in particular of bees) across the landscape.     

Species–area relationship within benthic habitat patches of a tropical floodplain river: An experimental test

The curvilinear relationship between species richness and habitat area (species–area relationship (SAR)) is a fundamental ecological pattern. The relationship is often viewed from a long‐term perspective across relatively large spatial scales, reflecting a balance between immigration and extinction dynamics. We explored whether predictions of SAR also manifest over short time periods (days) in benthic habitat patches of a dynamic floodplain river where littoral faunal assemblages are continuously assembled and disassembled with changing water levels. We examined the relationship of patch size with faunal abundance (i.e. fish and aquatic invertebrates), taxonomic richness, trophic group richness and overall assemblage composition. Strong taxa–area relationships emerged despite the relatively short experimental time period (21 days); larger patches had more taxa and trophic groups. For the smallest patches, taxonomic richness was especially sensitive to abundance of individuals; abundance of individuals was a less important predictor of taxonomic and trophic group richness for the largest patches. Despite the relatively short time frame for study within this temporally dynamic ecosystem, our findings indicate a strong SAR for fishes and macroinvertebrates inhabiting patchy habitats in the littoral zone of this tropical river.     

Intertidal assemblages associated with natural<Emphasis Type="Italic">corallina</Emphasis> turf and invasive mussel beds

There is considerable concern about conservation of biodiversity in highly disturbed and urbanized environments, although a very large proportion of biodiversity (i.e. the small and cryptic invertebrates) have been little studied in this regard. Many biogenic structures (e.g. coral reefs, mussel beds, foliose algae) provide habitat for a large number of small invertebrates. The features of these habitats to which these animals respond are complex and poorly documented. Invasive species are increasing in abundance and diversity in many disturbed estuaries, but most previous studies have concentrated on effects of invasive species on surrounding macroscopic assemblages. This study examines the assemblages of small invertebrates and algae living in natural patches of coralline turf and in patches of the invasive mussel, Mytilus galloprovincialis, on seawalls in Sydney Harbour. Although most taxa identified were common to both habitats, they were generally more abundant in turf than in the mussels, especially the more widespread and numerous taxa. Few taxa were unique to either habitat and those were generally sparse and patchy. In addition, there were relatively more smaller animals in the algal turf than in the mussels, although it is not known whether these were juveniles of adults present in both habitats, or different species. These data show that coralline turf and mussel beds do not provide similar intertidal habitat for associated assemblages and that overgrowth of natural biota by mussels may have strong indirect effects on associated assemblages. These warrant further experimental investigation, so that the effects of invasive species on local biodiversity can be better understood and managed.     

Environmental and spatial drivers of spider diversity at contrasting microhabitats

The relative importance of environmental and spatial drivers of animal diversity varies across scales, but identifying these scales can be difficult if a sampling design does not match the scale of the target organisms' interaction with their habitat. In this study, we quantify and compare the effects of environmental variation and spatial proximity on ground‐dwelling spider assemblages sampled from three distinct microhabitat types (open grassland, logs, trees) that recur across structurally heterogeneous grassy woodlands. We used model selection and multivariate procedures to compare the effects of different environmental attributes and spatial proximity on spider assemblages at each microhabitat type. We found that species richness and assemblage composition differed among microhabitat types. Bare ground cover had a negative effect on spider richness under trees, but a positive effect on spider richness in open grassland. Turnover in spider assemblages from open grassland was correlated with environmental distance, but not geographic distance. By contrast, turnover in spiders at logs and trees was correlated with geographic distance, but not environmental distance. Our study suggests that spider assemblages from widespread and connected open grassland habitat were more affected by environmental than spatial gradients, whereas spiders at log and tree habitats were more affected by spatial distance among these discrete but recurring microhabitats. Deliberate selection and sampling of small‐scale habitat features can provide robust information about the drivers of arthropod diversity and turnover in landscapes.     

Partitioning the diversity of riverine fish: the roles of habitat types and non-native species

1. Quantifying how biological diversity is distributed in the landscape is one of the central themes of conservation ecology. For this purpose, landscape classifications are being intensively used in conservation planning and biodiversity management, although there is still little information about their efficacy. 2. I used data from 158 running water sites in Hungary to examine the contribution of six a priori established habitat types to regional level diversity of fish assemblages. Three community measures [species richness, diversity (Shannon, Simpson indices), assemblage composition] were examined at two assemblage levels (entire assemblage, the native assemblage). The relative role of non‐native species was quantified to examine their contribution to patterns in diversity in this strongly human influenced landscape. 3. Additive diversity partitioning revealed the primary importance of beta diversity (i.e. among‐site factors) to patterns in species richness. Landscape‐scale patterns in species richness were best explained by between‐habitat type (beta₂: 41.2%), followed by within‐habitat type (beta₁: 37.7%) and finally within‐site (alpha: 21.1%) diversity. Diversity indices showed patterns different from species richness, indicating the importance of relative abundance distributions on the results. Exclusion of non‐natives from the analysis gave similar results to the entire‐assemblage level analysis. 4. Canonical analysis of principal coordinates, complemented with indicator species analysis justified the separation of fish assemblages among the habitat types, although classification error was high. Multivariate dispersion, a measure of compositional beta diversity, showed significant differences among the habitat types. Contrary to species diversity (i.e. richness, diversity indices), patterns in compositional diversity were strongly influenced by the exclusion of non‐natives from the analyses. 5. This study is the first to quantify how running water habitat types contribute to fish diversity at the landscape scale and how non‐native species influence this pattern. These results on riverine fish assemblages support the hypothesis that environmental variability (i.e. the diversity of habitat types) is an indication of biodiversity and can be used in large‐scale conservation designs. The study emphasises the joint application of additive diversity partitioning and multivariate statistics when exploring the contribution of landscape components to the overall biodiversity of the landscape mosaic.     

Spatial gradients in species diversity of microscopic animals: the case of bdelloid rotifers at high altitude

Aim Organisms smaller than 2 mm appear not to follow the spatial patterns in richness and diversity commonly observed in macroscopic organisms. We describe spatial patterns in species diversity in a group of microscopic organisms, bdelloid rotifers, living in moss and lichen patches, in order to test the hypotheses of no relationship between species richness and composition and spatial gradients, suggested by previously published patterns in microscopic organisms. Location Moss and lichen patches as habitats for bdelloids, on high‐elevation peaks at altitudes between 2984 and 4527 m a.s.l. across the Italian, French and Swiss Alps, with distances among sample sites ranging from 1 m to 420 km, in comparison with lower‐elevation samples at altitudes from 850 to 1810 m a.s.l. Methods We sampled species assemblages of bdelloid rotifers living in isolated moss and lichen patches in 47 sites. We described the observed α, β and γ diversities; the heterogeneity of species assemblages; and the estimated number of species (incidence‐based coverage estimator). Patterns in species distribution were analysed at three different levels: (1) habitat, comparing species richness on moss and lichen substrates, testing differences in α diversity and heterogeneity (anova ), species composition (analysis of similarities test), and γ diversity (rarefaction curves); (2) altitude, comparing the observed richness with previously published data from locations well below 2000 m; and (3) distances between sites, correlating the matrix of Jaccard dissimilarities and the matrix of geographical distances with a Mantel test. Results Both species richness and species composition of bdelloid rotifers differed significantly between mosses and lichens at high elevations, but no difference was found in the heterogeneity of species assemblages. Alpha diversity was significantly lower at high‐elevation than at low‐elevation sites, but the estimated number of species was not reduced when compared with sites at low elevations. Geographical distance between sites had no effect on species composition in either mosses or lichens. The distribution of species was highly heterogeneous, with a low similarity among assemblages. Main conclusions As expected, bdelloids appear to occupy habitats selectively. The altitudinal gradient in species richness for bdelloid rotifers is limited to a decrease in α diversity only; such a decrease is not caused by a lower number of species (low γ diversity) being able to tolerate harsh conditions, and high‐altitude species are not a subset of species living at lower elevations. The observed values of α, β and γ diversity at high altitudes in the Alps are compatible with the scenario of a very low number of available propagules because of the low density of patches of favourable habitat. Our results suggest that the geographical distribution of animals, and therefore biodiversity patterns, may be strongly influenced by animal size, as small organisms such as bdelloids appear to show spatial patterns that differ from those known in larger animals. Differences in body size should be taken into account carefully in future studies of biodiversity patterns.