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.     

An Examination of Scale-Dependent Resource Use by Eastern Hognose Snakes in Southcentral New Hampshire

ABSTRACT The decline of many snake populations is attributable to habitat loss, and knowledge of habitat use is critical to their conservation. Resource characteristics (e.g., relative availability of different habitat types, soils, and slopes) within a landscape are scale-dependent and may not be equal across multiple spatial scales. Thus, it is important to identify the relevant spatial scales at which resource selection occurs. We conducted a radiotelemetry study of eastern hognose snake (Heterodon platirhinos) home range size and resource use at different hierarchical spatial scales. We present the results for 8 snakes radiotracked during a 2-year study at New Boston Air Force Station (NBAFS) in southern New Hampshire, USA, where the species is listed by the state as endangered. Mean home range size (minimum convex polygon) at NBAFS (51.7 ± 14.7 ha) was similar to that reported in other parts of the species’ range. Radiotracked snakes exhibited different patterns of resource use at different spatial scales. At the landscape scale (selection of locations within the landscape), snakes overutilized old-field and forest edge habitats and underutilized forested habitats and wetlands relative to availability. At this scale, snakes also overutilized areas containing sandy loam soils and areas with lower slope (mean slope = 5.2% at snake locations vs. 6.7% at random locations). We failed to detect some of these patterns of resource use at the home range scale (i.e., within the home range). Our ability to detect resource selection by the snakes only at the landscape scale is likely the result of greater heterogeneity in macrohabitat features at the broader landscape scale. From a management perspective, future studies of habitat selection for rare species should include measurement of available habitat at spatial scales larger than the home range. We suggest that the maintenance of open early successional habitats as a component of forested landscapes will be critical for the persistence of eastern hognose snake populations in the northeastern United States.     

Ecological biogeography of Mexican bats: the relative contributions of habitat heterogeneity,beta diversity,and environmental gradients to species richness and composition patterns

Mexico has higher mammalian diversity than expected for its size and geographic position. High environmental hetero geneity throughout Mexico is hypothesized to promote high turnover rates (β‐diversity), thus contributing more to observed species richness and composition than within‐habitat (α) diversity. This is true if species are strongly associated with their environments, such that changes in environmental attributes will result in changes in species composition. Also, greater heterogeneity in an area will result in greater species richness. This hypothesis has been deemed false for bats, as their ability to fly would reduce opportunities for habitat specialization. If so, we would expect no significant relationships between 1) species composition and environmental variables, 2) species richness and environmental heterogeneity, 3) β‐diversity and environmental heterogeneity. We tested these predictions using 31 bat assemblages distributed across Mexico. Using variance partitioning we evaluated the relative contribution of vegetation, climate, elevation, horizontal heterogeneity (a variate including vegetation, climate, and elevational heterogeneity), spatial variation (lat‐long), and vertical hetero geneity (of vegetation strata) to variation in bat species composition and richness. Variation in vegetation explained 92% of the variation in species composition and was correlated with all other variables examined, indicating that bats respond directly to habitat composition and structure. Beta‐diversity and vegetational heterogeneity were significantly correlated. Bat species richness was significantly correlated with vertical, but not horizontal, heterogeneity. Nonetheless, neither horizontal nor vertical heterogeneity were random; both were related to latitude and to elevation. Variation in bat community composition and richness in Mexico were primarily explained by local landscape heterogeneity and environmental factors. Significant relationships between β‐diversity and environmental variation reveal differences in habitat specialization by bats, and explain their high diversity in Mexico. Understanding mechanisms acting along environmental or geographic gradients is as important for understanding spatial variation in community composition as studying mechanisms that operate at local scales.     

Contrasting effects of mosaic structure on alpha and beta diversity of bird assemblages in a human‐modified landscape

Habitat loss and fragmentation are key processes causing biodiversity loss in human‐modified landscapes. Knowledge of these processes has largely been derived from measuring biodiversity at the scale of ‘within‐habitat’ fragments with the surrounding landscape considered as matrix. Yet, the loss of variation in species assemblages ‘among’ habitat fragments (landscape‐scale) may be as important a driver of biodiversity loss as the loss of diversity ‘within’ habitat fragments (local‐scale). We tested the hypothesis that heterogeneity in vegetation cover is important for maintaining alpha and beta diversity in human‐modified landscapes. We surveyed bird assemblages in eighty 300‐m‐long transects nested within twenty 1‐km² vegetation ‘mosaics’, with mosaics assigned to four categories defined by the cover extent and configuration of native eucalypt forest and exotic pine plantation. We examined bird assemblages at two spatial scales: 1) within and among transects, and 2) within and among mosaics. Alpha diversity was the mean species diversity within‐transects or within‐mosaics and beta diversity quantified the effective number of compositionally distinct transects or mosaics. We found that within‐transect alpha diversity was highest in vegetation mosaics defined by continuous eucalypt forest, lowest in mosaics of continuous pine plantation, and at intermediate levels in mosaics containing eucalypt patches in a pine matrix. We found that eucalypt mosaics had lower beta diversity than other mosaic types when ignoring relative abundances, but had similar or higher beta diversity when weighting with species abundances. Mosaics containing both pine and eucalypt forest differed in their bird compositional variation among transects, despite sharing a similar suite of species. This configuration effect at the mosaic scale reflected differences in vegetation composition among transects. Maintaining heterogeneity in vegetation cover could help to maintain variation among bird assemblages across landscapes, thus partially offsetting local‐scale diversity losses due to fragmentation. Critical to this is the retention of remnant native vegetation.     

Estimated plant richness pattern across northwest South America provides similar support for the species-energy and spatial heterogeneity hypotheses

Explaining geographic variation in plant species richness at broad spatial scales has long been a major challenge. Many hypotheses have been proposed during the last 200 yr, but recent work has focused on a few major alternatives. Among these, two hypotheses contend that plant species richness reflects 1) variation in energy and water availability among sampling units (the species-energy hypothesis) and 2) habitat and topographic heterogeneity within sampling units (the spatial heterogeneity hypothesis). We used a large botanical database and regression models to simultaneously confront the predictions from both hypotheses against an estimate of vascular plant richness across northwest South America. This estimate provided similar support for both hypotheses, a result that may be seen as contrasting with the notion that variation in energy and water availability among sampling units is the main determinant of plant species richness. We discuss potential explanations for this apparent discrepancy. Regression models that incorporated the relative contributions of both hypotheses predicted that the highest plant species richness in northwest South America is found in topographically complex areas. In contrast to several of the most recent mapping efforts, lowland Amazonia was predicted to be a plant richness trough in the study region. We suggest that diverging portrayals of plant richness across northwest South America result from differences in estimates of the relative importance of the species-energy and the spatial heterogeneity hypotheses.     

Generation of Spatial Patterns in Boreal Forest Landscapes

Boreal forests are composed of a few plant species with contrasting traits with respect to ecosystem functioning and spatial patterning. Early successional deciduous species, such as birch and aspen, disperse seeds widely, do not tolerate low light and nitrogen availabilities, have rapidly decaying litter, and are highly preferred by herbivores. These later succeed to conifers, such as spruce and fir, which disperse seeds locally, tolerate low light levels and low nitrogen availability, have litter that decays slowly, and are unpalatable to most mammalian herbivores. Although there are also early successional conifers, such as jack pine and Scots pine, the aspen-birch-spruce-fir successional sequence is the most common over much of North America, and (without fir) in Fennoscandia and Siberia. The course of succession in these forests is controlled partly by seed dispersal and selective foraging by mammalian herbivores. Both of these processes are spatially dynamic, but little is known about how their spatial dynamics may affect ecosystem processes, such as nitrogen cycling or productivity. We present spatially explicit models that demonstrate the following: (a) Spatially explicit seed dispersal results in more clumped distribution of tree species and persistence of greater paper birch biomass than uniform seed rain across the landscape. Such results are consistent with current spatially explicit population models of dispersal and coexistence. (b) With localized seed dispersal, the concentrations of available soil nitrogen are distributed in larger patches with sharp transitions from low to high nitrogen availability near patch edges. In contrast, with a uniform seed rain, the distribution of soil nitrogen availability was more uniform and “hotspots” were more localized. Thus, the spatial pattern of an ecosystem process (nitrogen cycling) is determined by seed dispersal and competition for light among competing populations. (c) A dispersing herbivore, such as moose, that selectively forages on early successional deciduous species with high quality litter, such as aspen or birch, and discriminates against late successional conifers, such as spruce or fir, imposes higher-order repeated patterns of plant species and biomass distribution on the landscape. Thus, seed dispersal and herbivore foraging correlate properties in adjacent patches but in different ways, and different spatial patterns emerge. Other processes, such as insect outbreaks, fire, and water flow, also may correlate properties between adjacent patches and result in additional patterns. Received 8 February 1999; accepted 28 May 1999.     

Avian taxonomic and functional diversity in early stage of longleaf pine (Pinus palustris) stands restored at agricultural lands: variations in scale dependency

In agricultural landscapes, the longleaf pine initiative (LLPI) and the Bobwhite Quail Initiative (BQI) aim to restore longleaf pine forests and early successional habitats, respectively. The early stage of longleaf pine stands and grass and forb vegetation produced by a combination of both restoration programs (LLPI‐BQI) may form habitat conditions favorable to early successional bird species and other birds, increasing avian diversity. We investigated how the LLPI and BQI programs affected taxonomic and functional diversity of birds and abundance of early successional birds (grassland and scrub/shrub species), and what environmental characteristics were associated with the diversity and abundance of birds. Our study was performed at 41 fields in Georgia, United States, during 2001–2002 by considering environmental characteristics at two spatial scales: local‐scale vegetation features and restoration program type (LLPI or LLPI‐BQI) and landscape‐scale vegetation features and landscape heterogeneity. Functional evenness, species richness, and abundance of grassland and scrub/shrub species did not show a clear association with local‐ or landscape‐scale variables. Shannon‐Wiener diversity was slightly influenced by restoration program type (local‐scale variable) with higher value at LLPI‐BQI stands than at LLPI stands despite no significant differences in local vegetation features between those stands. Functional divergence was strongly positively associated with landscape‐scale variables. That is, niche differentiation increased with increasing shrub coverage within a landscape, reducing competition between abundant bird species and others. Our results suggest that although a combination of BQI and LLPI program may have a positive effect on avian taxonomic diversity, it is important to consider shrub vegetation cover within a landscape to improve functional diversity.     

Matrix composition and landscape heterogeneity structure multiple dimensions of biodiversity in temperate forest birds

Quantifying how human-modified landscapes shape the distribution of biodiversity is critical for developing effective conservation strategies. To address this, we evaluated three hypotheses (habitat area, habitat configuration and matrix heterogeneity hypotheses) that predict responses of biodiversity to landscape structure in human-modified landscapes. We compared characteristics of landscape structure that influence taxonomic (TD), functional (FD), and phylogenetic (PD) dimensions of biodiversity of breeding birds in temperate forests. Relationships between biodiversity and landscape structure were assessed at multiple spatial scales for 20 forest interior sites in northeastern USA. We assessed if relationships with landscape structure were consistent among dimensions and assemblages of different groups (residents, migrants and all birds). Relationships between dimensions of biodiversity and landscape structure were more prevalent for FD and PD than for TD. Forest amount and configuration were rarely associated with any dimensions of biodiversity. In contrast, the identity of the matrix and heterogeneity of the landscape were frequently associated with biodiversity, but relationships differed among groups of birds. For example, FD of all birds was associated positively with landscape diversity but FD of residents was associated negatively with landscape diversity, suggesting that landscape diversity surrounding forests may increase overall FD of birds but that not all groups of species respond similarly. Indeed, biodiversity of migrants was only weakly related to landscape structure. Differences among relationships to landscape structure for bird groups and spatial scales suggests that management plans should consider local decisions within a regional framework to balance potentially conflicting needs of species groups in human-dominated landscapes.     

Pattern of local plant species richness along a gradient of landscape topographical heterogeneity: result of spatial mass effect or environmental shift?

Several processes are hypothesised to mediate the relationship between local (microsite) plant species richness and the topographical heterogeneity of the surrounding landscape. In a topographically heterogeneous landscape with various habitats occurring close to each other, local species richness may be enriched by species from surrounding habitats due to the spatial mass effect (sink‐source dynamics). In contrast, increased habitat fragmentation due to spatial heterogeneity may have a negative effect on local species richness. The spatial mass effect is thought to be more pronounced in communities with a higher ratio of generalists, as generalists are more likely to establish viable populations in sink habitats. To reveal the pattern of local species richness along a gradient of landscape topographical heterogeneity at middle altitudes of the Bohemian Massif, we used 2551 forest vegetation plots stored in the Czech National Phytosociological Database. We developed an analytical approach relating the pattern of local species richness of vegetation types to the gradient of landscape topographical heterogeneity. An increase or decrease in species richness with increasing landscape heterogeneity was related to changes in the generalist/specialist ratio, and also to changes in soil reaction and productivity estimated through Ellenberg indicator values. Local species richness along a gradient of increasing landscape heterogeneity increased in nutrient‐poor vegetation and decreased in nutrient‐rich vegetation. Nutrient‐poor vegetation types, such as thermophilous and acidophilous oak forests, also had a high proportion of habitat generalists, supporting the hypothesis that increased richness in heterogeneous landscapes may result from the spatial mass effect. However, the same pattern may be explained by a shift in environmental conditions along the landscape heterogeneity gradient, such as increasing productivity of nutrient‐rich vegetation types or increasing soil reaction of most vegetation types in more heterogeneous landscapes. We discuss available evidence and conclude that these two explanations need not be mutually exclusive.     

Occupancy frequency distributions: patterns,artefacts and mechanisms

Numerous hypotheses have been proposed to explain the shape of occupancy frequency distributions (distributions of the numbers of species occupying different numbers of areas). Artefactual effects include sampling characteristics, whereas biological mechanisms include organismal, niche-based and meta-population models. To date, there has been little testing of these models. In addition, although empirically derived occupancy distributions encompass an array of taxa and spatial scales, comparisons between them are often not possible because of differences in sampling protocol and method of construction. In this paper, the effects of sampling protocol (grain, sample number, extent, sampling coverage and intensity) on the shape of occupancy distributions are examined, and approaches for minimising artefactual effects recommended. Evidence for proposed biological determinants of the shape of occupancy distributions is then examined. Good support exists for some mechanisms (habitat and environmental heterogeneity), little for others (dispersal ability), while some hypotheses remain untested (landscape productivity, position in geographic range, range size frequency distributions), or are unlikely to be useful explanations for the shape of occupancy distributions 'species specificity and adaptation to habitat, extinction-colonization dynamics). The presence of a core (class containing species with the highest occupancy) mode in occupancy distributions is most likely to be associated with larger sample units, and small homogenous sampling areas positioned well within and towards the range centers of a sufficient proportion of the species in the assemblage. Satellite (class with species with the lowest occupancy) modes are associated with sampling large, heterogeneous areas that incorporate a large proportion of the assemblage range. However, satellite modes commonly also occur in the presence of a core mode, and rare species effects are likely to contribute to the presence of a satellite mode at most sampling scales. In most proposed hypotheses, spatial scale is an important determinant of the shape of the observed occupancy distribution. Because the attributes of the mechanisms associated with these hypotheses change with spatial scale, their predictions for the shape of occupancy distributions also change. To understand occupancy distributions and the mechanisms underlying them, a synthesis of pattern documentation and model testing across scales is thus needed. The development of null models, comparisons of occupancy distributions across spatial scales and taxa, documentation of the movement of individual species between occupancy classes with changes in spatial scale, as well as further testing of biological mechanisms are all necessary for an improved understanding of the distribution of species and assemblages within their geographic ranges.     

海南鹿回头岸礁造礁石珊瑚群落结构动态Ⅱ.物种多样性

根据石珊瑚物种的总数、石珊瑚覆盖的百分率、物种多样性和均匀度,对鹿回头岸礁造礁石珊瑚群落结构类型所处在演替阶段的时间状态和生境的空间状态进行分析。在中等和深水区物种多样性的时间变异,随演替的发展而有所下降;浅水区不随演替而变化。在不同生境区珊瑚生长的聚群上,与时间的发展无关;每个演替时期,物种多样性的类型与生境的变化有关。     

Fine‐scale heterogeneity in beetle assemblages under co‐occurring Eucalyptus in the same subgenus

Aim Insect biodiversity is often positively associated with habitat heterogeneity. However, this relationship depends on spatial scale, with most studies focused on differences between habitats at large scales with a variety of forest tree species. We examined fine‐scale heterogeneity in ground‐dwelling beetle assemblages under co‐occurring trees in the same subgenus: Eucalyptus melliodora A. Cunn. ex Schauer and E. blakelyi Maiden (Myrtaceae). Location Critically endangered grassy woodland near Canberra, south‐eastern Australia. Methods We used pitfall traps and Tullgren funnels to sample ground‐dwelling beetles from the litter environment under 47 trees, and examined differences in diversity and composition at spatial scales ranging from 100 to 1000 m. Results Beetle assemblages under the two tree species had distinctive differences in diversity and composition. We found that E. melliodora supported a higher richness and abundance of beetles, but had higher compositional similarity among samples. In contrast, E. blakelyi had a lower abundance and species richness of beetles, but more variability in species composition among samples. Main conclusions Our study shows that heterogeneity in litter habitat under co‐occurring and closely related eucalypt species can influence beetle assemblages at spatial scales of just hundreds of metres. The differential contribution to fine‐scale alpha and beta diversity by each eucalypt can be exploited for conservation purposes by ensuring an appropriate mix of the two species in the temperate woodlands where they co‐occur. This would help not only to maximize biodiversity at landscape scales, but also to maintain heterogeneity in species richness, trophic function and biomass at fine spatial scales.     

The importance of small-scale heterogeneity in boreal forests: variation in diversity in forest-floor invertebrates across the succession gradient

Heterogeneity in species assemblages of forest-floor arthropods — carabid beetles, ants and spiders — within and between different forest age classes was studied in the southern Finnish taiga. The importance of processes operating on the local scale (within the movement radius of the species) vs on the regional scale (among forest stands) in determining the observed variation was assessed. Four data sets with different spatial resolutions in mesic forests in the same general study area were used. The material consists of 18 283 carabids of 51 species, 48 769 spiders of 212 species, and 126 718 worker ants of 23 species. Analyses of abundance variation and species complementarity among successional stages revealed that in all the three taxa species occurring in the mature forest were prevalent in the younger successional stages as well, constituting more than half of the catch in any age class. A great majority of carabid and spider species were widely distributed across the forest age classes, whereas ants include a higher proportion of species with a narrower amplitude across the succession gradient. Comparisons of similarity between samples at increasing distance from one another on the local scale within forest stands (a few tens of meters to a few hundreds of meters) showed a quite consistent pattern in carabids and spiders: there was more variation between sampling sites in young successional forests than in the mature forest. Furthermore, only in the mature forest a slight, albeit statistically not significant, negative relationship between similarity of samples and distance between sampling sites was detected. In carabids and spiders, comparisons between samples located at a distance of 10–15 m from each other showed considerable heterogeneity, the mean percentage similarity being c. 0.6 (in ants c. 0.8). On the regional scale, systematic variation between young and mature forest stands is a major element increasing the total diversity (species turnover c. 50% in carabids and spiders; compositional similarity c. 0.3–0.4 in carabids, 0.2–0.3 in spiders), but variation within forest stands on a spatial scale of 10–15 meters is another important component in the total heterogeneity. The results suggest that regional abundance variation is a primary factor influencing the composition of local assemblages; a set of hypotheses elaborating this conclusion is formulated. The result implies that maintenance of habitat heterogeneity on a small scale (10–15 m) is needed to preserve biodiversity in managed forests.     

Spatial variability in intertidal macroalgal assemblages on the North Portuguese coast: consistence between species and functional group approaches

Natural assemblages are variable in space and time; therefore, quantification of their variability is imperative to identify relevant scales for investigating natural or anthropogenic processes shaping these assemblages. We studied the variability of intertidal macroalgal assemblages on the North Portuguese coast, considering three spatial scales (from metres to 10 s of kilometres) following a hierarchical design. We tested the hypotheses that (1) spatial pattern will be invariant at all the studied scales and (2) spatial variability of macroalgal assemblages obtained by using species will be consistent with that obtained using functional groups. This was done considering as univariate variables: total biomass and number of taxa as well as biomass of the most important species and functional groups and as multivariate variables the structure of macroalgal assemblages, both considering species and functional groups. Most of the univariate results confirmed the first hypothesis except for the total number of taxa and foliose macroalgae that showed significant variability at the scale of site and area, respectively. In contrast, when multivariate patterns were examined, the first hypothesis was rejected except at the scale of 10 s of kilometres. Both uni- and multivariate results indicated that variation was larger at the smallest scale, and thus, small-scale processes seem to have more effect on spatial variability patterns. Macroalgal assemblages, both considering species and functional groups as surrogate, showed consistent spatial patterns, and therefore, the second hypothesis was confirmed. Consequently, functional groups may be considered a reliable biological surrogate to study changes on macroalgal assemblages at least along the investigated Portuguese coastline.     

Changes in the structure and heterogeneity of vegetation and microsite environments with the chronosequence of primary succession on a glacier foreland in Ellesmere Island, high arctic Canada

Primary plant succession was investigated on a well-vegetated glacier foreland on Ellesmere Island in high arctic Canada. A field survey was carried out on four glacier moraines differing in time after deglaciation to assess vegetation development and microsite modification in the chronosequence of succession. The results showed evidence of directional succession without species replacement, which is atypical in the high arctic, reflecting the exceptionally long time vegetation development. During this successional process, Salix arctica dominated throughout all moraines. The population structures of S. arctica on these moraines implied the population growth of this species with progressing succession. The population density of S. arctica reflected the abundance of vascular plants, suggesting that development of the plant community might be related to structural changes and the growth of constituting populations. Through such growths of the population and the whole community with progressing succession, the spatial heterogeneity of vegetation gradually declines. Moreover, this vegetation homogenization is accompanied by changes in the spatial heterogeneity of microsite environments, suggesting significant plant effects on the modification of microsite environments. Accordingly, it was concluded that the directional primary succession observed on this glacier foreland is characterized by the initial sporadic colonization of plants, subsequent population growths, and the community assembly of vascular plants, accompanied by microsite modification.     

Biome transitions as centres of diversity: habitat heterogeneity and diversity patterns of West African bat assemblages across spatial scales

It is widely accepted that species diversity is contingent upon the spatial scale used to analyze patterns and processes. Recent studies using coarse sampling grains over large extents have contributed much to our understanding of factors driving global diversity patterns. This advance is largely unmatched on the level of local to landscape scales despite being critical for our understanding of functional relationships across spatial scales. In our study on West African bat assemblages we employed a spatially explicit and nested design covering local to regional scales. Specifically, we analyzed diversity patterns in two contrasting, largely undisturbed landscapes, comprising a rainforest area and a forest‐savanna mosaic in Ivory Coast, West Africa. We employed additive partitioning, rarefaction, and species richness estimation to show that bat diversity increased significantly with habitat heterogeneity on the landscape scale through the effects of beta diversity. Within the extent of our study areas, habitat type rather than geographic distance explained assemblage composition across spatial scales. Null models showed structure of functional groups to be partly filtered on local scales through the effects of vegetation density while on the landscape scale both assemblages represented random draws from regional species pools. We present a mixture model that combines the effects of habitat heterogeneity and complexity on species richness along a biome transect, predicting a unimodal rather than a monotonic relationship with environmental variables related to water. The bat assemblages of our study by far exceed previous figures of species richness in Africa, and refute the notion of low species richness of Afrotropical bat assemblages, which appears to be based largely on sampling biases. Biome transitions should receive increased attention in conservation strategies aiming at the maintenance of ecological and evolutionary processes.     

Density dependence is prevalent in a heterogeneous subtropical forest

Although negative conspecific density dependence among neighbours is widely studied, the general prevalence of the effects is still poorly understood due to a lack of studies from zonal forests other than the tropics. In addition, the detection of density dependence may be confounded by the influence of habitat heterogeneity. Here we examined the spatial distributions of 47 common tree species (diameter at breast height≥1  cm) using the pair-correlation function g(r) in a fully mapped 24-ha subtropical forest in China. We first investigated whether habitat heterogeneity influenced tree distributions, and then examined the conspecific tree patterns and density dependence after removing the effects of habitat heterogeneity. We found that the forest plot exhibited strong large-scale heterogeneity in the distribution of both large adult trees of different growth forms and individual species. After the habitat heterogeneity was accounted for, 39 of the 47 species (83.0%) were found to exhibit density dependence predominantly at close distances among neighbors. Our findings highlight density dependence as a prevalent mechanism for regulating the population spatial structure of most tree species in the species-rich subtropical forest studied here. Furthermore, the occurrence of density dependence is closely associated with species abundance and the strength of conspecific aggregation at local scales. Abundant species with high strength of conspecific aggregation tend to show density dependence.     

Spatial and seasonal patterns of microsite light availability in a remnant fragment of deciduous riparian forest and their implication in the conservation ofArisaema heterophyllum, a threatened plant species

Seasonal and spatial patterns of light availability were investigated in the understory of a small fragment (approximately 1300 m²) of a riparian deciduous forest of the Kokai River in central Japan dominated byQuercus acutissima Carruth., with the aim to understand the characteristics of microsite light availability forArisaema heterophyllum Blume, a threatened plant species uniquely associated with the riparian habitat. Diffuse site factor, which is the ratio of PFD (photon flux density, 400–700 nm) at a microsite to the open sky reference under diffuse light condition, was shown to be a satisfactory index for the evaluation of light availability for the understory plants in the habitat. Diffuse site factor 1,0 1,000 understory microsites along a 20 m transect from the edge to the interior of the forest, showed conspicuous seasonal changes in both mean and variation. Light availability decreased with seasonal tree canopy regeneration, with the highest spatial heterogeneity being recorded during the time of canopy closure. Auto-correlations of microsite light availability between different seasons were considerably high, suggesting the stability of relative light availability for individual microsites throughout the growing season ofA. heterophyllum. Fairly high light availability during summer season, which surpassed 20% of the open sky reference in most microsites, would be important for the growth and persistence of summer herbaceous plants likeA. heterophyllum.