Species richness, environmental heterogeneity and area: a case study based on land snails in Skyros archipelago (Aegean Sea, Greece)

Aim To test the performance of the choros model in an archipelago using two measures of environmental heterogeneity. The choros model is a simple, easy‐to‐use mathematical relationship which approaches species richness as a combined function of area and environmental heterogeneity. Location The archipelago of Skyros in the central Aegean Sea (Greece). Methods We surveyed land snails on 12 islands of the archipelago. We informed the choros model with habitat data based on natural history information from the land snail species assemblage. We contrast this with habitat information taken from traditional vegetation classification to study the behaviour of choros with different measures of environmental heterogeneity. R² values and Akaike's information criterion (AIC) were used to compare the choros model and the Arrhenius species–area model. Path analysis was used to evaluate the variance in species richness explained by area and habitat diversity. Results Forty‐two land snail species were recorded, living in 33 different habitat types. The choros model with habitat types had more explanatory power than the classic species–area model and the choros model using vegetation types. This was true for all islands of the archipelago, as well as for the small islands alone. Combined effects of area and habitat diversity primarily explain species richness in the archipelago, but there is a decline when only small islands are considered. The effects of area are very low both for all the islands of the archipelago, and for the small islands alone. The variance explained by habitat diversity is low for the island group as a whole, but significantly increases for the small islands. Main conclusions The choros model is effective in describing species‐richness patterns of land snails in the Skyros Archipelago, incorporating ecologically relevant information on habitat occupancy and area. The choros model is more effective in explaining richness patterns on small islands. When using traditional vegetation types, the choros model performs worse than the classic species–area relationship, indicating that use of proxies for habitat diversity may be problematic. The slopes for choros and Arrhenius models both assert that, for land snails, the Skyros Archipelago is a portion of a larger biogeographical province. The choros model, informed by ecologically relevant habitat measures, in conjunction with path analysis points to the importance of habitat diversity in island species richness.     

Habitat heterogeneity drives bird species richness,nestedness and habitat selection by individual species in fluvial wetlands of the Paraná River,Argentina

We assessed the relationship between habitat heterogeneity and bird species richness and composition within wetlands of the floodplain of the Middle Paraná River, Argentina. Given the high habitat heterogeneity in these wetland systems, we sought to determine whether (i) there was a positive relationship between bird species richness and habitat heterogeneity; (ii) whether bird species richness was associated with certain types of individual habitat types; (iii) whether there was a pattern of species nestedness and turnover between sites as a function of habitat heterogeneity and composition, respectively; and (iv) whether individual species exhibited associations with habitat heterogeneity. Point counts were used to survey birds at 60 sites. We estimated the area of eight habitat types found within a 200‐m radius from the centre of each site and calculated number and Pielou's evenness of habitat types. These indices, together with area proportion of each habitat type, were used as explanatory factors of bird species richness in linear regression models. Habitat heterogeneity per se rather than area of individual habitat types was a more important predictor of species richness in these fluvial wetlands. Sites with more habitat types supported more bird species. Results showed that individual bird species were associated with different habitat types and, therefore, sites that contained more habitat types contained more species. Number of habitat types accounted for species nestedness between sites whereas composition of habitat types accounted for species turnover between sites. Results suggest that selection of heterogeneous sites by individual species could help explain the positive heterogeneity–species richness relationship. Our findings highlight the importance of habitat heterogeneity per se resulting from flood disturbances in maintaining bird richness in fluvial systems.     

基于植物-地形关系的物种丰富度空间格局预测——GAMs途径的一种应用

将基于样本调查数据的群落-生境因子回归分析与GIS支持下的植物属性空间格局预测结合起来,是国际上植被-环境关系定量研究的新途径。通用可加性模型（GAM）的非参数属性使之具有对不同数据类型的广泛适应性,成为这种“回归分析＋空间预测”途经的有效手段;不同程度上依赖于数字高程模型的环境空间数据集是实现空间预测的必要条件。介绍了这一新的研究途径,并应用于案例研究区域植物多样性指标空间格局的预测和分析。野外调查的一组样方地形特征指标和植物多样性指标（包括样方物种丰富度及乔木、灌木、草本、常绿木本、珍稀种类的丰富度）,分别作为预测变量和响应变量,建立GAM模型。结合研究区域10m分辨率的数字高程模型,对该区域植物物种丰富度的空间格局进行空间预测,并对预测模型和结果进行统计分析和检验。结果表明：（1）不同的多样性指标具有不同的模型结构和模拟效果,重复模拟的结果稳定性也不同,反映了所受地形因子影响的差异;（2）影响各多样性指标空间格局的地形变量主要是坡位和坡度等小尺度特征,大尺度海拔因素的影响并不显著;（3）模拟结果与独立检验数据的相关分析表明,对乔木种、草本种、珍稀种的模拟全部有效;对常绿种和样方物种总数的模拟部分有效;而对灌木种丰富度的预测基本失败。（4）模型预测变量有效性和全面性决定了模型对数据的解释能力,样本大小对模型的稳定性和可靠性也有显著影响。就地形因子对生境条件的代表性、模拟误差的来源及GAMs模型的优缺点和应用前景进行了讨论。     

Relative importance of isolation, area and habitat heterogeneity for vascular plant species richness of temporary wetlands in east-German farmland

The purpose of this investigation was to explore the determinants of vascular plant species richness for temporary, isolated wetland habitats which are influenced by hydrologic gradients and characterized by variation in habitat area. The dependent variables total plant species richness and the number of obligate wetland species were analyzed consecutively. In regression analyses habitat area explained between 11 and 15% of the variation in the dependent variables. Habitat area was correlated with the heterogeneity of the hydroperiods between the upper and lower parts of the hydrologic gradients. In multivariate regression analyses, habitat heterogeneity accounted for 70-77% of the variation in the dependent variables, and habitat area did not have a significant impact. The results are most consistent with the habitat heterogeneity hypothesis. I therefore concluded that area is a surrogate variable for habitat heterogeneity which directly enhances vascular plant species diversity. There was no significant impact of isolation on species richness. The data suggest that the expanses of agricultural fields are not an effective barrier to the dispersal of the studied plant species. Only 10 of,52 wetland species were negatively influenced by isolation. This group of species did not differ from the other wetland species with respect to dispersal strategies and longevity of seed banks. However, the longevity of the seed banks was generally high, and there was a dominance of species whose propagules are transported with the soil clinging to the feet of birds. The results are discussed in the context of accurate dispersal strategies and remnant populations, which may counteract the effects of isolation.     

Patterns of species richness for vascular plants in China's nature reserves

Explaining the heterogeneous distribution of biodiversity across the Earth has long been a challenge to ecologists and biogeographers. Here, we document the patterns of plant species richness for different taxonomic groups in China's nature reserves, and discuss their possible explanations at national and regional scales, using vascular plant richness data coupled with information on climate and topographical variables. We found that water deficit, energy and elevation range (a surrogate of habitat heterogeneity) represent the primary explanations for variation in plant species richness of the nature reserves across China. There are consistent relationships between species richness and climate and habitat heterogeneity for different taxonomic vascular plant groups at the national scale. Habitat heterogeneity is strongly associated with plant richness in all regions, whereas climatic constraints to plant diversity vary regionally. In the regions where energy is abundant or water is scarce, plant richness patterns were determined by water and habitat heterogeneity, whereas in the region with low energy inputs, water interacting with energy, and habitat heterogeneity determined its species richness pattern. Our results also suggest that energy variables alone do not represent the primary predictor of plant richness.     

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.     

Drivers of symbiont diversity in freshwater snails: a comparative analysis of resource availability,community heterogeneity,and colonization opportunities

Decades of community ecology research have highlighted the importance of resource availability, habitat heterogeneity, and colonization opportunities in driving biodiversity. Less clear, however, is whether a similar suite of factors explains the diversity of symbionts. Here, we used a hierarchical dataset involving 12,712 freshwater snail hosts representing five species to test the relative importance of potential factors in driving symbiont richness. Specifically, we used model selection to assess the explanatory power of variables related to host species identity, resource availability (average body size, host density), ecological heterogeneity (richness of hosts and other taxa), and colonization opportunities (wetland size and amount of neighboring wetland area) on symbiont richness in 146 snail host populations in California, USA. We encountered a total of 23 taxa of symbionts, including both obligatory parasites such as digenetic trematodes as well as more commensal, mutualistic, or opportunistic groups such as aquatic insect larvae, annelids, and leeches. After validating richness estimates per host population using species accumulative curves, we detected positive effects on symbiont richness from host body size, total richness of the aquatic community, and colonization opportunities. Neither snail density nor the richness of snail species accounted for significant variation in symbiont diversity. Host species identity also affected symbiont richness, with higher gamma and average alpha diversity among more common host species with higher local abundances. These findings highlight the importance of multiple, concurrent factors in driving symbiont richness that extend beyond epidemiological measures of host abundance or host diversity alone.     

The unusual suspect: Land use is a key predictor of biodiversity patterns in the Iberian Peninsula

Although land use change is a key driver of biodiversity change, related variables such as habitat area and habitat heterogeneity are seldom considered in modeling approaches at larger extents. To address this knowledge gap we tested the contribution of land use related variables to models describing richness patterns of amphibians, reptiles and passerines in the Iberian Peninsula. We analyzed the relationship between species richness and habitat heterogeneity at two spatial resolutions (i.e., 10 km × 10 km and 50 km × 50 km). Using both ordinary least square and simultaneous autoregressive models, we assessed the relative importance of land use variables, climate variables and topographic variables. We also compare the species–area relationship with a multi-habitat model, the countryside species–area relationship, to assess the role of the area of different types of habitats on species diversity across scales. The association between habitat heterogeneity and species richness varied with the taxa and spatial resolution. A positive relationship was detected for all taxa at a grain size of 10 km × 10 km, but only passerines responded at a grain size of 50 km × 50 km. Species richness patterns were well described by abiotic predictors, but habitat predictors also explained a considerable portion of the variation. Moreover, species richness patterns were better described by a multi-habitat species-area model, incorporating land use variables, than by the classic power model, which only includes area as the single explanatory variable. Our results suggest that the role of land use in shaping species richness patterns goes beyond the local scale and persists at larger spatial scales. These findings call for the need of integrating land use variables in models designed to assess species richness response to large scale environmental changes.     

A multi‐scale analysis of fragmentation effects on remnant plant species richness in Phoenix,Arizona

Aim Understanding complex ecological phenomena, such as the determinants of species richness, is best achieved by investigating their properties at different spatial scales. Factors significantly affecting the number of species occurring at one scale may not impact on richness at other scales. While this scale dependence has become increasingly recognized, there still remains a need to elucidate exactly how richness is structured across scales, and which mechanisms are influential for determining this important community property. This study explores how woody plant species richness varies in a fragmented system at multiple scales, and which factors are primarily responsible for these patterns. Location The study area is located in the Sonoran Desert within the bounds of metropolitan Phoenix, Arizona, which is the locus of the Central Arizona–Phoenix Long‐Term Ecological Research (CAP‐LTER) site. Methods Estimates of local and fragment plant species richness were generated from field data collected from 22 sites. Independent variables describing fragment sites were also calculated, including area, habitat heterogeneity, density of individuals, mean elevation, and extent of isolation. Structural equation modelling, multiple regression, and analysis of covariance were used to assess the contribution of independent variables to richness at the fragment and local scales. Results Fragment species richness was significantly influenced by area, though not isolation, habitat heterogeneity, mean elevation, or density of individuals. Local richness was not significantly related to fragment area, but was positively related to fragment richness, plant density, and elevation. Main conclusions The fragment species–area effect resulted from larger remnants supporting higher numbers of individuals at comparable densities, increasing richness through either passive sampling of progressively less common species and/or lower extinction rates among larger populations. Without using multi‐temporal data it is not possible to disentangle these mechanisms. We found that patterns evident at one scale are not necessarily apparent at other scales, as elevation and density of individuals significantly affected richness at the local scale but not at the fragment scale. These results lend support to the concept that mechanisms influencing the species richness of natural communities may be operable only within certain domains and that relevant scales should be specified.     

Influence of area,habitat and water chemistry on richness and composition of macrophyte assemblages in southern Brazilian wetlands

Questions: Two hypotheses were tested: (1) physical features, such as wetland surface area and habitat diversity, together with water chemistry, are important determinants of species richness and composition of macrophyte assemblages and (2) species richness and composition of macrophyte assemblages differ between wetlands of different types (i.e., palustrine versus lacustrine) and between wetlands of different hydrologies (i.e. permanent versus intermittent). Location: A subtropical coastal plain segment (2500 km²) of southern Brazil. Methods: Quarterly collections were carried out in 15 wetlands (2004–2005) in southern Brazil. Differences in richness over time were tested using repeated measures ANOVA. Stepwise multiple regression was performed to investigate relationships between total richness and environmental variables. Significance of differences between wetland types and hydroperiods on species composition was verified by MRPP (Multi‐Response Permutation Procedure). The influence of the environmental variables on species composition was assessed using CCA (Canonical Correspondence Analysis). Results: Macrophyte species richness changed with time, was not significantly different between wetland types, but was higher in permanent wetlands than in intermittent ones. Area, habitat diversity and soluble reactive phosphorus concentration explained 76% of the variation in species richness. Species composition was different between permanent and intermittent wetlands, although it was not significantly different between wetland types. Area, habitat diversity and water chemistry explained 50.1% of species composition. Conclusions: Species richness and composition of wetland macrophytes were mainly determined by area, habitat diversity and hydroperiod. These results can be used for the development of conservation and management programs in southern Brazil.     

松嫩平原不同生境蝴蝶群落多样性及其保护建议

随着气候变化加剧和人类活动影响,生物多样性变化及其保护逐渐受到广泛关注。蝴蝶作为开花植物的传粉媒介和生态环境监测及评价的关键指示者,其多样性变化能够在一定程度上反映生境状况,因此,有必要清晰认识不同生境中的蝴蝶多样性变化。为明确松嫩平原蝴蝶资源和不同生境的群落多样性差异,采用样线法于2016年5月-2018年8月对松嫩平原的割草草地、湿地、农田、放牧利用草地及恢复草地共五种生境类型进行调查研究。结果发现,调查共记录蝴蝶5108头,隶属于6科21属26种,其中牧女珍眼蝶（Coenonympha amaryllis）和红珠灰蝶（Plebejus argyrognomon）为优势种类,分别占蝴蝶个体总数的25.61%和31.66%,且在五种生境类型中均有分布。不同生境类型中,蝴蝶群落的物种丰富度指数和均匀度指数无明显差异,而恢复草地生境的蝴蝶群落Shannon-Wiener多样性指数较高,优势度指数较低。农田生境中的蝴蝶个体数量较少,且群落组成与其他四种生境之间均具有显著差异。五种生境类型中的蝴蝶数量和多样性均呈现一定的月动态和年动态变化趋势。除湿地和农田外,其余三种生境中蝴蝶物种和个体数量从5月到8月均持续升高。四种生境的蝴蝶物种数量、个体数量（除农田外）在2018年均出现明显下降趋势。物种丰富度指数等指标的月动态和年动态在不同生境类型间存在较大差异。这些结果表明,生境类型和人类活动与蝴蝶多样性变化关系密切,表现为单一生境中蝴蝶多样性较低,复杂生境有利于保护蝴蝶多样性。本研究有助于厘清松嫩平原蝴蝶资源的基础数据,并为该地区蝴蝶多样性保护和利用及评估该区域生态环境提供一定理论支撑。     

新疆开都河流域水生植物多样性及其影响因素

河流是一个连续的、流动的、独特而完整的系统,研究河流生态系统中水生植物的多样性分布格局及其影响因素对河流生态学研究具有重要意义。本文通过野外调查,研究了新疆开都河流域水生植物多样性、主要水生植物群落特征及与环境因子之间的关系,并利用水分-能量动态假说和栖息地异质性假说对该流域水生植物物种多样性的地理格局进行解释。结果表明: 开都河流域共有水生植物71种,隶属于24科39属;聚类分析可将开都河流域水生植物群落划分为10个主要群落类型,其中芦苇群落物种丰富度最高,狭叶香蒲群落和金鱼藻群落物种丰富度最低;流域水生植物群落Shannon指数与pH呈显著负相关,Simpson指数与pH、经度呈显著负相关,与海拔呈显著正相关;流域水生植物群落类型主要受海拔、水深及水温的影响;流域水生植物物种多样性随经纬度无明显变化规律。水分-能量动态假说和栖息地异质性假说共解释开都河流域水生植物多样性格局变量的31.4%,表明这两个假说对于该流域水生植物多样性格局的解释力并不高。     

Richness and composition of plants and birds on land‐bridge islands: effects of island attributes and differential responses of species groups

Aim To test relationships between the richness and composition of vascular plants and birds and attributes of habitat fragments using a model land‐bridge island system, and to investigate whether the effects of fragmentation differ depending on species natural history traits. Location Thousand Island Lake, China. Methods We compiled presence/absence data of vascular plant and bird species through exhaustive surveys of 41 islands. Plant species were assigned to two categories: shade‐intolerant and shade‐tolerant species; bird species were assigned to three categories: edge, interior, and generalist species. We analysed the relationships between island attributes (area, isolation, elevation, shape complexity, and perimeter to area ratio) and species richness using generalized linear models (GLMs). We also investigated patterns of composition in relation to island attributes using ordination (redundancy analysis). Results We found that island area explained a high degree of variation in the species richness of all species groups. The slope of the species–area relationship (z) was 0.16 for all plant species and 0.11 for all bird species. The lowest z‐value was for generalist birds (0.04). The species richness of the three plant species groups was associated with island area per se, while that of all, generalist, and interior birds was explained mainly by elevation, and that of edge bird species was associated primarily with island shape. Patterns of species composition were most strongly related to elevation, island shape complexity, and perimeter to area ratio rather than to island area per se. Species richness had no significant relationship with isolation, but species composition did. We also found differential responses among the species groups to changes in island attributes. Main conclusions Within the Thousand Island Lake system, the effects of fragmentation on both bird and plant species appear to be scale‐dependent and taxon‐specific. The number of plant species occurring on an island is strongly correlated with island area, and the richness of birds and the species composition of plants and birds are associated with variables related to habitat heterogeneity. We conclude that the effects of fragmentation on species diversity and composition depend not only on the degree of habitat loss but also on the specific patterns of habitat fragmentation.     

Importance of semi-natural habitats for the conservation of butterfly communities in landscapes dominated by pine plantations

While the area of plantation forests continues to increase worldwide, their contribution to the conservation of biodiversity is still controversial. There is a particular concern on the central role played by natural habitat remnants embedded within the plantation matrix in conserving species-rich insect communities. We surveyed butterflies in maritime pine plantation landscapes in south-western France in 83 plots belonging to seven habitat types (five successional stages of pine stands, native deciduous woodlands and herbaceous firebreaks). The effect of plot, habitat and landscape attributes on butterfly species richness, community composition and individual species were analysed with a General Linear Model (GLM), partial Canonical Correspondence Analysis (CCA) and the IndVal method. The most important factors determining butterfly diversity and community composition were the presence of semi-natural habitats (deciduous woodlands and firebreaks) at the landscape scale and the composition of understorey vegetation at the plot scale. Pure effects of plot variables explained the largest part of community variation (12.8%), but landscape factors explained an additional, independent part (6.7%). Firebreaks were characterized by a higher species richness and both firebreaks and deciduous woodlands harboured species not or rarely found in pine stands. Despite the forest-dominated landscape, typical forest butterflies were rare and mainly found in the deciduous woodlands. Threatened species, such as Coenonympha oedippus and Euphydryas aurinia, were found in pine stands and in firebreaks, but were more abundant in the latter. In the studied plantation forest, the conservation of butterflies depends mainly on the preservation of semi-natural habitats, an adequate understorey management and the maintenance of soil moisture levels.     

Ants and people: a test of two mechanisms potentially responsible for the large‐scale human population–biodiversity correlation for Formicidae in Europe

Aim: Recent coarse‐scale studies have shown positive relationships between the biodiversity of plants/vertebrates and the human population. Little is known about the generality of the pattern for invertebrates. Moreover, biodiversity and human population might correlate because they both covary with other factors such as energy availability and habitat heterogeneity. Here we test these two non‐mutually exclusive mechanisms with ant species‐richness data from the Fauna Europaea. Location Forty‐three European countries/regions. Methods We derived mixed models of total, native and exotic ant species richness as a function of human population size/density, controlling for country area, plant species richness (as a proxy for habitat heterogeneity), and mean annual temperature and precipitation (variables related to energy availability). Results Ant species richness increased significantly with increasing human population. This result was confirmed when controlling for variations in country area. Both for human population size/density and for ant species richness, there were positive correlations with temperature but not with precipitation. This finding is in agreement with the energy‐availability hypothesis. However, we observed a negative latitudinal gradient in ant and plant species richness, although not in human population size/density. Plant species richness was positively correlated with ant species richness but not with human population size/density. Thus, there is evidence that this type of habitat heterogeneity can play a role in the observed latitudinal gradient of ant species richness, but not in the positive correlation between ant species richness and human population. The results were confirmed for the 545 native and the 32 exotic ant species reported, and we observed a good correlation between exotic and native ant species richness. Main conclusions Ant species richness in European countries conforms to six macroecological patterns: (1) a negative latitudinal gradient; and a positive (2) species–energy relationship, (3) species–area relationship, (4) correlation with plant species richness, (5) exotic–native species richness correlation, and (6) species–people correlation. There is some evidence for the energy‐availability hypothesis, but little evidence for habitat heterogeneity as an explanation of the large‐scale human population–ant biodiversity correlation. This correlation has implications for the conservation of ant diversity in Europe.     

Breeding bird diversity in relation to environmental gradients in China

Geographic variation in species richness has been explained by different theories such as energy, productivity, energy–water balance, habitat heterogeneity, and freezing tolerance. This study determines which of these theories best account for gradients of breeding bird richness in China. In addition, we develop a best-fit model to account for the relationship between breeding bird richness and environment in China. Breeding bird species richness in 207 localities (3271 km² per locality on average) from across China was related to thirteen environmental variables after accounting for sampling area. The Akaike's information criterion (AIC) was used to evaluate model performance. We used Moran's I to determine the magnitude of spatial autocorrelation in model residuals, and used simultaneous autoregressive model to determine coefficients of determination and AIC of explanatory variables after accounting for residual spatial autocorrelation. Of all environmental variables examined, normalized difference vegetation index, a measure of plant productivity, is the best variable to explain the variance in breeding bird richness. We found that species richness of breeding birds at the scale examined is best predicted by a combination of plant productivity, elevation range, seasonal variation in potential evapotranspiration, and mean annual temperature. These variables explained 47.3% of the variance in breeding bird richness after accounting for sampling area; most of the explained variance in richness is attributable to the first two of the four variables.     

Avifaunal responses to habitat fragmentation in the threatened littoral forests of south-eastern Madagascar

Aim Madagascar's lowland forests are both rich in endemic taxa and considered to be seriously threatened by deforestation and habitat fragmentation. However, very little is known about how these processes affect biodiversity on the island. Herein, we examine how forest bird communities and functional groups have been affected by fragmentation at both patch and landscape scales, by determining relationships between species richness and individual species abundance and patch and landscape mosaic metrics. Location Littoral forest remnants within south‐eastern Madagascar. Methods We sampled 30 littoral forest remnants in south‐eastern Madagascar, within a landscape mosaic dominated by Erica spp. heathland. We quantified bird community composition within remnants of differing size, shape and isolation, by conducting point counts in November–December in 2001 and October–November 2002. Each remnant was characterized by measures of remnant area, remnant shape, isolation, and surrounding landscape complexity. We used step‐wise regression to test the relationship between bird species richness and landscape structural elements, after correcting for sampling effort. Relationships between bird species abundances and the landscape variables were investigated with Canonical Correspondence Analysis and binomial logistic regression modelling. Results Bird species richness and forest‐dependent bird species richness were significantly (P < 0.01) explained by remnant area but not by any measure of isolation or landscape complexity. The majority of forest‐dependent species had significant relationships with remnant area. Minimum area requirements for area‐sensitive species ranged from 15 to 150 ha, with the majority of species having area requirements > 30 ha. Surprisingly, there was no relationship between bird body size and minimum area requirement. Forest‐dependent canopy insectivorous species and large canopy frugivorous species were the most sensitive functional groups, with > 90% species sensitivity within each group. The distribution of four forest‐dependent species also appeared to be related to remnant shape where remnant area was < 100 ha. Main conclusions The majority of forest‐dependent species, including many that are considered widespread and common, were found to have significant relationships with fragment size, indicating that they are sensitive to processes associated with habitat loss and fragmentation. As deforestation and habitat fragmentation remain serious problems on the island, it follows that many forest‐dependent bird species will decline in abundance and become locally extinct. At the regional scale, we urge that large (> 200 ha) blocks of littoral forest are awarded protected status to preserve their unique bird community.     

Environmental effects on vertebrate species richness: testing the energy, environmental stability and habitat heterogeneity hypotheses

Background

Explaining species richness patterns is a central issue in biogeography and macroecology. Several hypotheses have been proposed to explain the mechanisms driving biodiversity patterns, but the causes of species richness gradients remain unclear. In this study, we aimed to explain the impacts of energy, environmental stability, and habitat heterogeneity factors on variation of vertebrate species richness (VSR), based on the VSR pattern in China, so as to test the energy hypothesis, the environmental stability hypothesis, and the habitat heterogeneity hypothesis.

Methodology/Principal Findings

A dataset was compiled containing the distributions of 2,665 vertebrate species and eleven ecogeographic predictive variables in China. We grouped these variables into categories of energy, environmental stability, and habitat heterogeneity and transformed the data into 100×100 km quadrat systems. To test the three hypotheses, AIC-based model selection was carried out between VSR and the variables in each group and correlation analyses were conducted. There was a decreasing VSR gradient from the southeast to the northwest of China. Our results showed that energy explained 67.6% of the VSR variation, with the annual mean temperature as the main factor, which was followed by annual precipitation and NDVI. Environmental stability factors explained 69.1% of the VSR variation and both temperature annual range and precipitation seasonality had important contributions. By contrast, habitat heterogeneity variables explained only 26.3% of the VSR variation. Significantly positive correlations were detected among VSR, annual mean temperature, annual precipitation, and NDVI, whereas the relationship of VSR and temperature annual range was strongly negative. In addition, other variables showed moderate or ambiguous relations to VSR.

Conclusions/Significance

The energy hypothesis and the environmental stability hypothesis were supported, whereas little support was found for the habitat heterogeneity hypothesis.     

Bird communities across varying landcover types in a Neotropical city

Urbanization poses a serious threat to local biodiversity, yet towns and cities with abundant natural features may harbor important species populations and communities. While the contribution of urban greenspaces to conservation has been demonstrated by numerous studies within temperate regions, few consider the bird communities associated with different landcovers in Neotropical cities. To begin to fill this knowledge gap, we examined how the avifauna of a wetland city in northern Amazonia varied across six urban landcover types (coastal bluespace; urban bluespace; managed greenspace; unmanaged greenspace; dense urban; and sparse urban). We measured detections, species richness, and a series of ground cover variables that characterized the heterogeneity of each landcover, at 114 locations across the city. We recorded >10% (98) of Guyana's bird species in Georgetown, including taxa of conservation interest. Avian detections, richness, and community composition differed with landcover type. Indicator species analysis identified 29 species from across dietary guilds, which could be driving community composition. Comparing landcovers, species richness was highest in managed greenspaces and lowest in dense urban areas. The canal network had comparable levels of species richness to greenspaces. The waterways are likely to play a key role in enhancing habitat connectivity as they traverse densely urbanized areas. Both species and landcover information should be integrated into urban land-use planning in the rapidly urbanizing Neotropics to maximize the conservation value of cities. This is imperative in the tropics, where anthropogenic pressures on species are growing significantly, and action needs to be taken to prevent biodiversity collapse.     

A functional relationship between species richness of spiders and lichens in spruce

Modern forestry has created stands with even age distribution of trees and fragmentation of the habitat. In boreal forests, the effects on biodiversity within many taxa need to be examined. We tested the hypothesis that species richness of foliose and fruticose lichens and spiders is positively related in the lower canopy of spruce (Picea abies) in forests with, or without, management in central Sweden. High species richness of lichens may increase the structural complexity of the microhabitat on spruce branches, and bring a higher species richness also in the spider community. In six areas, spruce branches were sampled in old-growth and managed boreal forest stands, respectively. Forest management did not affect the species richness of spiders or lichens, but an effect due to sampling area was found in the latter taxon. There was a significant covariation between species richness of lichens and spiders, and the hypothesised positive correlation was confirmed by separate analyses for each area and combining the probabilities. Moreover, regression analysis on mean values from each site revealed a positive relationship. We conclude that species richness of lichens and spiders covary on spruce branches for functional reasons, i.e. more lichen species promotes a more diverse spider community by increasing the structural heterogeneity. Our results might provide a shortcut for assessing biodiversity in boreal forests.