Alien and native birds in South Africa: patterns, processes and conservation

The spatial distribution of alien species richness often correlates positively with native species richness, and reflects the role of human density and activity, and primary productivity and habitat heterogeneity, in facilitating the establishment and spread of alien species. Here, we investigate the relationship between the spatial distribution of alien bird species, human density, and anthropogenic and natural environmental conditions. Next, we examined the relationship between the spatial distribution of alien bird species and native bird species richness. We examined alien species richness as a response variable, using correlative analyses that take spatial autocorrelation into account. Further, each alien bird species was examined as a response variable, using logistic regression procedures based on binary presence–absence data. A combination of human density and natural habitat heterogeneity best explained the spatial distribution of alien species richness. This contrasts with the results for individual alien species and with previous studies on other non-native taxa showing the importance of primary productivity and anthropogenic habitat modification as explanatory variables. In general, native species richness is an important correlate of the spatial distribution of alien species richness and individual alien species, with alien species being more similar to common species than to rare species.     

How,and how much,natural cover loss increases species richness

Aim The species–area relationship has been applied in the conservation context to predict monotonic species richness declines as natural area is converted to human‐dominated land covers. However, some conversion of natural cover could introduce new habitat types and allow new open habitat species to occur. Moreover, decelerating richness–area relationships suggest that, as natural area is converted to human‐dominated covers, more species will be added to the rare habitat than are lost from the common one. Area effects and increased habitat diversity could each lead to a peaked relationship between species richness and the relative amount of natural area. The purpose of this study is to quantify the effect on avian species richness of conversion of natural area to human‐dominated land cover. Location Ontario, Canada. Methods We evaluated the responses of total avian richness, forest bird richness and open habitat bird richness to remaining natural area within 993 quadrats, each of 100 km². We quantified the amount of natural land cover and land‐cover heterogeneity using remote sensing data. We used structural equation modelling (SEM) to disentangle the relationships among avian richness, natural area and land‐cover heterogeneity. Results Spatial variation in avian richness was a peaked function of remaining natural area, such that losses of up to 44% of the natural area increased avian richness. This partly reflects increased variety of land cover; however, SEM suggests that much of the increase in richness is due to pure area effects. Richness of forest species declined by two species over this range of natural cover loss while open habitat bird richness increased by approximately 20 species. The effect of natural area on species richness is consistent with the sum of species–area curves for natural habitat species and human‐dominated habitat species. Main conclusions At least in northern temperate forests, almost half of the natural land cover can be converted to human‐dominated forms before avian richness declines. Conversion of < 50% of regional natural area to human‐dominated land cover can benefit open‐area species richness with relatively few losses of forest obligate species. However, with > 50% natural area conversion, species begin to drop out of regional assemblages.     

What drives the positive correlation between human population density and bird species richness in Australia?

Aim To test six hypotheses that could explain or mediate the positive correlation between human population density (HPD) and bird species richness while controlling for biased sampling effort. These hypotheses were labelled as follows: productivity (net primary productivity, NPP); inherent heterogeneity (diversity of vegetation types); anthropogenic heterogeneity (diversity of land uses); conservation policy (proportion of conservation land); increased productivity (human‐induced productivity increases); and the reduced‐slope hypothesis (which predicts that humans have a negative impact on species numbers across the full range of variation in HPD). Location Australia. Methods All data were collected at a spatial resolution of 1° across mainland Australia. Bird species richness was from 2007 atlas data and random subsampling was used to account for biased sampling effort. HPD was from the 2006 census. All other data were from government produced geographic information system layers. The most important biotic or abiotic factors influencing patterns in both species richness and HPD were assessed using simultaneous autoregressive models and an information theoretic approach. Results NPP appeared to be one of the main factors driving spatial congruence between bird species richness and HPD. Inherent habitat heterogeneity was weakly related to richness and HPD, although an interaction between heterogeneity and NPP indicated that the former may be an important determinant of species richness in low‐productivity regions. There was little evidence that anthropogenic landscape heterogeneity or human‐induced changes in productivity influenced the relationship between species richness and HPD, but conservation policy appeared to act as an important mediating factor and species richness was positively related to the proportion of conservation land only in regions of high HPD. Main conclusions The spatial congruence between bird species richness and HPD occurs because both respond positively to productivity and, in certain circumstances, habitat heterogeneity. Our results suggest that conservation policy could mediate this relationship, but further research is required to determine the importance of conservation reserves in supporting species in regions densely populated by humans.     

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.     

Habitat heterogeneity overrides the species–area relationship

Aim The most obvious, although not exclusive, explanation for the increase of species richness with increasing sample area (the species–area relationship) is that species richness is ultimately linked to area-based increases in habitat heterogeneity. The aim of this paper is to examine the relative importance of area and habitat heterogeneity in determining species richness in nature reserves. Specifically, the work tests the hypothesis that species–area relationships are not positive if habitat heterogeneity does not increase with area. Location Sixteen nature reserves (area range 89–11,030 ha) in central Hungary. Methods Four-year faunistic inventories were conducted in the reserves involving c. 70 fieldworkers and 65 taxonomists. CORINE 50,000 land-cover maps were used for calculating the heterogeneity of the reserve landscape (number of habitat types, number of habitat patches and total length of edges). Results Large reserves were less heterogeneous than small reserves, probably because large reserves were established in large blocks of unproductive land whereas small reserves tended to be in more fertile land. In total, 3975 arthropod species were included in the analysis. The slope of the species–area relationship was positive only for Neuroptera and Trichoptera. There was no significant relationship in the other nine taxa examined (Collembola, Acari, Orthoptera, Thysanoptera, Coleoptera, Araneae, Diplopoda, Chilopoda, Diptera). The density (number of species ha⁻¹) of all species, however, showed a positive correlation with heterogeneity. Main conclusions The general lack of fit of species–area relationships in this study is inconsistent with most previous published studies. Importantly, and unlike many other studies, habitat heterogeneity was not correlated with reserve area in the studied system. In the absence of this source of covariation, stronger relationships were identified that suggested a fundamental link between species richness and habitat heterogeneity. The results indicate that habitat heterogeneity rather than area per se is the most important predictor of species richness in the studied system.     

Bird species richness patterns of northern Taiwan: primary productivity, human population density, and habitat heterogeneity

Energy, climate, habitat heterogeneity, and human activity are important correlates of spatial variation in species richness. We examined the correlation between species richness and these variables using the birds that breed in northern Taiwan. We conducted general linear models (GLMs) and spatial correlation models to examine the relationship between bird species richness (BSR) and environmental variables. We found that normalized difference vegetation index (NDVI) was the most important predictor of BSR. We suggest productivity is the primary process of BSR. Additionally, we hypothesized that scale dependency might exist in the relationship between BSR and NDVI in Taiwan. Human population density, the second most important factor, was inversely correlated with BSR. The factor and BSR did not have similar response to NDVI, which contradicted observations in most of the previous studies on human population vs. species richness. We proposed that the human population density had an effect on NDVI, which in turn had an effect on BSR. Moreover, we hypothesized that the contradiction between our study and the previous studies might arise from a higher level of human disturbance in Taiwan than in other areas. The necessity of conserving native species in intensively developed lowlands of Taiwan cannot be overemphasized. Number of land cover type was another significant predictor of BSR. Habitat heterogeneity may have an effect on BSR in Taiwan.     

Human impacts on environment–diversity relationships: evidence for biotic homogenization from butterfly species richness patterns

Aim Broad‐scale spatial variation in species richness relates to climate and physical heterogeneity but human activities may be changing these patterns. We test whether climate and heterogeneity predict butterfly species richness regionally and across Canada and whether these relationships change in areas of human activity. Location Canada. Methods We modelled the ranges of 102 butterfly species using genetic algorithms for rule‐set production (GARP). We then measured butterfly species richness and potentially important aspects of human activity and the natural environment. These were included in a series of statistical models to determine which factors are likely to affect butterfly species richness in Canada. We considered patterns across Canada, within predominantly natural areas, human‐dominated areas and particular ecozones. We examined independent observations of butterfly species currently listed under Canada's endangered species legislation to test whether these were consistent with findings from statistical models. Results Growing season temperature is the main determinant of butterfly species richness across Canada, with substantial contributions from habitat heterogeneity (measured using elevation). Only in the driest areas does precipitation emerge as a leading predictor of richness. The slope of relationships between all of these variables and butterfly species richness becomes shallower in human‐dominated areas, but butterfly richness is still highest there. Insecticide applications, habitat loss and road networks reduce butterfly richness in human‐dominated areas, but these effects are relatively small. All of Canada's at‐risk butterfly species are located in these human‐dominated areas. Main conclusions Temperature affects butterfly species richness to a greater extent than habitat heterogeneity at fine spatial scales and is generally far more important than precipitation, supporting both the species richness–energy and habitat heterogeneity hypotheses. Human activities, especially in southern Canada, appear to cause surprisingly consistent trends in biotic homogenization across this region, perhaps through range expansion of common species and loss of range‐restricted species.     

Determinants of species‐area relationships for marsh‐nesting birds

ABSTRACT To clarify the underlying causes of the species‐area relationship in marsh‐nesting birds, I studied eight freshwater tidal marshes of the Connecticut River that differed in area, degree of isolation, mudflat cover, water cover, tidal regime, and extent of individual plant communities. I measured these habitat variables on aerial infrared photos, and surveyed bird populations by mapping the distribution of all birds in marshes under 5 ha in area and establishing 50‐m radius plots in marshes over 5 ha. From surveys, I determined species richness, population densities, and total populations. Analysis revealed a positive relationship between species richness and area, but no correlation between area and habitat heterogeneity. Other habitat variables were poor predictors of species richness. The lack of a relationship between habitat and species richness appeared to be a consequence of most vegetation types present not being sufficiently distinct for birds to differentially associate with them. I also found no relationship between bird population density and area, suggesting that habitat quality in marshes did not improve with increasing size, and species evenness declined with increasing richness because greater richness was associated with the presence of more rare species. Larger marshes had more rare species, species with larger populations, and species with a minimum threshold area for occurrence. Thus, my results are consistent with theoretical predictions that larger populations are less prone to local extinction and, as individuals are added to a community, more rare species are present.     

Top-down pressure by generalist and specialist natural enemies in relation to habitat heterogeneity and resource availability

Habitat heterogeneity might promote the abundance and richness of natural enemies potentially leading to higher top-down pressure on herbivorous insects. Heterogeneous habitats could provide natural enemies with more abundant and alternative resources and a greater variety of micro-habitats. Natural enemies with different searching behaviours, e.g. generalists and specialists, could be affected in different ways by habitat heterogeneity, thus affecting their pressure on herbivorous insects.To understand how top-down pressure on herbivorous insects is promoted by habitat heterogeneity, it is crucial to investigate which parameters contributing to habitat heterogeneity affect not only the abundance and richness but also the searching behaviour of different natural enemies. We investigated the relationship between heterogeneity in forest habitats and the top-down pressure exerted by generalist predators and specialist parasitoids on larvae of the European pine sawfly (Neodiprion sertifer).We used forest stands with endemic or epidemic densities of resident sawfly populations. Within each stand we selected experimental trees to create variation in tree species diversity and density in their surrounding area, i.e. habitat heterogeneity. We found that a higher tree density increased the predation by generalists on sawfly larvae in stands with endemic sawfly densities. Parasitoids were less successful in stands with endemic sawfly densities. Total mortality depended on stand character and the proportion of pine around experimental trees.The explained variation in the response variables by the models is relatively low, indicating that other measures of heterogeneity, like understory vegetation and presence of dead wood could contribute to the observed variation. Also, interference between generalist and specialist enemies could affect the realized mortality pressure. Thus, the effect of tree species diversity in combination with these other measures of heterogeneity needs to be recognized to promote the presence and the activity of natural enemies in managed habitats.     

A continent‐scale test of multiple hypotheses on the abundances of Neotropical birds

Explaining variation in the abundance of species remains a challenge in ecology. We sought to explain variation in abundance of Neotropical forest birds using a dataset of population densities of 596 species. We tested a priori hypotheses for the roles of species traits, environmental factors, and species interactions. Specifically, we focused on four factors: 1) body mass (trait), 2) habitat type (environmental factor), 3) net primary productivity (NPP; environmental factor) and 4) species richness of competitors (species interaction). Body size explained much variation in density, although only when analyzed at higher taxonomic levels. Habitat type was a strong predictor of density. The relationship between density and productivity was weak. Densities were related negatively to the species richness of heterospecifics. This trend was particularly strong within closely related groups. Our results show that the influence of energetic factors such as body size and productivity depends on phylogeny, and that they act through indirect relations with other variables; alternative ecological factors such as habitat structure and species interactions play a more direct and stronger role in determining abundance than previously thought.     

The richness–heterogeneity relationship differs between heterogeneity measures within and among habitats

The positive monotonic relationship between habitat heterogeneity and species richness is a cornerstone of ecology. Recently, it was suggested that this relationship should be unimodal rather than monotonic due to a tradeoff between environmental heterogeneity and population sizes, which increases local species extinctions at high heterogeneity levels. Here, we studied the richness–heterogeneity relationship for an avian community using two different environmental variables, foliage‐height diversity and cover type diversity. We analyzed the richness–heterogeneity within different habitat types (grasslands, savannas, or woodlands) and at the landscape scale. We found strong evidence that both positive and unimodal relationships exist at the landscape scale. Within habitats we found positive relationships between richness and heterogeneity in grasslands and woodlands, and unimodal relationships in savannas. We suggest that the length of the environmental heterogeneity gradient (which is affected by both spatial scale and the environmental variable being analyzed) affects the type of the richness–heterogeneity relationship. We conclude that the type of the relationship between species richness and environmental heterogeneity is non‐ubiquitous, and varies both within and among habitats and environmental variables.     

Relative importance of the land‐use composition and intensity for the bird community composition in anthropogenic landscapes

Humans are changing the biosphere by exerting pressure on land via different land uses with variable intensities. Quantifying the relative importance of the land‐use composition and intensity for communities may provide valuable insights for understanding community dynamics in human‐dominated landscapes. Here, we evaluate the relative importance of the land‐use composition versus land‐use intensity on the bird community structure in the highly human‐dominated region surrounding Paris, France. The land‐use composition was calculated from a land cover map, whereas the land‐use intensity (reverse intensity) was represented by the primary productivity remaining after human appropriation (NPP_remaining), which was estimated using remote sensing imagery. We used variance partitioning to evaluate the relative importance of the land‐use composition versus intensity for explaining bird community species richness, total abundance, trophic levels, and habitat specialization in urban, farmland, and woodland habitats. The land‐use composition and intensity affected specialization and richness more than trophic levels and abundance. The importance of the land‐use intensity was slightly higher than that of the composition for richness, specialization, and trophic levels in farmland and urban areas, while the land‐use composition was a stronger predictor of abundance. The intensity contributed more to the community indices in anthropogenic habitats (farmland and urban areas) than to those in woodlands. Richness, trophic levels, and specialization in woodlands tended to increase with the NPP_remaining value. The heterogeneity of land uses and intensity levels in the landscape consistently promoted species richness but reduced habitat specialization and trophic levels. This study demonstrates the complementarity of NPP_remaining to the land‐use composition for understanding community structure in anthropogenic landscapes. Our results show, for the first time, that the productivity remaining after human appropriation is a determinant driver of animal community patterns, independent of the type of land use.     

Landscape‐ and small‐scale determinants of grassland species diversity: direct and indirect influences

Species richness is influenced both by mechanisms occurring at landscape scales, such as habitat availability, and local‐scale processes, that are related to abiotic conditions and plant–plant interactions. However, it is rarely tested to what extent local species richness can be explained by the combined effect of factors measured at multiple spatial scales. In this study, we quantified the simultaneous influence of historical landscape‐scale factors (past human population density, and past habitat availability – an index combining area and connectivity) and small‐scale environmental conditions (shrub cover, and heterogeneity of light, soil depth, and other soil environmental variables) on plant species richness in dry calcareous grasslands (alvars). By applying structural equation modelling (SEM) we found that both landscape conditions and local environmental factors had significant direct and indirect (i.e. through the modification of another factor), effects on species richness. At the landscape scale, we found a direct positive influence of historical habitat availability on species richness, and indirect positive influence of past human population (via its effects on historical habitat availability). At small scales, we found a positive direct influence of light heterogeneity and shrub cover on species richness. Conversely, we found that small‐scale soil environmental heterogeneity, which was mainly determined by soil depth heterogeneity, had a negative effect on species richness. Our study indicates that patterns of species richness in alvar grasslands are positively influenced by the anthropogenic management regime that maintained the landscape habitat conditions in the past. However, the abandonment of management, leading to shrub invasion and increased competition for light resources also influenced species richness. In contrast to the positive heterogeneity–diversity relationship we found that soil heterogeneity reduced species richness. Environmental heterogeneity, occurring at the plant neighbourhood scale (i.e. centimetres), can increase the isolation among suitable soil patches and thus hinder the normal functioning of populations. The combination of previous knowledge of the system with new ecological theories facilitates disentangling how species richness responds to complex relationships among factors operating at multiple scales.     

Deconstructing responses of dragonfly species richness to area,nutrients, water plant diversity and forestry

Understanding large-scale variation in species richness in relation to area, energy, habitat heterogeneity and anthropogenic disturbance has been a major task in ecology. Ultimately, variation in species richness results from variation in individual species occupancies. We studied whether the individual species occupancy patterns are determined by the same candidate factors as total species richness. We sampled 26 boreal forest ponds for dragonflies (Odonata) and studied the effects of shoreline length, water vascular plant species density (WVPSD), availability of nutrients, intensity of forestry, amount of Sphagnum peat cover and pH on dragonfly species richness and individual dragonfly species. WVPSD and pH had a strong positive effect on species richness. Removal of six dragonfly species experiencing strongest responses to WVPSD cancelled the relationship between species richness and WVPSD. By contrast, removal of nine least observed species did not affect the relationship between WVPSD and species richness. Thus, our results showed that relatively common species responding strongly to WVPSD shaped the observed species richness pattern whereas the effect of least observed, often rare, species was negligible. Also, our results support the view that, despite of the great impact of energy on species richness at large spatial scales, habitat heterogeneity can still have an effect on species richness in smaller scales, even overriding the effects of area.     

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.     

Contrasting effects of habitat reduction,conversion and alteration on neutral and non neutral biological communities

While habitat transformation driven by human activities is the main driver of current biodiversity changes, there is still no framework to explore and forecast the effects of different types of habitat changes on the richness and composition of biological communities. To tackle this issue, we modeled the dynamics of a regional meta‐community, composed either of ecologically equivalent species (neutral model) or of generalist and specialist species (specialization model), and explored the impact of the overall reduction, patch conversion or alteration of an original habitat into one or several other habitats of different total carrying capacity on the community metrics at equilibrium. Our simulations reveal strong interactions between the community model considered (neutral or specialization model) and the type of habitat change. Under neutrality, the impact of habitat changes on richness can be approached by a power law species–individual relationship (SIR), which may at constant density be simplified into the widely used power law species–area relationship (SAR), independent of the type of change. However, in the presence of generalist and specialist species, we found that 1) while habitat reduction in area also leads to approximately power law SIRs and SARs, 2) patch conversion and alteration have more complex effects on regional species richness, and 3) habitat alteration elicits the functional homogenization of communities, involving a decrease of their average level of specialization.     

Scale‐dependent determinants of plant species richness in a semi‐arid fragmented agro‐ecosystem

Aims: (1) Understanding how the relationship between species richness and its determinants depends on the interaction between scales at which the response and explanatory variables are measured. (2) Quantifying the relative contributions of local, intermediate and large‐scale determinants of species richness in a fragmented agro‐ecosystem. (3) Testing the hypothesis that the relative contribution of these determinants varies with the grain size at which species richness is measured. Location: A fragmented agro‐ecosystem in the Southern Judea Lowland, Israel, within a desert–Mediterranean transition zone. Methods: Plant species richness was estimated using hierarchical nested sampling in 81 plots, positioned in 38 natural vegetation patches within an agricultural matrix (mainly wheat fields) among three land units along a sharp precipitation gradient. Explanatory variables included position along that gradient, patch area, patch isolation, habitat heterogeneity and overall plant density. We used general linear models and hierarchical partitioning of variance to test and quantify the effect of each explanatory variable on species richness at four grain sizes (0.0625, 1, 25 and 225 m²). Results: Species richness was mainly affected by position along a precipitation gradient and overall plant density, and to a lesser extent by habitat heterogeneity. It was also significantly affected by patch area and patch isolation, but only for small grain sizes. The contribution of each explanatory variable to explained variance in species richness varied with grain size, i.e. scale‐dependent. The influence of geographic position and habitat heterogeneity on species richness increased with grain size, while the influence of plant density decreased with grain size. Main conclusions: Species richness is determined by the combined effect of several scale‐dependent determinants. Ability to detect an effect and effect size of each determinant varies with the scale (grain size) at which it is measured. The combination of a multi‐factorial approach and multi‐scale sampling reveals that conclusions drawn from studies that ignore these dimensions are restricted and potentially misleading.     

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.     

Does energy availability influence classical patterns of spatial variation in exotic species richness?

Aim At macroecological scales, exotic species richness is frequently positively correlated with human population density. Such patterns are typically thought to arise because high human densities are associated with increased introduction effort and/or habitat modification and disturbance. Exotic and native species richness are also frequently positively correlated, although the causal mechanisms remain unclear. Energy availability frequently explains much of the variation in species richness and we test whether such species–energy relationships may influence the relationships of exotic species richness with human population density and native species richness. Location Great Britain. Methods We first investigate how spatial variation in the distributions of the 10 exotic bird species is related to energy availability. We then model exotic species richness using native avian species richness, human population density and energy availability as predictors. Species richness is modelled using two sets of models: one assumes independent errors and the other takes spatial correlation into account. Results The probability of each exotic species occurring, in a 10‐km quadrat, increases with energy availability. Exotic species richness is positively correlated with energy availability, human population density and native species richness in univariate tests. When taking energy availability into account, exotic species richness is negligibly influenced by human population density, but remains positively associated with native species richness. Main conclusions We provide one of the few demonstrations that energy availability exerts a strong positive influence on exotic species richness. Within our data, the positive relationship between exotic species richness and human population density probably arises because both variables increase with energy availability, and may be independent of the influence of human density on the probability of establishment. Positive correlations between exotic and native species richness remain when controlling for the influence of energy on species richness. The relevance of such a finding to the debate on the relationship between diversity and invasibility is discussed.     

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.