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.     

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.     

Environmental heterogeneity as a universal driver of species richness across taxa,biomes and spatial scales

Environmental heterogeneity is regarded as one of the most important factors governing species richness gradients. An increase in available niche space, provision of refuges and opportunities for isolation and divergent adaptation are thought to enhance species coexistence, persistence and diversification. However, the extent and generality of positive heterogeneity–richness relationships are still debated. Apart from widespread evidence supporting positive relationships, negative and hump‐shaped relationships have also been reported. In a meta‐analysis of 1148 data points from 192 studies worldwide, we examine the strength and direction of the relationship between spatial environmental heterogeneity and species richness of terrestrial plants and animals. We find that separate effects of heterogeneity in land cover, vegetation, climate, soil and topography are significantly positive, with vegetation and topographic heterogeneity showing particularly strong associations with species richness. The use of equal‐area study units, spatial grain and spatial extent emerge as key factors influencing the strength of heterogeneity–richness relationships, highlighting the pervasive influence of spatial scale in heterogeneity–richness studies. We provide the first quantitative support for the generality of positive heterogeneity–richness relationships across heterogeneity components, habitat types, taxa and spatial scales from landscape to global extents, and identify specific needs for future comparative heterogeneity–richness research.     

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.     

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.     

Drivers of moth species richness on tropical altitudinal gradients: a cross-regional comparison

Aims We examine the role of species–area relationships (SARs), climatic parameters and phylogeny in shaping the altitudinal species richness patterns of moths. With respect to SARs, we investigate whether habitat heterogeneity is a probable mechanism for mediating area effects. We investigate the consistency of patterns by comparing several discrete regions. Location Nine mountainous regions in tropical Asia and the Malay Archipelago. Methods Presence‐only records for 292 species of the Lepidopteran family Sphingidae were used to measure interpolated species richness in 200‐m altitudinal bands. Species richness was correlated with area measures, which were calculated from both two‐dimensional map projections and three‐dimensional digital elevation models (DEMs). We used data simulations of homogeneous communities to test for effects of sample (i.e. habitat) heterogeneity as a mechanism causing SARs. Species richness patterns were compared among regions and between the two major sphingid clades, and were related to regional climatic characteristics. Results The area of altitudinal bands was a strong (statistical) explanation of species richness, particularly if area was calculated from three‐dimensional DEMs, but SARs often over‐predict species richness in lowland areas. There was no evidence for habitat heterogeneity as a mechanism of altitudinal SARs (tested for Borneo only). Species richness patterns varied considerably between the nine regions, which may, as an alternative to SARs, be explained by climatic differences such as (temperature) seasonality. Phylogenetic clades differed in species richness patterns exhibited. Main conclusion SARs provide strong empirical explanations for (regional) altitudinal patterns of species richness, but lack of evidence for the most likely mechanism cautions against a priori ‘corrections’ of species richness data for area. Furthermore, SARs are often not a sufficient explanation for the drop in species richness towards lowlands. Climate, or other collinear variables, may offer alternative explanations for altitudinal SARs. More research is needed to understand the mechanisms for SARs in an altitudinal context in order to evaluate their importance in the face of parameter collinearity.     

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.     

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.     

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.     

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.     

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.     

Species richness, environmental correlates, and spatial scale: a test using South African birds

Energy and habitat heterogeneity are important correlates of spatial variation in species richness, though few investigations have sought to determine simultaneously their relative influences. Here we use the South African avifauna to examine the extent to which species richness is related to these variables and how these relationships depend on spatial grain. Taking spatial autocorrelation and area effects into account, we find that primary productivity, precipitation, absolute minimum temperature, and, at coarser resolutions, habitat heterogeneity account for most of the variation in species richness. Species richness and productivity are positively related, whereas the relationship between potential evapotranspiration (PET) and richness is unimodal. This is largely because of the constraining effects of low rainfall on productivity in high-PET areas. The increase in the importance of vegetation heterogeneity as an explanatory variable is caused largely by an increase in the range of vegetation heterogeneity included at coarse resolutions and is probably also a result of the positive effects of environmental heterogeneity on species richness. Our findings indicate that species richness is correlated with, and hence likely a function of, several variables, that spatial resolution and extent must be taken into account during investigations of these relationships, and that surrogate measures for productivity should be interpreted cautiously.     

Island biogeography of bats in Baja California, Mexico: patterns of bat species richness in a near-shore archipelago

Aim We studied the relationship between the size and isolation of islands and bat species richness in a near‐shore archipelago to determine whether communities of vagile mammals conform to predictions of island biogeography theory. We compared patterns of species richness in two subarchipelagos to determine whether area per se or differences in habitat diversity explain variations in bat species richness. Location Islands in the Gulf of California and adjacent coastal habitats on the Baja California peninsula in northwest Mexico. Methods Presence–absence surveys for bats were conducted on 32 islands in the Gulf of California using acoustic and mist‐net surveys. We sampled for bats in coastal habitats of four regions of the Baja peninsula to characterize the source pool of potential colonizing species. We fitted a semi‐log model of species richness and multiple linear regression and used Akaike information criterion model selection to assess the possible influence of log₁₀ area, isolation, and island group (two subarchipelagos) on the species richness of bats. We compared the species richness of bats on islands with greater vegetation densities in the southern gulf (n = 20) with that on drier islands with less vegetation in the northern gulf (n = 12) to investigate the relationship between habitat diversity and the species richness of bats. Results Twelve species of bats were detected on islands in the Gulf of California, and 15 species were detected in coastal habitats on the Baja peninsula. Bat species richness was related to both area and isolation of islands, and was higher in the southern subarchipelago, which has denser vegetation. Log₁₀ area was positively related to bat species richness, which increased by one species for every 5.4‐fold increase in island area. On average, richness declined by one species per 6.25 km increase in isolation from the Baja peninsula. Main conclusions Our results demonstrate that patterns of bat species richness in a near‐shore archipelago are consistent with patterns predicted by the equilibrium theory of island biogeography. Despite their vagility, bats may be more sensitive to moderate levels of isolation than previously expected in near‐shore archipelagos. Differences in vegetation and habitat xericity appear to be associated with richness of bat communities in this desert ecosystem. Although observed patterns of species richness were consistent with those predicted by the equilibrium theory, similar relationships between species richness and size and isolation of islands may arise from patch‐use decision making by individuals (optimal foraging strategies).     

Do plant communities exist? Evidence from scaling‐up local species‐area relations to the regional level

Abstract. One long tradition in ecology is that discrete communities exist, at least in the sense that there are areas of relatively uniform vegetation, with more rapid change in species composition between them. The alternative extreme view is the Self‐similarity concept – that similar community variation occurs at all spatial scales. We test between these two by calculating species‐area curves within areas of vegetation that are as uniform as can be found, and then extrapolating the within‐community variation to much larger areas, that will contain many ‘communities’. Using the Arrhenius species‐area model, the extrapolations are remarkably close to the observed number of species at the regional/country level. We conclude that the type of heterogeneity that occurs within ‘homogeneous’ communities is sufficient to explain species richness at much larger scales. Therefore, whilst we can speak of ‘communities’ for convenience, the variation that certainly exists at the ‘community’ level can be seen as only a larger‐scale manifestation of micro‐habitat variation.     

The influence of habitat heterogeneity and latitude on gamma diversity of the Nearctic Simuliidae,a ubiquitous group of stream‐dwelling insects

Among the most prominent, large‐scale patterns of species richness are the increases in richness with decreasing latitude and with increasing habitat heterogeneity. Using the stream‐dwelling larval and pupal stages of North American black flies (Diptera: Simuliidae), we address 3 broad questions about species richness: (i) Does a significant latitude–richness relationship exist? (ii) How does habitat heterogeneity influence gamma diversity? (iii) What is the sign (positive or negative) of the latitude–richness and the heterogeneity–richness relationships? We found no evidence that habitat heterogeneity influences gamma diversity. The estimated peak species richness for black flies in North America was at 50–53°N, which also corresponds with peak generic richness. All plesiomorphic, extant lineages of the Simuliidae in the Western Hemisphere are found in cool mountainous environments of North America, suggesting that peak richness at 50–53°N might be a signature of this phylogenetic pattern and a reflection of underlying historical processes.     

Vascular plant species richness on wetland remnants is determined by both area and habitat heterogeneity

There is an ongoing ecological debate on whether area per se or habitat heterogeneity is the main driver for species richness. The wetland remnants in the Sanjiang Plain, NE China harbor a high biodiversity and play an important role for local ecosystems. Fifty-one wetland remnants were sampled to examine the effect of area and habitat heterogeneity on vascular plant species richness. Number of community types, elevation, water heterogeneity and soil resource heterogeneity were employed as habitat heterogeneity variables, but only water heterogeneity was identified as the proper surrogate for habitat heterogeneity. Compared with the classic species-area model, the choros model achieved better fitness when water heterogeneity and elevation were employed as habitat heterogeneity variables. Nevertheless, elevation was poorly correlated with species richness. It suggests, without a comprehensive analysis of habitat heterogeneity variables, the choros model might result in a misleading result. In this study, species richness was significantly influenced by water heterogeneity, area and number of community types. Water heterogeneity and area both controlled the number of community types, and they were the two main determinants of species richness. As area was significantly and positively correlated with water heterogeneity, the variance in species richness was mainly related to the mutual effect of area and water heterogeneity. The results of this study confirmed that the relationship between the area per se hypothesis and the habitat heterogeneity hypothesis was conjunct rather than mutually exclusive. In addition, it is critical that both area and water heterogeneity should be taken into account for biodiversity conservation and management in wetland remnants.     

An exploration of factors influencing lotic insect species richness

An understanding of factors that influence species richness of lotic insects is generally lacking. We present comparative data on aquatic insect species richness from several North American and other streams. Factors such as large sample numbers and drainage area (species area relationships) are not significant predictors of species richness across the streams we examined. We explore several hypotheses regarding the origins and maintenance of species richness using Upper Three Runs Creek (UTR), South Carolina, USA, as a reference stream. UTR has the highest species richness of any stream in the Western Hemisphere. Hypotheses examined included historical, regional and local processes such as: (1) Evolutionary time, (2) disturbance regime/environmental variability, (3) temperature/evolutionary-speed, (4) productivity, and (5) habitat heterogeneity. Of these hypotheses, we suggest that productivity and habitat heterogeneity appear to contribute most to the high species richness found in UTR. We believe that multidisciplinary analysis of other streams is necessary because without this crucial information our knowledge of, and desire to protect biodiversity in streams will be wanting.     

Patterns of invertebrate density and taxonomic richness across gradients of area,isolation, and vegetation diversity in a lake‐island system

Over the past half century, ecologists have tried to unravel the factors that drive species richness patterns in ecological communities. One influential theory is island biogeography theory (IBT), which predicts that island or habitat area and isolation are drivers of species richness. However, relatively few studies testing IBT have considered invertebrate or belowground communities, and it is unclear as to whether the predictions made by IBT hold for these communities. Other theories predict that habitat characteristics such as vegetation diversity may be important drivers of invertebrate species richness. To investigate patterns of invertebrate density and species richness across gradients of area, isolation, and vegetation diversity, we used a system of 30 lake islands in the boreal zone of northern Sweden. We assessed density and taxonomic richness of ground‐dwelling spiders, web‐building spiders, beetles, collembolans, mites, and nematodes, for all islands during two consecutive summers. For all invertebrate groups, both density and taxonomic richness were either neutrally or negatively related to island size, and either neutrally or positively related to island isolation. Meanwhile the density and taxonomic richness for several groups was positively related to vegetation diversity (i.e. habitat heterogeneity). In multiple regression analyses, island size was often the single best predictor for both invertebrate density and taxonomic richness, but in some cases island size and isolation in combination explained more variation than each factor considered singly. Contrary to IBT predictions, invertebrate density and richness was never positively related to island size or negatively related to island isolation. Instead, our results suggest that plant diversity (and thus habitat heterogeneity) was the main driver of the patterns that we found, although other factors could have some influence. We conclude that several factors, but not necessarily those predicted as important by IBT, are important in determining invertebrate abundance and species richness in island systems.