Direct and indirect effects of area,energy and habitat heterogeneity on breeding bird communities

Aim To compare the ability of island biogeography theory, niche theory and species–energy theory to explain patterns of species richness and density for breeding bird communities across islands with contrasting characteristics. Location Thirty forested islands in two freshwater lakes in the boreal forest zone of northern Sweden (65°55′ N to 66°09′ N; 17°43′ E to 17°55′ E). Methods We performed bird censuses on 30 lake islands that have each previously been well characterized in terms of size, isolation, habitat heterogeneity (plant diversity and forest age), net primary productivity (NPP), and invertebrate prey abundance. To test the relative abilities of island biogeography theory, niche theory and species–energy theory to describe bird community patterns, we used both traditional statistical approaches (linear and multiple regressions) and structural equation modelling (SEM; in which both direct and indirect influences can be quantified). Results Using regression‐based approaches, area and bird abundance were the two most important predictors of bird species richness. However, when the data were analysed by SEM, area was not found to exert a direct effect on bird species richness. Instead, terrestrial prey abundance was the strongest predictor of bird abundance, and bird abundance in combination with NPP was the best predictor of bird species richness. Area was only of indirect importance through its positive effect on terrestrial prey abundance, but habitat heterogeneity and spatial subsidies (emerging aquatic insects) also showed important indirect influences. Thus, our results provided the strongest support for species–energy theory. Main conclusions Our results suggest that, by using statistical approaches that allow for analyses of both direct and indirect influences, a seemingly direct influence of area on species richness can be explained by greater energy availability on larger islands. As such, animal community patterns that seem to be in line with island biogeography theory may be primarily driven by energy availability. Our results also point to the need to consider several aspects of habitat quality (e.g. heterogeneity, NPP, prey availability and spatial subsidies) for successful management of breeding bird diversity at local spatial scales and in fragmented or insular habitats.     

Linking species-area and species-energy relationships in Drosophila microcosms

Resource availability is an important constraint on community structure. Some authors have suggested it conceptually links two of the most basic patterns in ecology, the species–area relationship and the latitudinal gradient in species richness. I present the first experimental test of this conjecture, by manipulating both the area and resource concentration of artificial larval drosophilid fly habitats and then allowing colonization from a natural species pool. Both the abundance and species richness of these habitats depended upon the total quantity of resources available, regardless of whether those resources were contained within smaller high-quality habitats or larger poor-quality habitats. While the intercepts of species–area relationships varied with resource concentration, they all collapsed onto the same species–energy curve. These results support the view that energetic constraints are of fundamental importance in structuring ecological communities, and that such constraints may even help explain ecological patterns such as the species–area relationship that do not explicitly address resource availability.     

Disentangling the effects of area, energy and habitat heterogeneity on boreal forest bird species richness in protected areas

Aim  One of the few general laws in ecology is that species richness is a positive function of area. However, it has been proposed that area would merely be a proxy for energy. Additionally, habitat heterogeneity has been found to be an important factor determining species richness. Yet the relative importance of those relationships is little known, and it is still unclear how they are brought about. We aimed to dissect which factors drive the species richness of boreal forest birds, and to identify the most probable mechanisms.
Location  Forested protected areas in Finland.
Methods  Using bird line census data collected in 104 protected areas, we ran simultaneous autoregressive models to explain the species richness of forest birds. We explored the value of forest area, tree volume, tree growth, mean degree days and habitat heterogeneity as explanatory variables and used the species richness within different species groups, based on the predictions of hypothesized mechanisms, as a response variable.
Results  Energy, rather than area or habitat heterogeneity, seems to be the main driver of species richness in boreal forest birds. More specifically, productive energy was a better predictor of total species richness than solar energy. Among the tested hypothetical mechanisms, the sampling hypothesis received strong support. After accounting for sampling, solar energy had an effect on species richness.
Main conclusions  As productive energy, such as tree volume, is associated with species richness, high-energy areas should be prioritized in forest conservation planning. Reductions in productive energy may first lead to the disappearance of the rarest species due to the random sampling process. Climate change may result in increased species richness due to increasing amount of productive and solar energy in forests. However, the range shifts of bird species may not be fast enough to keep up with the temperature increases.     

Wasp community responses to habitat complexity in Sydney sandstone forests

Abstract Habitat structure and complexity affect the diversity and composition of fauna in a number of systems. We investigated patterns in wasp species richness, abundance and composition and also their associations with habitat complexity in Sydney sandstone forests, Australia. Pitfall and flight‐intercept traps collected dissimilar wasp assemblages. High complexity habitats supported greater abundance and species richness and a dissimilar composition of pitfall‐trapped wasps to low complexity habitats. Soil moisture, tree canopy cover, ground herb cover and shrub canopy cover all had significant positive associations with the species richness of pitfall‐trapped wasps. Although the five most abundant families of wasps we trapped are endoparasitoids of other arthropods, they showed a variety of preferences for habitat variables. The mechanisms driving associations between habitat complexity and patterns in wasp communities may also provide a basis for understanding factors influencing the regulation of arthropod assemblages by wasps in agricultural and natural landscapes.     

Global patterns of diversity among the specialist birds of riverine landscapes

1. Despite the growing view that biodiversity provides a unifying theme in river ecology, global perspectives on richness in riverine landscapes are limited. As a result, there is little theory or quantitative data on features that might have influenced global patterns in riverine richness, nor are there clear indications of which riverine landscapes are important to conservation at the global scale. As conspicuous elements of the vertebrate fauna of riverine landscapes, we mapped the global distributions of all of the world's specialist riverine birds and assessed their richness in relation to latitude, altitude, primary productivity and geomorphological complexity (surface configuration). 2. Specialist riverine birds, typical of high‐energy riverine landscapes and dependent wholly or partly on production from river ecosystems, occur in 16 families. They are represented by an estimated 60 species divided equally between the passerines and non‐passerines. Major radiation has occurred among different families on different continents, indicating that birds have evolved several times into the niches provided by riverine landscapes. 3. Continental richness varies from four species in Europe to 28 in Asia, with richness on the latter continent disproportionately larger than would be expected from a random distribution with respect to land area. Richness is greatest in mountainous regions at latitudes of 20–40°N in the riverine landscapes of the Himalayan mountains, where 13 species overlap in range. 4. Family, genus and species richness in specialist riverine birds all increase significantly with productivity and surface configuration (i.e. relief). However, family richness was the best single predictor of the numbers of species or genera. In keeping with the effect of surface configuration, river‐bird richness peaks globally at 1300–1400 m altitude, and most species occur typically on small, fast rivers where they feed predominantly on invertebrates. Increased lengths of such streams in areas of high relief and rainfall might have been responsible for species–area effects. 5. We propose the hypothesis that the diversity in channel forms and habitats in riverine landscapes, in addition to high temperature and primary productivity, have been prerequisites to the development of global patterns in the richness of specialist riverine organisms. We advocate tests of this hypothesis in other taxonomic groups. We draw attention, however, to the challenges of categorically defining riverine organisms in such tests because (i) rivers grade into many other habitat types across several different ecotones and (ii) `terrestrialisation' processes in riverine landscapes means that they offer habitat for organisms whose evolutionary origins are not exclusively riverine.     

Disentangling vegetation diversity from climate–energy and habitat heterogeneity for explaining animal geographic patterns

Broad‐scale animal diversity patterns have been traditionally explained by hypotheses focused on climate–energy and habitat heterogeneity, without considering the direct influence of vegetation structure and composition. However, integrating these factors when considering plant–animal correlates still poses a major challenge because plant communities are controlled by abiotic factors that may, at the same time, influence animal distributions. By testing whether the number and variation of plant community types in Europe explain country‐level diversity in six animal groups, we propose a conceptual framework in which vegetation diversity represents a bridge between abiotic factors and animal diversity. We show that vegetation diversity explains variation in animal richness not accounted for by altitudinal range or potential evapotranspiration, being the best predictor for butterflies, beetles, and amphibians. Moreover, the dissimilarity of plant community types explains the highest proportion of variation in animal assemblages across the studied regions, an effect that outperforms the effect of climate and their shared contribution with pure spatial variation. Our results at the country level suggest that vegetation diversity, as estimated from broad‐scale classifications of plant communities, may contribute to our understanding of animal richness and may be disentangled, at least to a degree, from climate–energy and abiotic habitat heterogeneity.     

Alien species richness is currently unbounded in all but the most urbanized bird communities

Urban areas suffer high pressure of introductions of alien species compared to other habitats due to intensive human activities. As trading globally continues to rise, more species will likely be introduced into urban areas. To determine whether this increase in introduction pressure will lead to increased alien species richness in urban areas, or whether other processes would act to impose an upper limit on species richness, we examined how the shape of the relationship between alien species richness and the number of introduced species over time (i.e. introduction pressure) varies along gradients of urbanization. We collected species composition data from urban bird surveys worldwide and used a global database of alien bird introductions to quantify how many species have been introduced over time at different sites. We found that urbanization gradually modified the shape of the studied relationship from linear to asymptotic. Only communities in extremely urbanized environments were associated with an asymptotic relationship, suggesting that alien bird richness has likely not reached its ecological limit in most urban areas. Our results show that urbanization can reduce the importance of introduction pressure in determining alien species richness. Additionally, the results predict that alien species richness will increase at finer spatial scales, especially if the introduced species can survive in urban areas outside of their native range.     

Intraspecific abundance–occupancy relationships: case studies of six bird species in Britain

Although acknowledged to be common, intraspecific relationships between local abundance and site occupancy have been examined in detail for few species. Here we report such analyses for six widespread species of breeding birds in Britain, using data from the Common Birds Census. These exhibit a range of temporal trends, including different combinations of increase and decrease in abundance and occupancy. Overall, two species have a statistically significant positive abundance–occupancy relationship on farmland but no relationship in woodland (collared dove, tree sparrow), one a significant positive relationship on farmland and in woodland (magpie), two a significant positive relationship on farmland and a negative one in woodland (redstart, song thrush), and one a significant negative abundance–occupancy relationship on farmland but no relationship in woodland (sparrowhawk). The population dynamics associated with these patterns are used to discern their underlying mechanisms.     

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.     

Species–energy relationship in the deep sea: a test using the Quaternary fossil record

Little is known about the processes regulating species richness in deep‐sea communities. Here we take advantage of natural experiments involving climate change to test whether predictions of the species–energy hypothesis hold in the deep sea. In addition, we test for the relationship between temperature and species richness predicted by a recent model based on biochemical kinetics of metabolism. Using the deep‐sea fossil record of benthic foraminifera and statistical meta‐analyses of temperature‐richness and productivity‐richness relationships in 10 deep‐sea cores, we show that temperature but not productivity is a significant predictor of species richness over the past c. 130 000 years. Our results not only show that the temperature‐richness relationship in the deep‐sea is remarkably similar to that found in terrestrial and shallow marine habitats, but also that species richness tracks temperature change over geological time, at least on scales of c. 100 000 years. Thus, predicting biotic response to global climate change in the deep sea would require better understanding of how temperature regulates the occurrences and geographical ranges of species.     

Noncrop features and heterogeneity mediate overwintering bird diversity in agricultural landscapes of southwest China

Farmland birds are of conservation concerns around the world. In China, conservation management has focused primarily on natural habitats, whereas little attention has been given to agricultural landscapes. Although agricultural land use is intensive in China, environmental heterogeneity can be highly variable in some regions due to variations in crop and noncrop elements within a landscape. We examined how noncrop heterogeneity, crop heterogeneity, and noncrop features (noncrop vegetation and water body such as open water) influenced species richness and abundance of all birds as well as three functional groups (woodland species, agricultural land species, and agricultural wetland species) in the paddy‐dominated landscapes of Erhai water basin situated in northwest Yunnan, China. Birds, crop, and noncrop vegetation surveys in twenty 1 km × 1 km landscape plots were conducted during the winter season (from 2014 to 2015). The results revealed that bird community compositions were best explained by amounts of noncrop vegetation and compositional heterogeneity of noncrop habitat (Shannon–Wiener index). Both variables also had a positive effect on richness and abundance of woodland species. Richness of agricultural wetland species increased with increasing areas of water bodies within the landscape plot. Richness of total species was also greater in the landscapes characterized by larger areas of water bodies, high proportion of noncrop vegetation, high compositional heterogeneity of noncrop habitat, or small field patches (high crop configurational heterogeneity). Crop compositional heterogeneity did not show significant effects neither on the whole community (all birds) nor on any of the three functional groups considered. These findings suggest that total bird diversity and some functional groups, especially woodland species, would benefit from increases in the proportion of noncrop features such as woody vegetation and water bodies as well as compositional heterogeneity of noncrop features within landscape.     

Resources and global avian assemblage structure in forests

Explaining spatial variation in a number of bird species, particularly from temperate to tropical regions, has been a longstanding challenge. We test at a global scale whether species‐rich forest assemblages are associated with division of a larger resource pool, a finer division of that pool, or some combination of the two. Species richness increases with increasing assemblage abundance, biomass and energy use. As assemblage abundance, biomass and energy use increase with increasing energy availability, and as per species numbers of individuals, biomass and energy use do not decrease with increasing energy availability, we provide direct evidence that the avian species–energy relationship in forests is associated foremost with an increase in the size of the resource pool and not with a finer level of its subdivision.     

Productivity gradients cause positive diversity–invasibility relationships in microbial communities

Models predict that community invasibility generally declines with species diversity, a prediction confirmed by small‐scale experiments. Large‐scale observations and experiments, however, find that diverse communities tend to be more heavily invaded than simple communities. One hypothesis states that large‐scale environmental heterogeneity, which similarly influences native and invasive species, can cause a positive correlation between diversity and invasibility, overriding the local negative effects of diversity on invasibility. We tested this hypothesis using aquatic microbial communities consisting of protists and rotifers consuming bacteria and nanoflagellates. We constructed a productivity gradient to simulate large‐scale environmental heterogeneity, started communities with the same number of species along this gradient, and subjected equilibrial communities to invasion by non‐resident consumer species. Both invaders and most resident species increased their abundances with resource enrichment, resulting in a positive correlation between diversity and invasibility. Intraspecific interference competition within resident species and the positive effect of enrichment on the number of available resources probably accounted for the higher invasibility with enrichment. Our results provide direct experimental evidence that environmental heterogeneity in productivity can cause a positive diversity–invasibility relationship.     

Energy availability, abundance, energy-use and species richness in forest bird communities: a test of the species–energy theory

Aim To test the ‘more individuals hypothesis’ as a mechanism for the positive association between energy availability and species richness. This hypothesis predicts that total density and energy use in communities is linearly related to energy availability, and that species richness is a positive function of increased density. We also evaluate whether similar energy–density patterns apply to different migratory groups (residents, short‐distance migrants and tropical migrants) separately. Location European and North American forest bird communities. Methods We collected published breeding bird census data from Europe and North America (n = 187). From each census data we calculated bird density (pairs 10 ha^?1), energy use by the community (the sum of metabolic needs of individuals, Watts 10 ha^?1) and geographical location with an accuracy of 0.5°. For each bird census data coordinate we extracted the corresponding monthly values of actual evapotranspiration (AET). From these values we calculated corresponding AET values that we expected to explain the density energy use of forest birds: total annual, breeding season (June) and winter AET. We used general linear modelling to analyse these data controlling for the area of census plots, forest type and census method. Results Total density and energy use in European and North American forest bird communities were linear functions of annual productivity, and increased density and energy use then translated into more species. Also resident bird density and energy consumption were positive functions of annual productivity, but the relationship between productivity and density as well as between productivity and energy use was weaker for migrants. Main conclusions Our results are consistent with the more individuals hypothesis that density and energy use in breeding forest bird communities is coupled tightly with the productivity of the environment, and that increased density and energy consumption results in more species. However, not all community members (migratory groups) are limited by productivity on the breeding grounds.     

Macro‐scale bird species richness patterns of the East Asian mainland and islands: energy,area and isolation

Aim To create a map of bird species richness (BSR) in East Asia and to examine the effect of area, isolation, primary productivity, topographic heterogeneity, and human population density on BSR. Location East Asia (from 70° E to 180° E longitude), including the eastern half of the Palaearctic Region, the entire Oriental Region, and the entire Wallacea Subregion. Methods The breeding ranges of 2406 terrestrial bird species were mapped and overlaid to create a species richness map. The BSR map was transformed into a 100 × 100 km quadrat system, and BSR was analysed in relation to land area, average normalized difference vegetation index (NDVI), elevation range, and average population density. Results In general, BSR declined from the Tropics to the Arctic. In mainland East Asia, however, BSR was highest around the Tropic of Cancer, and fluctuated between 30° and 50° N. Islands had lower BSR than adjacent mainland areas. The NDVI was strongly positively correlated with BSR in mainland areas and on islands. For mainland areas, NDVI explained 65% of the BSR variation, and topographic heterogeneity explained an additional 6% in ordinary least‐squares regression. On islands, NDVI explained 66% of BSR variation, island area explained 13%, and distance to mainland accounted for 1%. Main conclusions In East Asia, we suggest that primary productivity is the key factor underpinning patterns of BSR. Primary productivity sets the upper limits of the capacity of habitats to support bird species. In isolated areas such as islands and peninsulas, however, BSR might not reach the richness limits set by primary productivity because the degree of isolation and area size also can affect species richness. Other factors, such as spatial heterogeneity, biotic interactions, and perturbations, may also affect species richness. However, their effects are secondary and are not as strong as primary productivity, isolation, and area size.     

The terminology and use of species–area relationships: a response to Dengler (2009)

Dengler ( Journal of Biogeography , 2009, 36 , 728–744) addresses issues regarding species–area relationships (SARs), but fails to settles those issues. He states that only certain types of sampling schemes should be used to construct SARs, but is not consistent in the criteria that he uses to include some sampling schemes but not others. He argues that a sampling scheme of contiguous plots will be more accurate in extrapolating beyond the sampled area, but logic tells us that a dispersed sampling scheme is likely to be more accurate. Finally, he concludes that the 'true' SAR is a power function, but this conclusion is inconsistent with his results and with the results of others. Rather than defining a narrow framework for SARs, we need to recognize that the relationship between area and species richness is scale- and system-dependent. Different sampling schemes serve different purposes, and a variety of functional relationships are likely to hold. Further theoretical and empirical work is needed to resolve these issues fully.     

Bird diversity patterns in Neotropical temperate farmlands: The role of environmental factors and trophic groups in the spring and autumn

Productivity, habitat heterogeneity and environmental similarity are of the most widely accepted hypotheses to explain spatial patterns of species richness and species composition similarity. Environmental factors may exhibit seasonal changes affecting species distributions. We explored possible changes in spatial patterns of bird species richness and species composition similarity. Feeding habits are likely to have a major influence in bird–environment associations and, given that food availability shows seasonal changes in temperate climates, we expect those associations to differ by trophic group (insectivores or granivores). We surveyed birds and estimated environmental variables along line‐transects covering an E‐W gradient of annual precipitation in the Pampas of Argentina during the autumn and the spring. We examined responses of bird species richness to spatial changes in habitat productivity and heterogeneity using regression analyses, and explored potential differences between seasons of those responses. Furthermore, we used Mantel tests to examine the relationship between species composition similarity and both the environmental similarity between sites and the geographic distance between sites, also assessing differences between seasons in those relationships. Richness of insectivorous birds was directly related to primary productivity in both seasons, whereas richness of seed‐eaters showed a positive association with habitat heterogeneity during the spring. Species composition similarity between assemblages was correlated with both productivity similarity and geographic proximity during the autumn and the spring, except for insectivore assemblages. Diversity within main trophic groups seemed to reflect differences in their spatial patterns as a response to changes between seasons in the spatial patterns of food resources. Our findings suggest that considering different seasons and functional groups in the analyses of diversity spatial pattern could contribute to better understand the determinants of biological diversity in temperate climates.