Disentangling the relative effects of ambient energy,water availability,and energy–water balance on pteridophyte species richness at a landscape scale in China

Understanding what factors generate geographic variation in species richness is a fundamental goal of ecology and biogeography. Water and energy are considered as the major environmental factors influencing large-scale patterns of species richness, but their roles vary among taxa and regions. Pteridophytes are an ideal group of organisms for examining the relationship between species richness and their environment because the distribution of pteridophytes is usually in equilibrium with contemporary climate to a greater degree than those of seed plants and most terrestrial vertebrates partly due to the lightness of their spores, which is highly capable of long-distance dispersal by wind, and partly due to their single-spore reproduction strategy. Using correlation and regression analyses and structural equation modeling technique, we examine the relationship of pteridophyte species richness in 151 localities from across China with environmental factors representing energy, water, and energy–water balance. We found that pteridophyte species richness is correlated to water availability more strongly than to ambient energy. Furthermore, we found that of all environmental variables considered, energy–water balance has played the most important role in regulating pteridophyte species richness gradients in China.     

Contrasting environmental and regional effects on global pteridophyte and seed plant diversity

Pteridophytes (ferns and fern‐allies) represent the second‐largest group of vascular plants, but their global biogeography remains poorly studied. Given their functional biology, pteridophytes are expected to show a more pronounced relation to water availability and a higher dispersal ability compared to seed plants. We test these assertions and document the global pattern of pteridophyte richness across 195 mainland and 106 island regions. Using non‐spatial and spatial simple and multiple regression models, we analyze geographic trends in pteridophyte and seed plant richness as well as pteridophyte proportions in relation to environmental and regional variables. We find that pteridophyte and seed plant richness are geographically strongly correlated (all floras: r=0.68, mainland: r=0.82, island floras: r=0.77), but that the proportions of pteridophytes in vascular plant floras vary considerably (0–70%). Islands (mean=15.3%) have significantly higher proportions of pteridophytes than mainland regions (mean=3.6%). While the relative proportions of pteridophytes on islands show a positive relationship with geographic isolation, proportions in mainland floras increase most strongly along gradients of water availability. Pteridophyte richness peaks in humid tropical mountainous regions and is lowest in deserts, arctic regions, and on remote oceanic islands. Regions with Mediterranean climate, outstanding extra‐tropical centres of seed plant richness, are comparatively poor in pteridophytes. Overall, water‐energy variables and topographical complexity are core predictors of both mainland pteridophyte and seed plant richness. Significant residual richness across biogeographic regions points to an important role of idiosyncratic regional effects. Although the same variables emerge as core predictors of pteridophyte and seed plant richness, water availability is clearly a much stronger constraint of pteridophyte richness. We discuss the different limitations of gametophytes and sporophytes that might have limited the ability of pteridophytes to extensively diversify under harsh environmental conditions. Our results point to an important role of taxon‐specific functional traits in defining global richness gradients.     

Spatial scaling and transition in pneumatophore arthropod communities

Although most ecological variables are scale-dependent, few studies cover a broad range of spatial scales. Here, we consider South African mangrove pneumatophore arthropod communities (mites, crustaceans and insects), across seven spatial scales (from 10  cm to 100  km). We plot spatial autocorrelation in individual species, evaluate if resource and habitat availability determine spatial patterning, and identify the scales of community transition. Spatial autocorrelation in most ecological variables decreased with increasing spatial scale, with notable exceptions for the larger scales. Negative abundance autocorrelation was stronger at 10  km than at 100  km for common species, while the opposite was true for rare species. Spatial autocorrelation in species richness decreased from 1  m (strong positive) to 10  km (strong negative), but was not significant at the 100  km scale. These patterns reflect the patchy distribution of pneumatophores within mangrove forests, that of the forests along the coast, and the poor dispersal abilities of most of the arthropods sampled, in a highly dynamic environment. Although resource and habitat availability exhibited a similar autocorrelation pattern to that of the community, the total mass of pneumatophores did not appear to be an important determinant of community structure. Variations in the abundance of common species, as well as the restricted distribution of rare species caused assemblage structure to change gradually with increasing distance from 10 cm to 100 km, but only marginally from 10 to 100  km. We highlight the need for cross-scale studies in bridging the gap between two key ecological concepts: potential ecological niche and realized geographic range.     

What drives elevational patterns of diversity? A test of geometric constraints,climate and species pool effects for pteridophytes on an elevational gradient in Costa Rica

Aim We studied pteridophyte species richness between 100 m and 3400 m along a Neotropical elevational gradient and tested competing hypotheses for patterns of species richness. Location Elevational transects were situated at Volcán Barva in the Braulio Carrillo National Park and La Selva Biological Station (100–2800 m) and Cerro de la Muerte (2700–3400 m), both on the Atlantic slope of Costa Rica, Central America. Method We analysed species richness on 156 plots of 20 × 20 m and measured temperature and humidity at four elevations (40, 650, 1800 and 2800 m). Species richness patterns were regressed against climatic variables (temperature, humidity, precipitation and actual evapotranspiration), regional species pool, area and predicted species number of a geometric null model (the mid‐domain effect, MDE). Results The species richness of the 484 recorded species showed a hump‐shaped pattern with elevation with a richness peak at mid‐elevations (c. 1700 m). The MDE was the single most powerful explanatory variable in linear regression models, but species richness was also associated strongly with climatic variables, especially humidity and temperature. Area and species pool were associated less strongly with observed richness patterns. Main conclusions Geometric models and climatic models exclusive of geometric constraints explained comparable amounts of the elevational variation in species richness. Discrimination between these two factor complexes is not possible based on model fits. While overall fits of geometric models were high, large‐ and small‐ranged species were explained by geometric models to different extents. Species with narrow elevational ranges clustered at both ends of the gradient to a greater extent than predicted by the MDE null models used here. While geometric models explained much of the pattern in species richness, we cannot rule out the role of climatic factors (or vice versa) because the predicted peak in richness from geometric models, the empirical peak in richness and the overlap in favourable environmental conditions all coincide at middle elevations. Mid‐elevations offer highest humidity and moderate temperatures, whereas at high elevations richness is reduced due to low temperatures, and at low elevations by reduced water availability due to high temperatures.     

Spatial scale, abundance and the species-energy relationship in British birds

1. The spatial scale of analysis may influence the nature, strength and underlying drivers of macroecological patterns, one of the most frequently discussed of which is the relationship between species richness and environmental energy availability. 2. It has been suggested that species-energy relationships are hump-shaped at fine spatial grains and consistently positive at larger regional grains. The exact nature of this scale dependency is, however, the subject of much debate as relatively few studies have investigated species-energy relationships for the same assemblage across a range of spatial grains. Here, we contrast species-energy relationships for the British breeding avifauna at spatial grains of 1 km x 1 km, 2 km x 2 km and 10 km x 10 km plots, while maintaining a constant spatial extent. 3. Analyses were principally conducted using data on observed species richness. While survey work may fail to detect some species, observed species richness and that estimated using nonparametric techniques were strongly positively correlated with each other, and thus exhibit very similar spatial patterns. Moreover, the forms of species-energy relationships using observed and estimated species richness were statistically indistinguishable from each other. 4. Positive decelerating species-energy relationships arise at all three spatial grains. There is little evidence that the explanatory power of these relationships varies with spatial scale. However, ratios of regional (large-scale) to local (small-scale) species richness decrease with increasing energy availability, indicating that local richness responds to energy with a steeper gradient than does regional richness. Local assemblages thus sample a greater proportion of regional richness at higher energy levels, suggesting that spatial turnover of species richness is lower in high-energy regions. Similarly, a crude measure of temporal turnover, the ratio of cumulative species richness over a 4-year period to species richness in a single year, is lower in high-energy regions. These negative relationships between turnover and energy appear to be causal as both total and mean occupancy per species increases with energy. 5. While total density in 1 km x 1 km plots correlates positively with energy availability, such relationships are very weak for mean density per species. This suggests that the observed association between total abundance and species richness may not be mediated by population extinction rates, as predicted by the more individuals hypothesis. 6. The sampling mechanism suggests that species-energy relationships arise as high-energy areas support a greater number of individuals, and that random allocation of these individuals to local areas from a regional assemblage will generate species-energy relationships. While randomized local species-energy relationships are linear and positive, predicted richness is consistently greater than that observed. The mismatch between the observed and randomized species-energy relationships probably arises as a consequence of the aggregated nature of species distributions. The sampling mechanism, together with species spatial aggregation driven by limited habitat availability, may thus explain the species-energy relationship observed at this spatial scale.     

Abundance, species richness and energy availability in the North American avifauna

Aim To determine how species richness, abundance, biomass, energy use and mean number of individuals per species scale with environmental energy availability in wintering and breeding avian assemblages, and to contrast assemblages of (i) common and rare species and (ii) breeding residents and migrants. To assess whether such patterns are compatible with the ‘more individuals hypothesis’ (MIH) that high‐energy areas are species‐rich because they support larger populations that are buffered against extinction. Location The North American continent (latitudinal range 23.4 °?48.1 °N; longitudinal range 124.2°?68.7° W). Methods Avian species richness, abundance, biomass and energy use were calculated for 295 Resident Bird Count plots. Environmental energy availability was measured using ambient temperature and the Normalized Difference Vegetation Index (NDVI), a close correlate of plant productivity. Analyses took plot area into account, and were conducted (with and without taking habitat type into account) using general linear models and spatial mixed models. Results Positive species–energy relationships were exhibited by both wintering and breeding assemblages, but were stronger in the former. The structure of winter assemblages responded more strongly to temperature than NDVI, while breeding assemblages tended to respond more strongly to NDVI. Breeding residents responded to annual measures of energy availability while breeding migrants and the winter assemblage responded more strongly to seasonal measures. In the winter assemblage, rare and common species exhibited species–energy relationships of a similar strength, but common breeding species exhibited a much stronger relationship than rare breeding species. In both breeding and wintering assemblages, abundance, biomass and energy use increased with energy availability and species richness. Energy availability was a poor predictor of the mean number of individuals per species. Main conclusions The nature of the species–energy relationship varies seasonally and with the manner in which energy availability is measured. Our data suggest that residents are less able to respond to seasonal fluxes in resource availability than long‐distance migrants. Increasing species richness and energy availability is associated with increasing numbers of individuals, biomass and energy use. While these observations are compatible with the MIH our data provide only equivocal support for this hypothesis, as the rarest species do not exhibit the strongest species–energy relationships.     

Effects of Forest Fragmentation on Pteridophyte Diversity in a Tropical Rain Forest in Brazil

The impacts of forest fragmentation on the pteridophyte communities of the Una region of Bahia, Brazil, were investigated by comparing species richness and ensemble diversity among areas of large forest fragments (>900 ha), small forest fragments (<100 ha), and landscape matrix. We inventoried the pteridophytes below 1 m in height in interiors of small fragments, interiors of large fragments (control areas), edges of fragments, edges of continuous forest, capoeiras (initial stages of forest regeneration) and cabrucas (cocoa plantations). All ferns were collected following the plot method (plots of 120×10 m, each). Sampling units were established in the six main ecotypes of the Una region. These units were allocated within three sampling blocks of 5 per 5 km, which were chosen in order to include the largest forest patches that still remain. Results suggest that fragmentation has a negative impact on species richness at the matrix and the edges of forest remnants. A similar negative matrix end edge effect is reported for diversity of those sites measured by the α Log-series Index. However, small forest fragments have pteridophyte species richness and diversity rates similar to large ones so they should be considered of utmost importance to the conservation of forest-related species in the region.     

Patterns of distribution for southern Australian marine echinoderms and decapods

Aim To relate patterns of distribution of marine echinoderms and decapods around southern Australia to major ecological and historical factors. Location Shallow‐water (0–100 m) marine waters off southern Australia, south of 30° S. Methods (1) Record the presence/absence of known echinoderm and decapod species in cells of c. 1° latitude and longitude, along the coast of southern mainland Australia and Tasmania. (2) Describe patterns in species composition, species richness and endemism through gradient analysis, ordination and cluster analysis. (3) Relate these patterns to distance and temperature gradients, the area of continental shelf, the average size of species range, and known historical factors. Results Species composition varied with both latitude and longitude. Species richness was relatively constant from east to west but graded with latitude from high in the warm‐temperate regions around Perth and Sydney to low in cool‐temperate southern Tasmania. Species richness was not related to the area of continental shelf or average species range size. Species turnover was not correlated with rates of temperature change. It was problematic to separate distance from temperature gradients, but there was evidence that the southern distribution limits of some species are related to minimum sea surface temperature. Within the taxonomic groups surveyed, evolutionary radiation has been largely limited to a few cosmopolitan species‐rich genera. Main conclusions There are historical as well as ecological hypotheses explaining the latitudinal gradient of marine species richness in southern Australia: (1) the continual invasion and speciation of species of tropical origin as Australia has split from Gondwana and drifted northward; (2) progressive extinction of some Gondwanan cool‐temperate species at the limits of their range; (3) low level of immigration of additional cool‐temperate species; and (4) some in situ endemic speciation.     

Latitudinal patterns of range size and species richness of New World woody plants

Aim  Relationships between range size and species richness are contentious, yet they are key to testing the various hypotheses that attempt to explain latitudinal diversity gradients. Our goal is to utilize the largest data set yet compiled for New World woody plant biogeography to describe and assess these relationships between species richness and range size.
Location  North and South America.
Methods  We estimated the latitudinal extent of 12,980 species of woody plants (trees, shrubs, lianas). From these estimates we quantified latitudinal patterns of species richness and range size. We compared our observations with expectations derived from two null models.
Results   Peak richness and the smallest- and largest-ranged species are generally found close to the equator. In contrast to prominent diversity hypotheses: (1) mean latitudinal extent of tropical species is greater than expected; (2) latitudinal extent appears to be decoupled from species richness across New World latitudes, with abrupt transitions across subtropical latitudes; and (3) mean latitudinal extents show equatorial and north temperate peaks and subtropical minima. Our results suggest that patterns of range size and richness appear to be influenced by three broadly overlapping biotic domains (biotic provinces) for New World woody plants.
Main conclusions  Hypotheses that assume a direct relationship between range size and species richness may explain richness patterns within these domains, but cannot explain gradients in richness across the New World.     

Complex effects of scale on the relationships of landscape pattern versus avian species richness and community structure in a woodland savanna mosaic

Landscape pattern metrics are widely used for predicting habitat and species diversity. However, the relationship between landscape pattern and species diversity is typically measured at a single spatial scale, even though both landscape pattern, and species occurrence and community composition are scale‐dependent. While the effects of scale on landscape pattern are well documented, the effects of scale on the relationships between spatial pattern and species richness and composition are not well known. Here, our main goal was to quantify the effects of cartographic scale (spatial resolution and extent) on the relationships between spatial pattern and avian richness and community structure in a mosaic of grassland, woodland, and savanna in central Wisconsin. Our secondary goal was to evaluate the effectiveness of a newly developed tool for spatial pattern analysis, multiscale contextual spatial pattern analysis (MCSPA), compared to existing landscape metrics. Landscape metrics and avian species richness had quadratic, exponential, or logarithmic relationships, and these patterns were generally consistent across two spatial resolutions and six spatial extents. However, the magnitude of the relationships was affected by both resolution and extent. At the finer resolution (10‐m), edge density was consistently the best predictor of species richness, followed by an MCSPA metric that measures the standard deviation of woody cover across extents. At the coarser resolution (30‐m), NDVI was the best predictor of species richness by far, regardless of spatial extent. Another MCSPA metric that denotes the average woody cover across extents, together with percent of woody cover, were always the best predictors of variation in avian community structure. Spatial resolution and extent had varying effects on the relationships between spatial pattern and avian community structure. We therefore conclude that cartographic scale not only affects measures of landscape pattern per se, but also the relationships among spatial pattern, species richness, and community structure, often in complex ways, which reduces the efficacy of landscape metrics for predicting the richness and diversity of organisms.     

Multi-scale analysis of plant species richness in Serengeti grasslands

Aim To assess scale dependence between environmental factors and plant species richness. Additionally, we aimed to identify the scales at which niche relations and habitat heterogeneity, as hypothesized by A. Shmida & M.V. Wilson (1985) Journal of Biogeography, 12 , 1–20, operate in the savanna grasslands that were the focus of this study. Location Savanna grassland plant communities of Serengeti National Park, Tanzania. Methods Plant species richness was sampled in 102 modified Whittaker plots and tested for associations with two climate factors, mean annual rainfall (MAP) and potential evapotranspiration (PET), and two landscape variables, plot aspect (ASP) and topographic variation (TOPO), using multiple regressions. Scale dependence was assessed by conducting regressions after altering three aspects of spatial scale: grain, extent and focus. Grain was altered by analysing plant richness at 1, 10, 10² and 10³ m²; extent was investigated by restricting the maximum distance between samples to 75, 100, 125 and 150 km; and focus was manipulated by averaging samples spatially according to geographical land regions. Within the context of our data, we assumed that niche relations were represented by climate factors and habitat heterogeneity by landscape factors. Results Across all 102 plots, plant species richness between 1 and 10² m² had a negative relation to PET and a weak positive relation to MAP. Plant species richness at 10³ m² had a positive association with TOPO and weaker associations with climate factors. ASP stayed in the model between grains of 10 and 10³ m², but had a very weak positive association with richness. When the focus was changed to land regions, associations between plant species richness and explanatory variables strengthened, but were not qualitatively different. At spatial extents of 75 and 100 km, PET was the strongest correlate of plant species richness across all spatial grains. At spatial extents ≥ 125 km, PET explained the majority of the model variance at spatial grains ≤ 10² m², whereas TOPO explained equal amounts or more of the model variance at spatial grains of 10³ m². Main conclusions Both climate and topographic variation explained plant species richness in Serengeti grasslands, but specific patterns depended on grain, extent and, to a lesser degree, focus. Consistent with the ideas of Shmida & Wilson (1985) , determinants of plant species richness shifted from niche relations to habitat heterogeneity between spatial grains of 1 and 10³ m², although this occurred only at relatively large spatial extents (≥ 150 km). Finally, the signs, strength and shape of plant species richness relationships in Serengeti closely match those that describe macro‐scale patterns of woody plant species richness across the entire African continent.     

Primary productivity is weakly related to floristic alpha and beta diversity across Australia

Aim Ecosystem functions such as productivity may be influenced not only by the biological diversity at each location (α‐diversity) but also by the biological turnover between locations (β‐diversity). We perform a continental‐scale test of the strength and direction of relationships between gross primary productivity (GPP) and both α‐ and β‐diversity. Location Continental Australia. Methods Species occurrence records were used to quantify the taxonomic α‐diversity of vascular plants in approximately 11,000 1 km × 1 km grid cells across Australia, and to calculate the average β‐diversity within a 10‐km radius around each cell. The magnitude and variability of monthly, MODIS‐derived remotely sensed GPP (2001–12) were summarized for continental Australia, as were rainfall and temperature over the same period. Generalized additive models were then used to test whether the magnitude or variability of GPP were distinctly influenced by either biodiversity measure, over and above the influence of environmental conditions. Results Precipitation and temperature explained large proportions of deviance in the magnitude (75.6%) and variability (38.3%) of GPP across the Australian continent. GPP was marginally more strongly related to species richness than it was to species turnover. However, neither diversity measure provided substantial increases in the explanatory power of GPP models over and above that of environment‐only models (always < 1%). Main conclusions The relationship between primary productivity and taxonomic α‐ and β‐diversity was weak for the Australian flora. Our findings question the generality of key assumptions, predictions and results in the literature regarding the strength of empirical relationships between productivity and biodiversity across multiple biological levels (α‐, β‐ and γ‐diversity) at macroecological scales.     

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.     

Species richness patterns of obligate subterranean beetles (Insecta: Coleoptera) in a global biodiversity hotspot – effect of scale and sampling intensity

We studied species richness patterns of obligate subterranean (troglobiotic) beetles in the Dinaric karst of the western Balkans, using five grid sizes with cells of 80 × 80, 40 × 40, 20 × 20, 10 × 10, and 5 × 5 km. The same two hotspots could be recognized at all scales, although details differed. Differences in sampling intensity were not sufficient to explain these patterns. Correlations between number of species and number of sampled localities increased with increasing cell size. Additional species are expected to be found in the region, as indicated by jackknife 1, jackknife 2, Chao2, bootstrap, and incidence‐based coverage (ICE) species richness estimators. All estimates increased with increasing cell size, except Chao2, with the lowest prediction at the intermediate 20 × 20 km cell size. Jackknife 2 and ICE gave highest estimates and jackknife 1 and bootstrap the lowest. Jackknife 1 and bootstrap estimates changed least with cell size, while the number of single cell species increased. In the highly endemic subterranean fauna with many rare species, bootstrap may be most appropriate to consider. Positive autocorrelation of species numbers was highest at 20 × 20 km scale, so we used this cell size for further analyses. At this scale we added 137 localities with less positional accuracy to 1572 previously considered, and increased 254 troglobiotic species considered to 276. Previously discovered hotspots and their positions did not change, except for a new species‐rich cell which appeared in the south‐eastern region. There are two centres of troglobiotic species richness in the Dinaric karst. The one in the north‐west exhibited high species richness of Trechinae (Carabidae), while in the south‐east, the Leptodirinae (Cholevidae) were much more diverse. These centres of species richness should serve as the starting point for establishing a conservation network of important subterranean areas in Dinaric karst.     

Mapping patterns of ferns species richness through the use of herbarium data

This paper aims to analyse the spatial patterns of sampling effort and species richness of pteridophyte in a well-investigated region as Tuscany, Italy, by using data stored from a geodatabase storing information on the specimens preserved in the main herbaria of the region. A total of 6,905 records about pteridophyte specimens were extracted from the geodatabase, and 5,638 of such specimens were studied through the use of spatial statistical techniques. The data about the sampling effort and species richness were analysed in relation to topographical variables to assess any significant relationship. Specimen-based rarefaction techniques were used to compare areas with different number of detected species. The analysis of the sampling effort data showed a nonhomogeneous distribution of herbarium data, with some areas being intensively sampled and others being almost unsampled. Thus, the geographical distribution of specimens was extremely clustered. The comparison across geographical areas through specimen-based rarefaction curves showed great differences in species richness and sampling completeness. The analysis of the residuals of species–area relationships evidenced that the distance to water bodies was the only significant topographical variable in controlling species diversity.     

Productivity–species richness relationship changes from unimodal to positive linear with increasing spatial scale in the Inner Mongolia steppe

Productivity–species diversity relationships have been a controversial research topic in ecology with scale believed to be among the main reasons for discovering different relationships. We collected data on species diversity (richness) and productivity (peak above-ground biomass) of the Stipa breviflora association in the Inner Mongolia grassland to examine spatial scale dependency and possible underlying mechanisms responsible for the relationships found. One local and seven different landscape scales (the first level corresponds in extent to a 100 × 100 km area, which is increased consecutively by 100 km resulting in the 700 × 700 km area at the highest level) were considered. We found that: (1) unimodal relationships dominated the local scale, but this varied depending on the position along successional gradients; (2) a positive linear relationship was common at larger spatial scales; (3) biotic processes were the most likely primary factor underlying local scale unimodal relationships, but environmental heterogeneity (precipitation patterns) was the main determinant of relationships found at larger spatial scales; (4) our study contributed to other empirical evidence and predictions of theoretical models regarding scale dependency of productivity–species richness relationships; (5) while earlier research demonstrated positive linear species richness–productivity relationships across a number of ecological scales in the Inner Mongolia steppe, our study specifically tested a spectrum of geographical scales to confirm the scale-dependency of this relationship. Lastly, our study emphasized the critical role played by precipitation patterns in controlling biodiversity and grassland ecosystem functioning, which maintains the relatively high level of biodiversity and stable ecosystem processes.     

Relationship between avian range limits and plant transition zones in Maine

Aim To determine if vegetation complexity associated with transition zones may be a contributing factor affecting bird species distributions in Maine, USA, and in increased numbers of bird species at about 45° north latitude in northeastern North America. Location Maine, USA; North America north of Mexico. Methods We delineated the ranges within Maine (86,156 km²) of 186 bird species and 240 woody plants using literature and expert review. Maps showing species richness and numbers of range limits, at 324 km² resolution, were developed for woody plants and groups of breeding birds: forest specialists, forest generalists, and those that used barren and urban habitats, early successional areas, and wetlands or open water. Two plant transition zones for Maine were identified previously, with the north–south transition zone mapped across eastern North America. Patterns in bird distribution maps were compared to woody plant maps and to transition zones. Results When the distributions of forest specialists were compared to the north–south vegetation transition zone in Maine, they were spatially coincident, but were not for other groups. Forest specialists had more species with range limits in the state (61%) than generalists (13%) or any other group. At a continental‐scale, the vegetation transition zone within eastern North America agreed fairly well with the areas of highest bird richness. Main conclusions A bird transition zone occurs in Maine and across eastern North America, akin to and overlapping the vegetation transition zone. Seasonality is likely the primary source of the inverse gradient in bird richness in the eastern USA, as reported by others. However, vegetation structure and habitat selection at very broad spatial scales appear to contribute to the reversed gradient. North of the vegetation transition zone, forest structure is simpler and coniferous forests more dominant, and this may contribute to reduced bird species richness. However, the northern (> 49°) typical gradient in bird species richness has been related to many hypotheses, and several are likely involved in the genesis of the gradient.     

Relative importance of water, energy, and heterogeneity in determining regional pteridophyte and seed plant richness in China

Environmental variables, such as ambient energy, water availability, and environmental heterogeneity have been frequently proposed to account for species diversity gradients. How taxon-specific functional traits define large-scale richness gradients is a fundamental issue in understanding spatial patterns of species diversity, but has not been well documented. Using a large dataset on the regional flora from China, we examine the contrast spatial patterns and environmental determinants between pteridophytes and seed plants which differ in dispersal capacity and environmental requirements. Pteridophyte richness shows more pronounced spatial variation and stronger environmental associations than seed plant richness. Water availability generally accounts for more spatial variance in species richness of pteridophytes and seed plants than energy and heterogeneity do, especially for pteridophytes which have high dependence on moist and shady environments. Thus, pteridophyte richness is disproportionally affected by water-related variables; this in turn results in a higher proportion of pteridophytes in regional vascular plant floras (pteridophyte proportion) in wet regions. Most of the variance in seed plant richness, pteridophyte richness, and pteridophyte proportion explained by energy is included in variation that water and heterogeneity account for, indicating the redundancy of energy in the study extent. However, heterogeneity is more important for determining seed plant distributions. Pteridophyte and seed plant richness is strongly correlated, even after the environmental effects have been removed, implying functional linkages between them. Our study highlights the importance of incorporating biological traits of different taxonomic groups into the studies of macroecology and global change biology.     

Habitat availability does not explain the species richness patterns of European lentic and lotic freshwater animals

Aim In Europe, the relationships between species richness and latitude differ for lentic (standing water) and lotic (running water) species. Freshwater animals are highly dependent on suitable habitat, and thus the distribution of available habitat should strongly influence large‐scale patterns of species richness. We tested whether habitat availability can account for the differences in species richness patterns between European lentic and lotic freshwater animals. Location Europe. Methods We compiled occurrence data of 1959 lentic and 2445 lotic species as well as data on the amount of lentic and lotic habitats across 25 pre‐defined biogeographical regions of European freshwaters. We used the range of elevation of each region as a proxy for habitat diversity. We investigated the relationships between species richness, habitat availability and habitat diversity with univariate and multiple regression analyses. Results Species richness increased with habitat availability for lentic species but not for lotic species. Species richness increased with elevational range for lotic species but decreased for lentic species. For both groups, neither habitat availability nor diversity could account for previously reported latitudinal patterns in species richness. For lotic species, richness declined with latitude, whereas there was no relationship between habitat availability and latitude. For lentic species, richness showed a hump‐shaped relationship with latitude, whereas available habitat increased with latitude. Main conclusions Habitat availability and diversity are poor predictors of species richness of the European freshwater fauna across large scales. Our results indicate that the distributions of European freshwater animals are probably not in equilibrium and may still be influenced by history, namely the recurrent European glaciations and possible differences in post‐glacial recolonization. The distributions of lentic species appear to be closer to equilibrium than those of lotic species. This lends further support to the hypothesis that lentic species have a higher propensity for dispersal than lotic species.