Beta‐diversity gradients of butterflies along productivity axes

Aim Several lines of evidence suggest that beta diversity, or dissimilarity in species composition, should increase with productivity: (1) the latitudinal species richness gradient is most closely related to productivity and associated latitudinal beta‐diversity relationships have been described, and (2) the scale dependence of the productivity–diversity relationship implies that there should be a positive productivity–beta‐diversity relationship. However, such a pattern has not yet been demonstrated at broad scales. We test if there is a gradient of increasing beta diversity with productivity. Location Canada. Methods Canada was clustered into regions of similar productivity regimes along three remotely sensed productivity axes (minimum and integrated annual productivity, seasonality of productivity) and elevation. The overall (β_j), turnover (β_sim) and nestedness (β_nes) components of beta diversity within each productivity regime were estimated with pairwise dissimilarity metrics and related to cluster productivity with partial linear regression and with spatial autoregression. Tests were performed for all species, productivity breadth‐based subsets (e.g. species occurring in many and a moderate number of productivity regimes), and pre‐ and post‐1970 butterfly records. Beta diversity between adjacent clusters along the productivity gradients was also evaluated. Results Within‐cluster β_j and β_sim increased with productivity and decreased with seasonality. The converse was true for β_nes. All species subsets responded similarly; however, productivity–beta‐diversity relationships were weaker for the post‐1970 temporal subset and strongest for species of moderate breadth. Between‐cluster beta diversity (β_j) and nestedness (β_nes) declined with productivity. Main conclusions As predicted, beta diversity of communities within productivity regimes was observed to increase with productivity. This pattern was driven largely by a gradient of species turnover. Therefore, beta diversity may make an important contribution to the broad‐scale gradient of species richness with productivity. However, this species richness gradient dominates regional beta diversity between productivity regimes, resulting in decreasing between‐productivity dissimilarity with productivity driven by a concurrent decline in nestedness.     

Measuring fractions of beta diversity and their relationships to nestedness: a theoretical and empirical comparison of novel approaches

Beta diversity and nestedness are central concepts of ecology and biogeography and evaluation of their relationships is in the focus of contemporary ecological and conservation research. Beta diversity patterns are originated from two distinct processes: the replacement (or turnover) of species and the loss (or gain) of species leading to richness differences. Nested distributional patterns are generally thought to have a component deriving from beta diversity which is independent of replacement processes. Quantification of these phenomena is often made by calculating a measure of beta diversity, and the resulting value being subsequently partitioned into a contribution by species replacement plus a fraction shared by beta diversity and nestedness. Three methods have been recently proposed for such partitioning, all of them based on pairwise comparisons of sites. In this paper, the performance of these methods was evaluated on theoretical grounds and tested by a simulation study in which different gradients of dissimilarity, with known degrees of species replacement and species loss, were created. Performance was also tested using empirical data addressing land‐use induced changes in endemic arthropod communities of the Terceira Island in the Azores. We found that the partitioning of β_cc (dissimilarity in terms of the Jaccard index) into two additive fractions, β_‐3 (dissimilarity due to species replacement) plus β_rich (dissimilarity due to richness differences) reflects the species replacement and species loss processes across the simulated gradients in an ecologically and mathematically meaningful way, whilst the other two methods lack mathematical consistency and prove conceptually self‐contradictory. Moreover, the first method identified a selective local extinction process for endemic arthropods, triggered by land‐use changes, while the latter two methods overweighted the replacement component and led to false conclusions. Their basic flaw derives from the fact that the proposed replacement and nestedness components (deemed to account for species loss) are not scaled in the same way as the measure that accounts for the total dissimilarity (Sørensen and Jaccard indices). We therefore recommend the use of β_cc=β_‐3+β_rich, since its components are scaled in the same units and their responses are proportional to the replacement and the gain/loss of species.     

Rethinking the relationship between nestedness and beta diversity: a comment on Baselga (2010)

Baselga [Partitioning the turnover and nestedness components of beta diversity. Global Ecology and Biogeography, 19 , 134–143, 2010] proposed pairwise (β_nes) and multiple‐site (β_NES) beta‐diversity measures to account for the nestedness component of beta diversity. We used empirical, randomly created and idealized matrices to show that both measures are only partially related to nestedness and do not fit certain fundamental requirements for consideration as true nestedness‐resultant dissimilarity measures. Both β_nes and β_NES are influenced by matrix size and fill, and increase or decrease even when nestedness remains constant. Additionally, we demonstrate that β_NES can yield high values even for matrices with no nestedness. We conclude that β_nes and β_NES are not true measures of the nestedness‐resultant dissimilarity between sites. Actually, they quantify how differences in species richness that are not due to species replacement contribute to patterns of beta diversity. Finally, because nestedness is a special case of dissimilarity in species composition due to ordered species loss (or gain), the extent to which differences in species composition is due to nestedness can be measured through an index of nestedness.     

Dispersal ability modulates the strength of the latitudinal richness gradient in European beetles

Aim A debate exists as to whether present‐day diversity gradients are governed by current environmental conditions or by changes in environmental conditions through time. Recent studies have shown that latitudinal richness gradients might be partially caused by incomplete post‐glacial recolonization of high‐latitude regions; this leads to the prediction that less mobile taxa should have steeper gradients than more mobile taxa. The aim of this study is to test this prediction. Location Europe. Methods We first assessed whether spatial turnover in species composition is a good surrogate for dispersal ability by measuring the proportion of wingless species in 19 European beetle clades and relating this value to spatial turnover (β_sim) of the clade. We then linearly regressed β_sim values of 21 taxa against the slope of their respective diversity gradients. Results A strong relationship exists between the proportion of wingless species and β_sim, and β_sim was found to be a good predictor of latitudinal richness gradients. Main conclusions Results are consistent with the prediction that poor dispersers have steeper richness gradients than good dispersers, supporting the view that current beetle diversity gradients in Europe are affected by post‐glacial dispersal lags.     

The need for richness‐independent measures of turnover when delineating biogeographical regions

Delineating biogeographical regions is one of the primary steps when analysing biogeographical patterns. In their proposed quantitative framework, Kreft & Jetz (2010, Journal of Biogeography, 37 , 2029–2053) recommended the use of the β_sim index to delineate biogeographical regions because this turnover measure is weakly affected by differences in species richness between localities. A recent study by Carvalho et al. (2012, Global Ecology and Biogeography, 21 , 760–771) critiziced the use of β_sim in ecological and biogeographical studies, and proposed the β_‐3 index. Here we used simple numerical examples and an empirical case study (European freshwater fishes) to highlight potential pitfalls associated with the use of β_‐3 for bioregionalization. We show that β_‐3 is not a richness‐independent measure of species turnover. We also show that this index violates the ‘complementarity’ property, namely that localities without species in common have the largest dissimilarity, which is an essential prerequisite for beta diversity studies.     

Testing the performance of beta diversity measures based on incidence data: the robustness to undersampling

Aim Researchers measuring beta diversity have rarely concerned themselves with the problems of how complete the species lists of studied communities are, and of how the varying degrees of completeness can actually change estimates of beta diversity. No comprehensive assessment has been made regarding the behaviour of most beta diversity indices when applied to incomplete samples, a situation which is more common than usually recognized. Our objective was to assess the behaviour and robustness of a number of beta diversity measures for incidence data from undersampled communities. Location Mainland Portugal and the Azorean archipelago (North Atlantic). Methods Data from intensive sampling of spiders in mainland Portugal and arthropods in Azores were collected. We examined the properties of 15 beta diversity measures developed for incidence data. We simulated varying degrees of completeness, whereas computing beta diversity for selected pairs of samples. The robustness of these beta diversity accumulation curves was assessed for the purpose of finding the best measures for undersampled communities. Results The Harrison et al.β_‐2 and the Williams β_‐3 are particularly robust to undersampling. These measures are also insensitive to differences of alpha diversity (species richness) between communities, and therefore to nestedness. Colwell & Coddington β_cc and the related Jaccard β_j and Gaston et al.β_g performed best of the measures sensitive to alpha diversity. They performed poorly, however, when compared communities exhibited very low values of beta diversity. In such cases, the Routledge β_r performed the best. Main conclusions No index was found to perform without bias in all circumstances. Overall, β_‐2, β_‐3 and β_cc (or related measures β_j and β_g) are recommended as they seem to be the most robust to undersampling.     

Beta diversity in relation to dispersal ability for vascular plants in North America

Aim To test the hypothesis that plant species with a higher dispersal ability have a lower beta diversity. Location North America north of Mexico. Method Propagules of pteridophytes (ferns and their allies) are more vagile than propagules of spermatophytes (gymnosperms and angiosperms), and thus pteridophytes have a higher dispersal ability than do spermatophytes. The study area was divided into 71 geographical units distributed in five latitudinal zones. Species lists of pteridophytes and spermatophytes were compiled for each geographical unit. Three measures of beta diversity were used: β_sim, which is one minus the Simpson index of similarity, β_slope, which is the slope of the relationship between Simpson index and geographical distance, and β_{0.5‐distance}, which is the distance that halves the similarity from its initial value. Results Average β_sim is higher for spermatophytes than for pteridophytes, regardless of whether the data are analysed for the whole continent or for latitudinal zones. Average β_sim decreases with increasing latitude for both spermatophytes and pteridophytes. The difference in average β_sim between the two plant groups increases with increasing latitude, indicating that beta diversity decreases with increasing latitude faster for pteridophytes than for spermatophytes. When the Simpson index is regressed against geographical distance, the regression slope (β_slope) is steeper for spermatophytes than for pteridophytes, and the slope decreases with increasing latitude for both plant groups. Similarly, β_{0.5‐distance} was shorter for spermatophytes than for pteridophytes in each latitudinal zone and increased with increasing latitude for both plant groups. The results of the analyses using the three different measures of beta diversity are consistent. Main conclusions The fact that beta diversity is lower for pteridophytes with vagile propagules than for spermatophytes with less vagile propagules suggests that beta diversity is negatively related to dispersal ability.     

The effect of species composition dissimilarity on plant–herbivore network structure is not consistent over time

The structural organization of several antagonistic networks has been demonstrated to be largely conserved through time and space even when species beta-diversity is high. This might occur either because species are replaced by others that fulfill similar network roles or because interaction probabilities are given by species relative abundances rather than by their functional traits. Alternatively, if species-specific traits are important drivers of realized interactions, any change in species composition should promote a certain degree of network structural dissimilarity. Here, we used a spatial-temporal system comprising asteraceous plants and flower head herbivores from remnants of Brazilian Cerrado to investigate whether the relationship between spatial beta-diversity of species and network structural dissimilarity changes over time. We measured species beta-diversity using Sørensen's dissimilarity index (β_sor) and its components of species replacement (β_-3) and richness differences (β_rich). Network structural dissimilarity was estimated using three different metrics: connectance, modularity, and web asymmetry. We show that, in general, the effect of species beta-diversity on network structure was time-dependent: While some periods presented a positive relationship between spatial beta-diversity and network structural dissimilarity, others presented no significant relationship. This indicates that functionally similar species may present different turnover rates at distinct periods, and different non-exclusive processes affect plant–herbivore network organization across time.     

Plant species richness and diversity along an altitudinal gradient in the Sierra Nevada, Mexico

This article presents an analysis of plant species richness and diversity and its association with climatic and soil variables along a 1300‐m elevation gradient on the Cerro Tláloc Mountain in the northern Sierra Nevada in Mexico. Two 1000‐m² tree sampling plots were created at each of 21 selected sampling sites, as well as two 250‐m² plots for shrubs and six 9‐m² plots for herbaceous plants. Species richness and diversity were estimated for each plant life form, and beta diversity between sites was estimated along the gradient. The relationship between species richness and diversity and environmental variables was modelled using simple linear correlation and regression trees. Species richness and diversity showed a unimodal pattern with a bias towards high values in the lower half of the elevation gradient under study. This response was consistent for all three life forms. Beta diversity increased steadily along the elevation gradient, being lower between contiguous sites at intermediate elevations and high – the species replacement rate was nearly 100%– between sites at the extremes of the gradient. Few species were adapted to the full spectrum of environmental variation along the elevation gradient studied. The regression tree suggests that differences in species richness are mainly influenced by elevation (temperature and humidity) and soil variables, namely A₂ permanent wilting point, organic matter and horizon field capacity and A₁ horizon Mg²⁺.     

Patterns of plant beta‐diversity along elevational and latitudinal gradients in mountain forests of China

Biodiversity patterns and their underlying mechanisms have long been focal topics of study for ecologists and biogeographers. However, compared with spatial variation in species richness (α‐ and γ‐diversity), β‐diversity, or the dissimilarity of species composition between two or more sites has until recently received limited attention. In this study, we explored the large‐scale patterns of altitudinal turnover (β‐diversity) of plants in montane forests of China, based on systematic inventories of 1153 plots from 46 mountains distributed over ?30 degrees of latitude (21.9–51.7°N) and ?4100 m of altitude (160–4250 m). The β‐diversity of trees and shrubs declined significantly with increasing latitude. Along the altitudinal gradient, β‐diversity of both trees and shrubs showed non‐significant trends in most mountains. Differences in climate explained ?30.0% of the variation in tree β‐diversity (27.7, 36.5, and 26.2% for the Jaccard's, β_j, Sorenson's, β_s, and Simpson's dissimilarity, β_sim, respectively), with mean annual temperature being most important, and ≤ 10.0% of that in shrub β‐diversity (10.0, 8.2, and 7.0% for β_j, β_s, and β_sim, respectively), with annual actual evapotranspiration and annual precipitation as the main predictors. However, climatic controls of β‐diversity varied dramatically in different biogeograpical regions. The β‐diversity of trees exhibited stronger, whereas that of shrubs showed weaker, climatic patterns in temperate and arid than subtropical regions. These results suggest that mechanisms causing patterns of β‐diversity may differ between latitudinal and altitudinal gradients, and among biogeographical regions; as a result, caution should be exercised in drawing close parallels between patterns and causes of β‐diversity along latitudinal and altitudinal gradients and among regions.     

Independence of total species richness and richness difference are not the same thing

Leprieur and Oikonomou (2014) criticized that a replacement index β_− 3, a partitioned component of Jaccard index β_jac, was not richness independent and should not be used in biogeographic and ecological studies. However, Leprieur and Oikonomou failed to recognize the difference between richness and richness difference. Independence of total species richness is not equal to independence of richness difference. Theoretically and ideally, it is true that β_− 3 (and β_jac as well) is not independent of richness difference while Simpson index (β_sim) is fully independent of richness difference. However, all these indices actually are independent of total species richness. At last, it is worth mentioning that the ideal independence studied here is easily violated in computational simulation and real-world settings.     

From plots to islands: species diversity at different scales

Aim To investigate how plant diversity of whole islands (‘gamma’) is related to alpha and beta diversity patterns among sampling plots within each island, thus exploring aspects of diversity patterns across scales. Location Nineteen islands of the Aegean Sea, Greece. Methods Plant species were recorded at both the whole‐island scale and in small 100 m² plots on each island. Mean plot species richness was considered as a measure of alpha diversity, and six indices of the ‘variation’‐type beta diversity were also applied. In addition, we partitioned beta diversity into a ‘nestedness’ and a ‘replacement’ component, using the total species richness recorded in all plots of each island as a measure of ‘gamma’ diversity. We also applied 10 species–area models to predict the total observed richness of each island from accumulated plot species richness. Results Mean alpha diversity was not significantly correlated with the overall island species richness or island area. The range of plot species richness for each island was significantly correlated with both overall species richness and area. Alpha diversity was not correlated with most indices of beta diversity. The majority of beta diversity indices were correlated with whole‐island species richness, and this was also true for the ‘replacement’ component of beta diversity. The rational function model provided the best prediction of observed island species richness, with Monod’s and the exponential models following closely. Inaccuracy of predictions was positively correlated with the number of plots and with most indices of beta diversity. Main conclusions Diversity at the broader scale (whole islands) is shaped mainly by variation among small local samples (beta diversity), while local alpha diversity is not a good predictor of species diversity at broader scales. In this system, all results support the crucial role of habitat diversity in determining the species–area relationship.     

Phylogeographic patterns of Merodon hoverflies in the Eastern Mediterranean region: revealing connections and barriers

We investigated the phylogeographic patterns of Merodon species (Diptera, Syrphidae) in the Eastern Mediterranean. Ten species were sampled on five different islands and mainland sites as a minimum. All samples were screened for their mtDNA COI barcode haplotype diversity, and for some samples, we additionally generated genomic fingerprints. The recently established zoogeographic distribution categories classify these species as having (1) Balkan distribution; (2) Anatolian distribution; (3) continental areas and large islands distribution; and (4) with wide distribution. The ancestral haplotypes and their geographical localities were estimated with statistical parsimony (TCS). TCS networks identified as the ancestral haplotype samples that originated from localities situated within the distributional category of the species in question. Strong geographical haplotype structuring was detected for many Merodon species. We were particularly interested to test the relative importance of current (Aegean Sea) and past Mid‐Aegean Trench) barriers to dispersal for Merodon flies in the Aegean. We employed phylogenetic β‐diversity (Pβ_total) and its partition in replacement (Pβ_repl) and richness difference (Pβ_rich) to test the importance of each explanatory variable (interisland distance, MAT, and island area) in interisland differences using partial Mantel tests and hierarchical partitioning of variation. β‐Analyses confirmed the importance of both current and past barriers to dispersal on the evolution of group. Current interisland distance was particularly important to explain the replacement of haplotypes, while the MAT was driving differences in richness of haplotypes, revealing the MAT as a strong past barrier whose effects are still visible today in the phylogenetic history of the clade in the Aegean. These results support the hypothesis of a highly restricted dispersal and gene flow among Merodon populations between islands since late Pleistocene. Additionally, patterns of phylogeographic structure deduced from haplotype connections and ISSR genome fingerprinting data revealed a few putative cases of human‐mediated transfers of Merodon spp.     

Patterns of beta diversity in Europe: the role of climate,land cover and distance across scales

Aim We test the prediction that beta diversity (species turnover) and the decay of community similarity with distance depend on spatial resolution (grain). We also study whether patterns of beta diversity are related to variability in climate, land cover or geographic distance and how the independent effects of these variables depend on the spatial grain of the data. Location Europe, Great Britain, Finland and Catalonia. Methods We used data on European birds, plants, butterflies, amphibians and reptiles, and data on British plants, Catalonian birds and Finnish butterflies. We fitted two or three nested grids of varying resolutions to each of these datasets. For each grid we calculated differences in climate, differences in land‐cover composition (CORINE) and beta diversity (β_sim, β_Jaccard) between all pairs of grid cells. In a separate analysis we looked specifically at pairs of adjacent grid cells (the first distance class). We then used variation partitioning to identify the magnitude of independent statistical associations (i.e. independent effects in the statistical sense) of climate, land cover and geographic distance with spatial patterns of beta diversity. Results Beta diversity between grid cells at any given distance decreased with increasing grain. Geographic distance was always the most important predictor of beta diversity for all pairwise comparisons at the extent of Europe. Climate and land cover had weaker but distinct and grain‐dependent effects. Climate was more important at relatively coarse grains, whereas land‐cover effects were stronger at finer grains. In the country‐wide analyses, climate and land cover were more important than geographic distance. Climatic and land‐cover models performed poorly and showed no systematic grain dependence for beta diversity between adjacent grid cells. Main conclusions We found that relationships between geographic distance and beta diversity, as well as the environmental correlates of beta diversity, are systematically grain dependent. The strong independent effect of distance indicates that, contrary to the current belief, a substantial fraction of species are missing from areas with a suitable environment. Moreover, the effects of geographic distance (at continental extents) and land cover (at fine grains) indicate that any species distribution modelling should take both environment and dispersal limitation into account.     

Patterns and gradients of diversity in South Patagonian ombrotrophic peat bogs

Many north‐hemispherical mires seemingly untouched by drainage and cultivation are influenced by a diffuse sum of man‐made environmental changes, such as atmospherical nitrogen deposition that mask general patterns in species richness and functional group responses along resource gradients. To obtain insights into natural diversity‐environment relationships, we studied the vegetation and the peat chemistry of pristine bog ecosystems in southern Patagonia along a west–east transect across the Andes. The studied bog ecosystems covered a floristic gradient from hyperoceanic blanket bogs dominated by cushion building vascular plants via a transitional mixed type to Sphagnum‐dominated raised bogs east of the mountain range. To test the influence of resource availability on diversity patterns, species richness and functional groups were related to environmental variables by calculating general regression models and generalized additive models. Species richness showed strong linear correlations to peat chemical features and the general regression model resulted in three major environmental variables (water level, total nitrogen, NH₄Cl soluble calcium), altogether explaining 76% of variance. Functional group response illustrated a clear separation along environmental gradients. Mosses dominated at the low end of a nitrogen gradient, whereas cushion plants had their optimum at intermediate levels, and graminoids dominated at high nitrogen contents. Further shifts were related to NH₄Cl soluble calcium and water level. The models documented partly non‐linear relationships between functional group response and trophical peat properties. Within the three bog types, the calculated models differed remarkably illustrating the scale‐dependency of the explanatory factors. Our findings confirmed several general patterns of species richness and functional shifts along resource gradients in a surprisingly clear way and underpin the significance of undisturbed peatlands as reference systems for testing of ecological theory and for conservation and ecological restoration in landscapes with strong human impact.     

Temporal changes in fish diversity in lotic and lentic environments along a reservoir cascade

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Energy and speleogenesis: Key determinants of terrestrial species richness in caves

The aim of this study was to unravel the relative role played by speleogenesis (i.e., the process in which a cave is formed), landscape‐scale variables, and geophysical factors in the determination of species richness in caves. Biological inventories from 21 caves located in the southeastern Iberian Peninsula along with partial least square (PLS) regression analysis were used to assess the relative importance of the different explanatory variables. The caves were grouped according to the similarity in their species composition; the effect that spatial distance could have on similarity was also studied using correlation between matrices. The energy and speleogenesis of caves accounted for 44.3% of the variation in species richness. The trophic level of each cave was the most significant factor in PLS regression analysis, and epigenic caves (i.e., those formed by the action of percolating water) had significantly more species than hypogenic ones (i.e., those formed by the action of upward flows in confined aquifers). Dissimilarity among the caves was very high (multiple‐site β_sim = 0.92). Two main groups of caves were revealed through the cluster analysis, one formed by the western caves and the other by the eastern ones. The significant—but low—correlation found between faunistic dissimilarity and geographical distance (r = .16) disappeared once the caves were split into the two groups. The extreme beta‐diversity suggests a very low connection among the caves and/or a very low dispersal capacity of the species. In the region under study, two main factors are intimately related to the richness of terrestrial subterranean species in caves: the amount of organic material (trophic level) and the formation process (genesis). This is the first time that the history of a cave genesis has been quantitatively considered to assess its importance in explaining richness patterns in comparison with other factors more widely recognized.     

Spatial gradients in species diversity of microscopic animals: the case of bdelloid rotifers at high altitude

Aim Organisms smaller than 2 mm appear not to follow the spatial patterns in richness and diversity commonly observed in macroscopic organisms. We describe spatial patterns in species diversity in a group of microscopic organisms, bdelloid rotifers, living in moss and lichen patches, in order to test the hypotheses of no relationship between species richness and composition and spatial gradients, suggested by previously published patterns in microscopic organisms. Location Moss and lichen patches as habitats for bdelloids, on high‐elevation peaks at altitudes between 2984 and 4527 m a.s.l. across the Italian, French and Swiss Alps, with distances among sample sites ranging from 1 m to 420 km, in comparison with lower‐elevation samples at altitudes from 850 to 1810 m a.s.l. Methods We sampled species assemblages of bdelloid rotifers living in isolated moss and lichen patches in 47 sites. We described the observed α, β and γ diversities; the heterogeneity of species assemblages; and the estimated number of species (incidence‐based coverage estimator). Patterns in species distribution were analysed at three different levels: (1) habitat, comparing species richness on moss and lichen substrates, testing differences in α diversity and heterogeneity (anova ), species composition (analysis of similarities test), and γ diversity (rarefaction curves); (2) altitude, comparing the observed richness with previously published data from locations well below 2000 m; and (3) distances between sites, correlating the matrix of Jaccard dissimilarities and the matrix of geographical distances with a Mantel test. Results Both species richness and species composition of bdelloid rotifers differed significantly between mosses and lichens at high elevations, but no difference was found in the heterogeneity of species assemblages. Alpha diversity was significantly lower at high‐elevation than at low‐elevation sites, but the estimated number of species was not reduced when compared with sites at low elevations. Geographical distance between sites had no effect on species composition in either mosses or lichens. The distribution of species was highly heterogeneous, with a low similarity among assemblages. Main conclusions As expected, bdelloids appear to occupy habitats selectively. The altitudinal gradient in species richness for bdelloid rotifers is limited to a decrease in α diversity only; such a decrease is not caused by a lower number of species (low γ diversity) being able to tolerate harsh conditions, and high‐altitude species are not a subset of species living at lower elevations. The observed values of α, β and γ diversity at high altitudes in the Alps are compatible with the scenario of a very low number of available propagules because of the low density of patches of favourable habitat. Our results suggest that the geographical distribution of animals, and therefore biodiversity patterns, may be strongly influenced by animal size, as small organisms such as bdelloids appear to show spatial patterns that differ from those known in larger animals. Differences in body size should be taken into account carefully in future studies of biodiversity patterns.     

Declines in littoral species richness across both spatial and temporal nutrient gradients: a palaeolimnological study of two taxonomic groups

1. Using a palaeolimnological approach in shallow lakes, we quantified the species richness responses of diatoms and Cladocera to phosphorus enrichment. We also examined differences in species richness responses between littoral and pelagic assemblages of our focal communities. To address both spatial and temporal relationships, our study includes an analysis of both surface sediments from 40 lakes and of a lake sediment record spanning c. 120 years. The objective of our study was to determine whether similar species richness patterns occurred across trophic levels, as well as along spatial and temporal gradients. 2. We found that both diatom and Cladocera species richness estimates significantly declined with increasing phosphorus across space and through time. When the assemblages were subdivided according to known habitat preferences, littoral biodiversity maintained a negative trend, whereas pelagic species richness tended to show no relationship with phosphorus. 3. Negative productivity–diversity patterns have been observed across almost all palaeolimnological studies that span large productivity gradients. This congruence in patterns is most likely due to the similarity in data collection methods and in focal communities studied. The contrasting responses between littoral and pelagic assemblages may be explained by the differences in physical habitat and the pool of taxa in each of these environments. Consistent with the literature, we found statistical support for the idea that littoral diversity declines could be explained by an interaction between macrophytes and nutrients along strong trophic gradients. The general lack of a diversity response in our pelagic assemblages could be attributable to the limited pool of subfossil taxa. The response of the pelagic diatom could also be related to their broad range of nutrient tolerances. 4. The observed negative response of species richness to phosphorus enrichment, particularly in the littoral assemblages, has implications for ecosystems functioning because communities with reduced biodiversity often are less resilient to anthropogenic change.