The tangled link between β‐ and γ‐diversity: a Narcissus effect weakens statistical inferences in null model analyses of diversity patterns

Understanding the structure of and spatial variability in the species composition of ecological communities is at the heart of biogeography. In particular, there has been recent controversy about possible latitudinal trends in compositional heterogeneity across localities (β‐diversity). A gradient in the size of the regional species pool alone can be expected to impose a parallel gradient on β‐diversity, but whether β‐diversity also varies independently of the size of the species pool remains unclear. A recently suggested methodological approach to correct latitudinal β‐diversity gradients for the species pool effect is based on randomization null models that remove the effects of gradients in α‐ and γ‐diversity on β‐diversity. However, the randomization process imposes constraints on the variability of α‐diversity, which in turn force γ‐ and β‐diversity to become interdependent, such that any change in one is mirrored in the other. We argue that simple null model approaches are inadequate to discern whether correlations between α‐, β‐ and γ‐diversity reflect processes of ecological interest or merely differences in the size of the species pool among localities. We demonstrate that this kind of Narcissus effect may also apply to other metrics of spatial or phylogenetic species distribution. We highlight that Narcissus effects may lead to artificially high rejection rates for the focal pattern (Type II errors) and caution that these errors have not received sufficient attention in the ecological literature.     

Environmental heterogeneity among lakes promotes hyper β‐diversity across phytoplankton communities

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Hot,dry and different: Australian lizard richness is unlike that of mammals,amphibians and birds

Aims (1) To map the species richness of Australian lizards and describe patterns of range size and species turnover that underlie them. (2) To assess the congruence in the species richness of lizards and other vertebrate groups. (3) To search for commonalities in the drivers of species richness in Australian vertebrates. Location Australia. Methods We digitized lizard distribution data to generate gridded maps of species richness and β‐diversity. Using similar maps for amphibians, mammals and birds, we explored the relationship between species richness and temperature, actual evapotranspiration, elevation and local elevation range. We used spatial eigenvector filtering and geographically weighted regression to explore geographical patterns and take spatial autocorrelation into account. We explored congruence between the species richness of vertebrate groups whilst controlling for environmental effects. Results Lizard richness peaks in the central deserts (where β‐diversity is low) and tropical north‐east (where β‐diversity is high). The intervening lowlands have low species richness and β‐diversity. Generally, lizard richness is uncorrelated with that of other vertebrates but this low congruence is strongly spatially structured. Environmental models for all groups also show strong spatial heterogeneity. Lizard richness is predicted by different environmental factors from other vertebrates, being highest in dry and hot regions. Accounting for environmental drivers, lizard richness is weakly positively related to richness of other vertebrates, both at global and local scales. Main conclusions Lizard species richness differs from that of other vertebrates. This difference is probably caused by differential responses to environmental gradients and different centres of diversification; there is little evidence for inter‐taxon competition limiting lizard richness. Local variation in habitat diversity or evolutionary radiations may explain weak associations between taxa, after controlling for environmental variables. We strongly recommend that studies of variation in species richness examine and account for non‐stationarity.     

Global patterns of stream detritivore distribution: implications for biodiversity loss in changing climates

Aim We tested the hypothesis that shredder detritivores, a key trophic guild in stream ecosystems, are more diverse at higher latitudes, which has important ecological implications in the face of potential biodiversity losses that are expected as a result of climate change. We also explored the dependence of local shredder diversity on the regional species pool across latitudes, and examined the influence of environmental factors on shredder diversity. Location World‐wide (156 sites from 17 regions located in all inhabited continents at latitudes ranging from 67° N to 41° S). Methods We used linear regression to examine the latitudinal variation in shredder diversity at different spatial scales: alpha (α), gamma (γ) and beta (β) diversity. We also explored the effect of γ‐diversity on α‐diversity across latitudes with regression analysis, and the possible influence of local environmental factors on shredder diversity with simple correlations. Results Alpha diversity increased with latitude, while γ‐ and β‐diversity showed no clear latitudinal pattern. Temperate sites showed a linear relationship between γ‐ and α‐diversity; in contrast, tropical sites showed evidence of local species saturation, which may explain why the latitudinal gradient in α‐diversity is not accompanied by a gradient in γ‐diversity. Alpha diversity was related to several local habitat characteristics, but γ‐ and β‐diversity were not related to any of the environmental factors measured. Main conclusions Our results indicate that global patterns of shredder diversity are complex and depend on spatial scale. However, we can draw several conclusions that have important ecological implications. Alpha diversity is limited at tropical sites by local factors, implying a higher risk of loss of key species or the whole shredder guild (the latter implying the loss of trophic diversity). Even if regional species pools are not particularly species poor in the tropics, colonization from adjacent sites may be limited. Moreover, many shredder species belong to cool‐adapted taxa that may be close to their thermal maxima in the tropics, which makes them more vulnerable to climate warming. Our results suggest that tropical streams require specific scientific attention and conservation efforts to prevent loss of shredder biodiversity and serious alteration of ecosystem processes.     

β‐diversity scaling patterns are consistent across metrics and taxa

β‐diversity (variation in community composition) is a fundamental component of biodiversity, with implications for macroecology, community ecology and conservation. However, its scaling properties are poorly understood. Here, we systematically assessed the spatial scaling of β‐diversity using 12 empirical large‐scale datasets including different taxonomic groups, by examining two conceptual types of β‐diversity and explicitly considering the turnover and nestedness components. We found highly consistent patterns across datasets. Multiple‐site β‐diversity (i.e. variation across multiple sites) scaling curves were remarkably consistent, with β‐diversity decreasing with sampled area according to a power law. For pairwise dissimilarities, the rates of increase of dissimilarity with geographic distance remained largely constant across scales, while grain size (or scale level) had a stronger effect on overall dissimilarity. In both analyses, turnover was the main contributor to β‐diversity, following total β‐diversity patterns closely, while the nestedness component was largely insensitive to scale changes. Our results highlight the importance of integrating both inter‐ and intraspecific aggregation patterns across spatial scales, which underpin substantial differences in community structure from local to regional scales.     

The latitudinal gradient of species diversity among North American grasshoppers (Acrididae) within a single habitat: a test of the spatial heterogeneity hypothesis

Abstract. The spatial heterogeneity hypothesis predicts a positive relationship between habitat complexity and species diversity: the greater the heterogeneity of a habitat, the greater the number of species in that habitat. On a regional scale, this hypothesis has been proposed to explain the increases in species diversity from the poles to the tropics: the tropics are more diverse because they contain more habitats. On the local scale, the spatial heterogeneity hypothesis suggests that the tropics are more diverse because they contain more microhabitats. The positive relationship between habitat heterogeneity and species diversity, on the local scale, is well documented. In this paper, we test whether habitat heterogeneity on the local scale can explain the latitudinal gradient of species diversity on the regional scale. We determined the latitudinal gradient of species diversity of 305 species of North American grasshoppers using published distribution maps. We compared the slope of this multihabitat (regional-scale) gradient with the slope of a within-habitat (local-scale) gradient in the prairie grasslands. Our results show no significant difference between the slopes at the two scales. We tested the generality of our results by comparing multi- and within-habitat latitudinal gradients of species diversity for ants, scorpions and mammals using data from the literature. These results are in accordance with those from grasshoppers. We can therefore reject the local-scale spatial heterogeneity hypothesis as a mechanism explaining the regional-scale latitudinal gradient of species diversity. We discuss alternative mechanisms that produce this gradient.     

Local-regional relationships and the geographical distribution of species

Aim Local‐regional (LR) species diversity plots were conceived to assess the contribution of regional and local processes in shaping the patterns of biological diversity, but have been used also to explore the scaling of diversity in terms of its alpha, beta, and gamma components. Here we explore the idea that patterns in the geographical ranges of species over a continent can determine the shape of small region to large region (SRLR) plots, which are equivalent to LR plots when comparing the diversity of sites at two regional scales. Location To test that idea, we analysed the diversity patterns at two regional scales for the mammals of North America, defined as the mainland from Alaska and Canada to Panama. Method We developed a theoretical model relating average range size of species over a large‐scale region with its average regional point species diversity (RPD). Then, we generated a null model of expected SRLR plots based on theoretical predictions. Species diversities at two scales were modelled using linear and saturation functions for Type I and Type II SRLR relationships, respectively. We applied the models to the case of North American mammals by examining the regional diversity and the RPD for 21 large‐scale quadrats (with area equal to 160,000 km²), arranged along a latitudinal gradient. Results Our model showed that continental and large‐scale regional patterns of distribution of species can generate both types of SRLR relationship, and that these patterns can be reflected in LR plots without invoking any kind of local processes. We found that North American nonvolant mammals follow a Type I SRLR relationship, whereas bats follow a Type II pattern. This difference was linked to patterns in which species of the two mammalian groups distribute in geographical space. Conclusion Traditional LR plots and the new SRLR plots are useful tools in exploring the scaling of species diversity and in showing the relationship between distribution and diversity. Their usefulness in comparing the relative role of local and regional processes is, however, very limited.     

Environmental heterogeneity and dispersal limitation explain different aspects of β‐diversity in Neotropical fish assemblages

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Ecological correlates of mammal β‐diversity in Amazonian land‐bridge islands: from small‐ to large‐bodied species

Aim

Mega hydroelectric dams have become one of the main drivers of biodiversity loss in the lowland tropics. In these reservoirs, vertebrate studies have focused on local (α) diversity measures, whereas between‐site (β) diversity remains poorly assessed despite its pivotal importance in understanding how species diversity is structured and maintained. Here, we unravel the patterns and ecological correlates of mammal β‐diversity, including both small (SM) and midsized to large mammal species (LM) across 23 islands and two continuous forest sites within a mega hydroelectric reservoir.

Location

Balbina Hydroelectric Dam, Central Brazilian Amazonia.

Methods

Small mammals were sampled using live and pitfall traps (48,350 trap‐nights), and larger mammals using camera traps (8,160 trap‐nights). β‐diversity was examined for each group using multiplicative diversity decomposition of Hill numbers, which considers the importance of rare, common and dominant species, and tested to what extent those were related to a set of environmental characteristics measured at different spatial scales.

Results

β‐diversity for both mammal groups was higher when considering species presence–absence. When considering species abundance, β‐diversity was significantly higher for SM than for LM assemblages. Habitat variables, such as differences in tree species richness and percentage of old‐growth trees, were strong correlates of β‐diversity for both SMs and LMs. Conversely, β‐diversity was weakly related to patch and landscape characteristics, except for LMs, for which β‐diversity was correlated with differences in island sizes.

Main conclusions

The lower β‐diversity of LMs between smaller islands suggests subtractive homogenization of this group. Although island size plays a major role in structuring mammal α‐diversity in several land‐bridge islands, local vegetation characteristics were additional key factors determining β‐diversity for both mammal groups. Maintaining the integrity of vegetation characteristics and preventing the formation of a large set of small islands within reservoirs should be considered in long‐term management plans in both existing and planned hydropower development in lowland tropical forests.

     

Agriculture erases climate‐driven β‐diversity in Neotropical bird communities

Earth is experiencing multiple global changes that will, together, determine the fate of many species. Yet, how biological communities respond to concurrent stressors at local‐to‐regional scales remains largely unknown. In particular, understanding how local habitat conversion interacts with regional climate change to shape patterns in β‐diversity—differences among sites in their species compositions—is critical to forecast communities in the Anthropocene. Here, we study patterns in bird β‐diversity across land‐use and precipitation gradients in Costa Rica. We mapped forest cover, modeled regional precipitation, and collected data on bird community composition, vegetation structure, and tree diversity across 120 sites on 20 farms to answer three questions. First, do bird communities respond more strongly to changes in land use or climate in northwest Costa Rica? Second, does habitat conversion eliminate β‐diversity across climate gradients? Third, does regional climate control how communities respond to habitat conversion and, if so, how? After correcting for imperfect detection, we found that local land‐use determined community shifts along the climate gradient. In forests, bird communities were distinct between sites that differed in vegetation structure or precipitation. In agriculture, however, vegetation structure was more uniform, contributing to 7%–11% less bird turnover than in forests. In addition, bird responses to agriculture and climate were linked: agricultural communities across the precipitation gradient shared more species with dry than wet forest communities. These findings suggest that habitat conversion and anticipated climate drying will act together to exacerbate biotic homogenization.     

Latitudinal gradient of nestedness and its potential drivers in stream detritivores

Understanding what mechanisms shape the diversity and composition of biological assemblages across broad‐scale gradients is central to ecology. Litter‐consuming detritivorous invertebrates in streams show an unusual diversity gradient, with α‐diversity increasing towards high latitudes but no trend in γ‐diversity. We hypothesized this pattern to be related to shifts in nestedness and several ecological processes shaping their assemblages (dispersal, environmental filtering and competition). We tested this hypothesis, using a global dataset, by examining latitudinal trends in nestedness and several indicators of the above processes along the latitudinal gradient. Our results suggest that strong environmental filtering and low dispersal in the tropics lead to often species‐poor local detritivore assemblages, nested in richer regional assemblages. At higher latitudes, dispersal becomes stronger, disrupting the nested assemblage structure and resulting in local assemblages that are generally more species‐rich and non‐nested subsets of the regional species pools. Our results provide evidence that mechanisms underlying assemblage composition and diversity of stream litter‐consuming detritivores shift across latitudes, and provide an explanation for their unusual pattern of increasing α‐diversity with latitude. When we repeated these analyses for whole invertebrate assemblages of leaf litter and for abundant taxa showing reverse or no diversity gradients we found no latitudinal patterns, suggesting that function‐based rather than taxon‐based analyses of assemblages may help elucidate the mechanisms behind diversity gradients.     

Multi‐scale patterns of moss and lichen richness on the Antarctic Peninsula

Mosses and lichens are the dominant macrophytes of the Antarctic terrestrial ecosystem. Using occurrence data from existing databases and additional published records, we analyzed patterns of moss and lichen species diversity on the Antarctic Peninsula at both a regional scale (1°latitudinal bands) and a local scale (52 and 56 individual snow‐ and ice‐free coastal areas for mosses and lichens, respectively) to test hypothesized relationships between species diversity and environmental factors, and to identify locations whose diversity may be particularly poorly represented by existing collections and online databases. We found significant heterogeneity in sampling frequency, number of records collected, and number of species found among analysis units at the two spatial scales, and estimated species richness using projected species accumulation curves to account for potential biases stemming from sample heterogeneity. Our estimates of moss and lichen richness for the entire Antarctic Peninsula region were within 20% of the total number of known species. Area, latitude, spatial isolation, mean summer temperature, and penguin colony size were considered as potential covariates of estimated species richness. Moss richness was correlated with isolation and latitude at the local scale, while lichen richness was correlated with summer mean temperature and, for 17 sites where penguins where present with <20 000 breeding pairs, penguin colony size. At the regional scale, moss richness was correlated with temperature and latitude. Lichen richness, by contrast, was not significantly correlated with any of the variables considered at the regional scale. With the exception of temperature, which explained 91% of the variation in regional moss diversity, explained variance was very low. Our results show that patterns of moss and lichen biodiversity are highly scale‐dependent and largely unexplained by the biogeographic variables found important in other systems.     

Continental and regional ranges of North American mammals: Rapoport's rule in real and null worlds

Aim To assess the relationship between species richness and distribution within regions arranged along a latitudinal gradient we use the North American mammalian fauna as a study case for testing theoretical models. Location North America. Methods We propose a conceptual framework based on a fully stochastic mid‐domain model to explore geographical patterns of range size and species richness that emerge when the size and position of species ranges along a one‐dimensional latitudinal gradient are randomly generated. We also analyse patterns for the mammal fauna of North America by comparing empirical results from a biogeographical data base with predictions based on randomization null models. Results We confirmed the validity of Rapoport's rule for the mammals of North America by documenting gradients in the size of the continental ranges of species. Additionally, we demonstrated gradients of mean regional range size that parallel those of continental range. Our data also demonstrated that mean range size, measured both as a continental or a regional variable, is significantly correlated with the geographical pattern in species richness. All these patterns deviated sharply from null models. Main conclusions Rapoport's statement of an areographic relationship between species distribution and richness is highly relevant in modern discussions about ecological patterns at the geographical scale.     

Latitudinal patterns of forest ecosystem stability across spatial scales as affected by biodiversity and environmental heterogeneity

Our planet is facing a variety of serious threats from climate change that are unfolding unevenly across the globe. Uncovering the spatial patterns of ecosystem stability is important for predicting the responses of ecological processes and biodiversity patterns to climate change. However, the understanding of the latitudinal pattern of ecosystem stability across scales and of the underlying ecological drivers is still very limited. Accordingly, this study examines the latitudinal patterns of ecosystem stability at the local and regional spatial scale using a natural assembly of forest metacommunities that are distributed over a large temperate forest region, considering a range of potential environmental drivers. We found that the stability of regional communities (regional stability) and asynchronous dynamics among local communities (spatial asynchrony) both decreased with increasing latitude, whereas the stability of local communities (local stability) did not. We tested a series of hypotheses that potentially drive the spatial patterns of ecosystem stability, and found that although the ecological drivers of biodiversity, climatic history, resource conditions, climatic stability, and environmental heterogeneity varied with latitude, latitudinal patterns of ecosystem stability at multiple scales were affected by biodiversity and environmental heterogeneity. In particular, α diversity is positively associated with local stability, while β diversity is positively associated with spatial asynchrony, although both relationships are weak. Our study provides the first evidence that latitudinal patterns of the temporal stability of naturally assembled forest metacommunities across scales are driven by biodiversity and environmental heterogeneity. Our findings suggest that the preservation of plant biodiversity within and between forest communities and the maintenance of heterogeneous landscapes can be crucial to buffer forest ecosystems at higher latitudes from the faster and more intense negative impacts of climate change in the future.     

Biodiversity of soft-sediment benthic communities from Italian transitional waters

Aim For conservation purposes, it is important to understand the forces that shape biodiversity in transitional waters (TWs) and to evaluate the effects of small‐scale latitudinal changes. To this end, we analysed data on soft‐sediment macroinvertebrates from nine Italian TWs in order to (1) investigate the structure and distribution of the benthic fauna and their relationships with environmental and geographical variables, and (2) examine species richness and β‐diversity at various spatial scales. Location European Transition Waters Ecoregion 6. Methods Using a data set collected along a 7° latitudinal cline between 45°28′ N and 39°56′ N, we used Spearman’s rank correlation analysis to evaluate the relationships between species richness and both environmental and geographical variables, and linear regression analysis to show the relationships between α‐, β‐ and γ‐diversity. Three measures were used to assess β‐diversity: Whittaker’s β_W, and two similarity indices, namely the Bray‐Curtis similarity index and Δ_s. Using multivariate analyses, we determined the similarity in composition of the benthic community between sites and compared the biotic ordination with abiotic (geographical and environmental) characteristics. Results Two hundred and sixty‐eight species were recorded from 46 sites. Of these, 53.4% were restricted to one TW. Annelida was the dominant taxonomic group, followed by Crustacea and Mollusca. The α‐diversity was highly variable (5–87 species) and was correlated with latitude. The γ‐diversity, measured at the TW scale, was correlated significantly with α‐diversity. The β‐diversity increased with spatial scale and habitat heterogeneity. In the community pattern identified by multivariate analysis, TWs were segregated by latitude and biogeography, and this reflected different climatic conditions. Main conclusions We found that α‐diversity increased when moving from higher to lower latitudes, and that it depended on both regional and local factors. In addition, we detected latitudinal variations in the extent of regional influence on local species richness. The observed distribution pattern of TW faunas depended mostly on climate type. We suggest that the distribution of annelidan species could be used as a proxy for assessing general community patterns for Italian TWs.     

A phylogenetic perspective on species diversity, β‐diversity and biogeography for the microbial world

There is an increasing interest to combine phylogenetic data with distributional and ecological records to assess how natural communities arrange under an evolutionary perspective. In the microbial world, there is also a need to go beyond the problematic species definition to deeply explore ecological patterns using genetic data. We explored links between evolution/phylogeny and community ecology using bacterial 16S rRNA gene information from a high‐altitude lakes district data set. We described phylogenetic community composition, spatial distribution, and β‐diversity and biogeographical patterns applying evolutionary relatedness without relying on any particular operational taxonomic unit definition. High‐altitude lakes districts usually contain a large mosaic of highly diverse small water bodies and conform a fine biogeographical model of spatially close but environmentally heterogeneous ecosystems. We sampled 18 lakes in the Pyrenees with a selection criteria focused on capturing the maximum environmental variation within the smallest geographical area. The results showed highly diverse communities nonrandomly distributed with phylogenetic β‐diversity patterns mainly shaped by the environment and not by the spatial distance. Community similarity based on both bacterial taxonomic composition and phylogenetic β‐diversity shared similar patterns and was primarily structured by similar environmental drivers. We observed a positive relationship between lake area and phylogenetic diversity with a slope consistent with highly dispersive planktonic organisms. The phylogenetic approach incorporated patterns of common ancestry into bacterial community analysis and emerged as a very convenient analytical tool for direct inter‐ and intrabiome biodiversity comparisons and sorting out microbial habitats with potential application in conservation studies.     

Spatial patterns of association at local and regional scales in coastal sand dune communities

Questions: Are positive understorey‐dominant associations important in physically severe dune communities and does the strength of positive associations vary with disturbance at the local scale and with stress at the regional scale? Do associational patterns observed at the neighbourhood scale predict diversity at higher scales? Location: Coastal sand dunes, Aquitaine (France). Methods: Associational patterns with five dominant species were recorded along a local gradient of disturbance and a 240‐km long regional gradient. Density, richness, cover and variance ratio of understorey species were recorded in quadrats located in dominant and in open areas. Spatial pattern of dominant plant species was recorded using a distance‐based method. Results: Positive understorey‐dominant associations were most frequent at both regional and local scale, although negative associations with understorey species were observed for one of the five dominants. At the regional scale, there was a shift in the magnitude of spatial associations, with higher positive associations in the most stressful sites, whereas spatial associations where not affected by the local disturbance gradient. Positive associations were not related to the size of the dominants but rather influenced by the identity of the dominant species. Conclusions: Our study highlights the potential crucial role of facilitation together with the importance of turnover of the dominants in explaining large‐scale variation in diversity. However, because positive associations may also be attributed to environmental heterogeneity or co‐occurrence of microhabitat preferences of species, experiments are needed to fully assess the relative importance of facilitation versus other drivers of community diversity.     

Applying the dark diversity concept to plants at the European scale

Large‐scale biodiversity maps are essential to macroecology. However, between‐region comparisons can be more useful if patterns of observed species richness are supplemented by variations in dark diversity – the absent portion of the species pool. We aim to quantify and map plant diversity across Europe by using a measure that accounts for both observed and dark diversity. To do this we need to delimit suitable species pools, and evaluate the potential and limitation of a large‐scale dataset. We used Atlas Florae Europaeae (ca 20% of European plant species mapped within 50 × 50 km grid cells) and defined for each grid cell several species pools by applying various geographical and environmental filters: geographic species pool (number of species within 500 km radius), biogeographic species pool (additionally incorporating species distribution patterns, i.e. dispersion fields), site‐specific species pool (additionally integrating environmental preferences of species based on species co‐occurrence). We integrated dark diversity and observed diversity at a relative scale to calculate the completeness of site diversity: logistic expression of observed and dark diversity. We tested whether our results are robust against regional variation in data availability. We used independent regional databases to test if Atlas Florae Europaeae is a representative subset of total species richness. Environmental filtering was the most influential determinant of species pool size with more species filtered out in southern Europe. Both observed and dark diversity adhered to the well‐known latitudinal gradient, but completeness of site diversity varied throughout Europe with no latitudinal trend. Dark diversity patterns were fairly insensitive to variations in regional sampling intensity. Atlas Florae Europaeae represented well the total variation in plant diversity. In summary, dark diversity and completeness of site diversity add valuable information to broad‐scale diversity patterns since observed diversity is expressed at a relative scale.     

Aquatic insect β‐diversity is not dependent on elevation in Southern Rocky Mountain streams

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