Type and spatial structure of distribution data and the perceived determinants of geographical gradients in ecology: the species richness of African birds

Aim Studies exploring the determinants of geographical gradients in the occurrence of species or their traits obtain data by: (1) overlaying species range maps; (2) mapping survey‐based species counts; or (3) superimposing models of individual species’ distributions. These data types have different spatial characteristics. We investigated whether these differences influence conclusions regarding postulated determinants of species richness patterns. Location Our study examined terrestrial bird diversity patterns in 13 nations of southern and eastern Africa, spanning temperate to tropical climates. Methods Four species richness maps were compiled based on range maps, field‐derived bird atlas data, logistic and autologistic distribution models. Ordinary and spatial regression models served to examine how well each of five hypotheses predicted patterns in each map. These hypotheses propose productivity, temperature, the heat–water balance, habitat heterogeneity and climatic stability as the predominant determinants of species richness. Results The four richness maps portrayed broadly similar geographical patterns but, due to the nature of underlying data types, exhibited marked differences in spatial autocorrelation structure. These differences in spatial structure emerged as important in determining which hypothesis appeared most capable of explaining each map's patterns. This was true even when regressions accounted for spurious effects of spatial autocorrelation. Each richness map, therefore, identified a different hypothesis as the most likely cause of broad‐scale gradients in species diversity. Main conclusions Because the ‘true’ spatial structure of species richness patterns remains elusive, firm conclusions regarding their underlying environmental drivers remain difficult. More broadly, our findings suggest that care should be taken to interpret putative determinants of large‐scale ecological gradients in light of the type and spatial characteristics of the underlying data. Indeed, closer scrutiny of these underlying data — here the distributions of individual species — and their environmental associations may offer important insights into the ultimate causes of observed broad‐scale patterns.     

From richer to poorer: successful invasion by freshwater fishes depends on species richness of donor and recipient basins

Evidence for the theory of biotic resistance is equivocal, with experiments often finding a negative relationship between invasion success and native species richness, and large‐scale comparative studies finding a positive relationship. Biotic resistance derives from local species interactions, yet global and regional studies often analyze data at coarse spatial grains. In addition, differences in competitive environments across regions may confound tests of biotic resistance based solely on native species richness of the invaded community. Using global and regional data sets for fishes in river and stream reaches, we ask two questions: (1) does a negative relationship exist between native and non‐native species richness and (2) do non‐native species originate from higher diversity systems. A negative relationship between native and non‐native species richness in local assemblages was found at the global scale, while regional patterns revealed the opposite trend. At both spatial scales, however, nearly all non‐native species originated from river basins with higher native species richness than the basin of the invaded community. Together, these findings imply that coevolved ecological interactions in species‐rich systems inhibit establishment of generalist non‐native species from less diverse communities. Consideration of both the ecological and evolutionary aspects of community assembly is critical to understanding invasion patterns. Distinct evolutionary histories in different regions strongly influence invasion of intact communities that are relatively unimpacted by human actions, and may explain the conflicting relationship between native and non‐native species richness found at different spatial scales.     

Factors controlling the spatial species richness pattern of four groups of terrestrial vertebrates in an area between two different biogeographic regions in northern Spain

Aim To examine the influence of environmental variables on species richness patterns of amphibians, reptiles, mammals and birds and to assess the general usefulness of regional atlases of fauna. Location Navarra (10,421 km²) is located in the north of the Iberian Peninsula, in a territory shared by Mediterranean and Eurosiberian biogeographic regions. Important ecological patterns, climate, topography and land‐cover vary significantly from north to south. Methods Maps of vertebrate distribution and climatological and environmental data bases were used in a geographic information systems framework. Generalized additive models and partial regression analysis were used as statistical tools to differentiate (A) the purely spatial fraction, (B) the spatially structured environmental fraction and (C) the purely environmental fraction. In this way, we can evaluate the explanatory capacity of each variable, avoiding false correlations and assessing true causality. Final models were obtained through a stepwise procedure. Results Energy‐related features of climate, aridity and land‐cover variables show significant correlation with the species richness of reptiles, mammals and birds. Mammals and birds exhibit a spatial pattern correlated with variables such as aridity index and vegetation land‐cover. However, the high values of the spatially structured environmental fraction B and the low values of the purely environmental fraction A suggest that these predictor variables have a limited causal relationship with species richness for these vertebrate groups. An increment in land‐cover diversity is correlated with an increment of specific richness in reptiles, mammals and birds. No variables were found to be statistically correlated with amphibian species richness. Main conclusions Although aridity and land‐cover are the best predictor variables, their causal relationship with species richness must be considered with caution. Historical factors exhibiting a similar spatial pattern may be considered equally important in explaining the patterns of species richness. Also, land‐cover diversity appears as an important factor for maintaining biological diversity. Partial regression analysis has proved a useful technique in dealing with spatial autocorrelation. These results highlight the usefulness of coarsely sampled data and cartography at regional scales to predict and explain species richness patterns for mammals and birds. The accuracy of models appears to be related to the range perception of each group and the scale of the information.     

Spatiotemporal scaling of plant species richness and functional diversity in a temperate semi‐natural grassland

The accumulation of biodiversity in space and time has been modelled extensively using the species–area relationship and the species–time relationship, respectively. Recently, these models have been combined into time–area curves in order to investigate spatiotemporal scaling of species richness. This study expands on previous research by applying these spatiotemporal models to functional diversity. Understanding spatiotemporal dynamics of ecological traits is important due to their crucial role in ecosystem functioning and mediating species responses to environmental change. We present a new function based on the semi‐logarithmic species–area relationship, which was applied with a power function to vegetation survey data from Scottish machair grassland for both species richness and two measures of functional diversity. When taking a whole‐study approach using non‐linear mixed effects models, the semi‐logarithmic function used here shows a positive time–area interaction for species richness, contrasting with the negative interaction of the power law found in previous investigations. Although there was a negative time–area interaction for functional diversity measures at the whole‐study scale, parameter estimates were inconsistent at the individual site level. Overall, the results reveal differing spatiotemporal dynamics of species and their traits and suggest that the appropriate scale for space‐for‐time substitutions depends on the aspect of biodiversity being investigated. The new model developed in this study, and the novel application to functional diversity, opens up future possible research into spatiotemporal dynamics of biodiversity.     

Elevational gradients and species richness: do methods change pattern perception?

Aim Species richness patterns along elevational gradients have been documented extensively. Yet, the implications of differences in how the data are compiled are seldom explored. We investigate the effect of grain size on the richness–elevation relationship. Grain size varies among the principal methods used to collect or aggregate species occurrences: localized sites, elevational ‘bins’ and interpolation of species ranges. Assumptions of sampling and species distributions also vary among these methods. Methodology can influence the pattern that is perceived and comparability of results. We compare patterns from all three methods explicitly using the same suite of observations, based on museum records and field surveys of non‐flying small mammals. Our assessment is enhanced by comparing patterns resulting from each method for each of six adjacent mountain ranges. Location Utah, North America. Methods We document elevational species richness patterns using generalized linear models (GLMs), comparing the general shape of the trend as well as curvature, location and magnitude of peak richness across methods, both within and among gradients. We also introduce a new procedure to test for richness peaks using site‐based occurrences. Results We find a general congruence of the richness–elevation relationship, depicting a hump‐shaped pattern with a second‐order polynomial GLM showing a significant fit to nearly all gradient‐methodology combinations. However, underlying characteristics of the trend may vary with grain size. As grain size coarsens, maximum species richness increases and elevation of the mode slightly decreases. Results for curvature vary, but degree of curvature tends to increase as grain size coarsens. The richness–elevation patterns are independent of sampling effects. Main conclusions The perceived elevational diversity pattern for small mammals along these mountain ranges is not scale‐dependent. Differences in how the data are compiled are not reflected in major differences in patterns, even when local samples are neither uniformly spaced nor sampled with the same intensity. This result lends confidence to the assertion that patterns documented in similar studies with different methodologies and for which sampling is sufficiently comprehensive are good indicators of diversity. However, consistency of results from more than one compilation method may help to address issues of scale‐dependence, more so when these comparisons are made explicit.     

Niches and neutral processes contribute to the resource–diversity relationships of stream detritivores

1. Explaining resource–diversity relationships is a long‐standing goal in ecology, and there is currently little consensus as to the relative contributions of neutral versus a variety of proposed niche‐related mechanisms. 2. The resource–diversity relationship of insect detritivores was examined in a survey of 25 small, parallel streams flowing into the Bay of Fundy in eastern Canada, with the objective of determining whether neutral processes (sampling effects) could account for the observed patterns. 3. Detritivore taxonomic richness showed a positive, but decelerating relationship with quantity of detritus. Richness also increased with catchment area and with stream permanence. 4. Species distribution patterns were significantly nested, and low resource streams (little detritus) tended to have species with large ranges (i.e. found in many or most streams). 5. Sampling effects could explain only part of the positive relationship between richness and detrital resources, but accounted for the species richness–area relationship. 6. Two mechanisms that could potentially increase niche space as resource abundance increased were rejected: there was no evidence that riparian forest diversity or beta diversity increased with detrital resources. 7. Two niche‐related mechanisms were consistent with existing data, but will require further testing. First, flood disturbance may decrease species richness by eliminating species that require benign habitat, and lowering detritus retention, producing a positive correlation between detritivore richness and resources. Second, large wood in streams located in older riparian forest may increase habitat heterogeneity (number of niches) and the retention of organic matter, again leading to a positive relationship between detritivore diversity and detrital resources. 8. It was concluded that the positive ‘productivity–diversity’ relationship for stream detritivores was most likely produced in part by sampling effects, but also by ecological processes (disturbance and succession) that simultaneously influence resource level and niche availability.     

Ecological realism and mechanisms by which diversity begets stability

We investigated how ecological realism might impact the outcome of three experimental manipulations of species richness to determine whether the patterns and the mechanisms underlying richness–variability relationships differ as ecological communities are increasingly exposed to external forces that may drive richness–variability patterns in nature. To test for such an effect, we conducted experiments using rock pool meio‐invertebrate communities housed in three experimental venues: controlled laboratory microcosms, artificially constructed rock pools in the field, and naturally occurring rock pools in the field. Our results showed that experimental venue can have a strong effect on the outcome of richness manipulation experiments. As ecological realism increased, the strength of the relationship between species richness and community variability declined from 32.9% in the laboratory microcosms to 16.8% in the artificial pools to no effect of species richness on community variability in the natural rock pools. The determinants of community variability also differed as ecological realism increased. In laboratory microcosms, community variability was driven solely by mechanisms related to increasing species richness. In artificial rock pools, community variability was driven by a combination of direct and indirect environmental factors as well as mechanisms related to increasing species richness. In the natural rock pools community variability was independent of species richness and was only related to environmental factors. In summary, we found that stabilizing mechanisms associated with species interactions were influential in establishing species richness–variability relations only in the less realistic experimental venues (the laboratory microcosms and the artificial rock pools in the field), and that these mechanisms diminished in importance as ecological realism and complexity of the experimental venue increased. Our results suggest that the effects of diversity might be more difficult to detect in natural systems due to the combined effects of biotic and abiotic forcing, which can mask our ability to detect richness effects.     

Patterns of diversity, depth range and body size among pelagic fishes along a gradient of depth

Studies of geographical patterns of diversity have focused largely on compiling and analysing data to evaluate alternative hypotheses for the near‐universal decrease in species richness from the equator to the poles. Valuable insights into the mechanisms that promote diversity can come from studies of other patterns, such as variation in species distributions with elevation in terrestrial systems or with depth in marine systems. To obtain such insights, we analysed and interpreted data on species diversity, depth of occurrence and body size of pelagic fishes along an oceanic depth gradient. We used a database on pelagic marine fishes native to the north‐east Pacific Ocean between 40°N and 50°N. We used data from the Pacific Rim Fisheries Program that were obtained from commercial, management and scientific surveys between 1999 and 2000. Depth of occurrence and maximum body length were used to assess the distributions of 409 species of pelagic fishes along a depth gradient from 0 to 8000 m. A presence–absence matrix was used to classify the depth range of each species into 100‐m intervals. Atmar & Patterson's (1995 ) software was used to quantify the degree of nestedness of species distributions. Pelagic fish species diversity decreased steeply with increasing depth; diversity peaked at less than 200 m and more than half of the species had mean depths of occurrence between 0 and 300 m. The distribution of species showed a very strong nested subset pattern along the depth gradient. Whereas species with narrow ranges were generally restricted to shallow waters, wide‐ranging species occurred from near the surface to great depths. The relationship between maximum body size and mean depth range differed between teleost and elasmobranch fishes: being positive for teleosts, but negative for elasmobranches. Results support hypotheses that some combination of high productivity and warm temperature promote high species diversity, and reject those that would attribute the pattern of species richness to the mid‐domain effect, habitat area, or environmental constancy. The data provided a clear example of Rapoport's rule, a negative correlation between average depth range and species diversity.     

A new conceptual and methodological framework for exploring and explaining pattern in presence – absence data

A conceptual framework is proposed to evaluate the relative importance of beta diversity, nestedness and agreement in species richness in presence – absence data matrices via partitioning pairwise gamma diversity into additive components. This is achieved by calculating three complementary indices that measure similarity, relative species replacement, and relative richness difference for all pairs of sites, and by displaying the results in a two‐dimensional simplex diagram, or ternary plot. By summing two terms at a time, three one‐dimensional simplices are derived correspondig to different contrasts: beta diversity versus similarity, species replacement versus nestedness and, finally, richness difference versus richness agreement. The simplex diagrams can be used to interpret underlying data structures by showing departure from randomness towards well‐interpretable directions, as demonstrated by artificial and actual examples. In particular, one may appreciate how far data structure deviates from three extreme model situations: perfect nestedness, anti‐nestedness and perfect gradient. Throughout the paper, we pay special attention to the measurement and interpetation of beta diversity and nestedness for pairs of sites, because these concepts have been in focus of ecological reseach for decades. The novel method can be used in community ecology, conservation biology, and biogeography, whenever the objective is to recover explanatory ecological processes behind patterns conveyed by presence–absence data.     

Orchids of the West Indies: predictability of diversity and endemism

Aim We examined phytogeographical patterns of West Indian orchids, and related island area and maximum elevation with orchid species richness and endemism. We expected strong species–area relationships, but that these would differ between low and montane island groups. In so far as maximum island elevation is a surrogate for habitat diversity, we anticipated a strong relationship with maximum elevation and both species richness and endemism for montane islands. Location The West Indies. Methods Our data included 49 islands and 728 species. Islands were classified as either montane (≥ 300 m elevation) or low (< 300 m). Linear and multivariate regression analyses were run to detect relationships between either area or maximum island elevation and species richness or the number of island endemic species. Results For all 49 islands, the species–area relationship was strong, producing a z‐value of 0.47 (slope of the regression line) and explaining 46% of the variation. For 18 relatively homogeneous, low islands we found a non‐significant slope of z = −0.01 that explained only 0.1% of the variation. The 31 montane islands had a highly significant species–area relationship, with z = 0.49 and accounting for 65% of the variation. Species numbers were also strongly related to maximum island elevation. For all islands < 750 km², we found a small‐island effect, which reduced the species–area relationship to a non‐significant z = 0.16, with only 5% of the variation explained by the model. Species–area relationships for montane islands of at least 750 km² were strong and significant, but maximum elevation was the best predictor of species richness and accounted for 79% of the variation. The frequency of single‐island endemics was high (42%) but nearly all occurred on just nine montane islands (300 species). The taxonomic distribution of endemics was also skewed, suggesting that seed dispersability, while remarkable in some taxa, is very limited in others. Montane island endemics showed strong species–area and species–elevation relationships. Main conclusions Area and elevation are good predictors of orchid species diversity and endemism in the West Indies, but these associations are driven by the extraordinarily strong relationships of large, montane islands. The species richness of low islands showed no significant relationship with either variable. A small‐island effect exists, but the montane islands had a significant relationship between species diversity and maximum elevation. Thus, patterns of Caribbean orchid diversity are dependent on an interplay between area and topographic diversity.     

Using soundscape recordings to estimate bird species abundance, richness, and composition

ABSTRACT Point counts are the most frequently used technique for sampling bird populations and communities, but have well‐known limitations such as inter‐ and intraobserver errors and limited availability of expert field observers. The use of acoustic recordings to survey birds offers solutions to these limitations. We designed a Soundscape Recording System (SRS) that combines a four‐channel, discrete microphone system with a quadraphonic playback system for surveying bird communities. We compared the effectiveness of SRS and point counts for estimating species abundance, richness, and composition of riparian breeding birds in California by comparing data collected simultaneously using both methods. We used the temporal‐removal method to estimate individual bird detection probabilities and species abundances using the program MARK. Akaike's Information Criterion provided strong evidence that detection probabilities differed between the two survey methods and among the 10 most common species. The probability of detecting birds was higher when listening to SRS recordings in the laboratory than during the field survey. Additionally, SRS data demonstrated a better fit to the temporal‐removal model assumptions and yielded more reliable estimates of detection probability and abundance than point‐count data. Our results demonstrate how the perceptual constraints of observers can affect temporal detection patterns during point counts and thus influence abundance estimates derived from time‐of‐detection approaches. We used a closed‐population capture–recapture approach to calculate jackknife estimates of species richness and average species detection probabilities for SRS and point counts using the program CAPTURE. SRS and point counts had similar species richness and detection probabilities. However, the methods differed in the composition of species detected based on Jaccard's similarity index. Most individuals (83%) detected during point counts vocalized at least once during the survey period and were available for detection using a purely acoustic technique, such as SRS. SRS provides an effective method for surveying bird communities, particularly when most species are detected by sound. SRS can eliminate or minimize observer biases, produce permanent records of surveys, and resolve problems associated with the limited availability of expert field observers.     

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.     

Putting insects on the map: near‐global variation in sphingid moth richness along spatial and environmental gradients

Despite their vast diversity and vital ecological role, insects are notoriously underrepresented in biogeography and conservation, and key broad‐scale ecological hypotheses about them remain untested – largely due to generally incomplete and very coarse spatial distribution knowledge. Integrating records from publications, field work and natural history collections, we used a mixture of species distribution models and expert estimates to provide geographic distributions and emergent richness patterns for all ca 1000 sphingid moth species found outside the Americas in high spatial detail. Total sphingid moth richness, the first for a higher insect group to be documented at this scale, shows distinct maxima in the wet tropics of Africa and the Oriental with notable decay toward Australasia. Using multivariate models controlling for spatial autocorrelation, we found that primary productivity is the dominant environmental variable associated with moth richness, while temperature, contrary to our predictions, is an unexpectedly weak predictor. This is in stark contrast to the importance we identify for temperature as a niche variable of individual species. Despite divergent life histories, both main sub‐groups of moths exhibit these relationships. Tribal‐level deconstruction of richness and climatic niche patterns indicate idiosyncratic effects of biogeographic history for some of the less species‐rich tribes, which in some cases exhibit distinct richness peaks away from the tropics. The study confirms, for a diverse insect group, overall richness associations of remarkable similarity to those documented for vertebrates and highlights the significant within‐taxon structure that underpins emergent macroecological patterns. Results do not, however, meet predictions from vertebrate‐derived hypotheses on how thermoregulation affects the strength of temperature–richness effects. Our study thus broadens the taxonomic focus in this data‐deficient discourse. Our procedures of processing incomplete, scattered distribution data are a template for application to other taxa and regions.     

The causes of species richness patterns across space, time, and clades and the role of "ecological limits"

A major goal of research in ecology and evolution is to explain why species richness varies across habitats, regions, and clades. Recent reviews have argued that species richness patterns among regions and clades may be explained by "ecological limits" on diversity over time, which are said to offer an alternative explanation to those invoking speciation and extinction (diversification) and time. Further, it has been proposed that this hypothesis is best supported by failure to find a positive relationship between time (e.g., clade age) and species richness. Here, I critically review the evidence for these claims, and propose how we might better study the ecological and evolutionary origins of species richness patterns. In fact, ecological limits can only influence species richness in clades by influencing speciation and extinction, and so this new "alternative paradigm" is simply one facet of the traditional idea that ecology influences diversification. The only direct evidence for strict ecological limits on richness (i.e., constant diversity over time) is from the fossil record, but many studies cited as supporting this pattern do not, and there is evidence for increasing richness over time. Negative evidence for a relationship between clade age and richness among extant clades is not positive evidence for constant diversity over time, and many recent analyses finding no age-diversity relationship were biased to reach this conclusion. More comprehensive analyses strongly support a positive age-richness relationship. There is abundant evidence that both time and ecological influences on diversification rates are important drivers of both large-scale and small-scale species richness patterns. The major challenge for future studies is to understand the ecological and evolutionary mechanisms underpinning the relationships between time, dispersal, diversification, and species richness patterns.     

Global patterns of specialization and coexistence in bird assemblages

Aim Increased specialization has been hypothesized to facilitate local coexistence and thus high species richness, but empirical evaluations of the richness–specialization relationships have been relatively scant. Here, we provide a first assessment of this relationship for terrestrial bird assemblages at global extent and from fine to coarse grains. Location World‐wide. Methods We use two indices of specialization that describe species‐level resource use: diet and habitat specialization. The relationship between richness and mean assemblage‐level specialization was independently assessed at realm, biome‐realm, 12,100 km² equal‐area grid cells and fine‐grained scales. To identify assemblages that are diverse relative to environmental conditions we: (1) applied quantile regressions, (2) statistically accounted for other environmental variables which may constrain richness, and (3) parsed the data according to the residuals of a model relating species richness to the environmental variables. Results Assemblage species richness increases with both measures of specialization at all scales. Statistically, richness appears constrained by levels of specialization, with the highest richness values only found in specialized assemblages. Richness is positively associated with specialization even after accounting for gradients in resource availability. Net primary productivity and assemblage specialization have complementary statistical effects on assemblage species richness. Contrary to expectations based on niche partitioning of local resources, the relationship between specialization and richness is steep even at coarse scales. Main conclusions The results demonstrate that for an entire clade, totalling > 9000 species, specialization and species richness are related, at least for diverse assemblages. The strong patterns observed across scales suggest that this relationship does not solely originate from (1) limits on coexistence in present‐day assemblages, or (2) increased specialization in richer assemblages imposed by species’ abilities to partition ecological space. Instead, regional‐scale influences on the species pool may determine much of the observed relationship between richness and specialization. Although causal attribution is not straightforward, these findings support the idea that, for the scale of our analysis, specialization may be related to the past origination of high‐diversity assemblages, rather than their contemporary assembly.     

Climatic control of dispersal–ecological specialization trade‐offs: a metacommunity process at the heart of the latitudinal diversity gradient?

We outline the potentially important role of dispersal in linking diversity patterns at different spatial and temporal scales, and the resulting potential to link hypotheses explaining macroscale patterns of diversity. We do this by proposing a possible mechanism linking climate to diversity patterns: we argue that climate, via effects of continuity of habitat availability in space and time, mediates a dispersal–ecological specialization trade‐off at the metacommunity level that leads to latitudinal trends in dispersal ability, ecological specialization, range sizes, speciation and species richness, ultimately driving the latitudinal diversity gradient. This trade‐off constitutes a possible mechanism for the strong macroscale correlation between climate and species richness that is consistent with recent ideas about niche conservatism and gradient lengths, as well as other leading hypotheses. We present an overview of predictions derived from our ideas. Of these, some have already been tested and supported and others are still open to debate or need testing. Together they provide a unique set of predictions that allows falsification.     

Biodiversity–productivity relationship in ponds: Community and metacommunity patterns along time and environmental gradients

Primary production correlates with diversity in various ways. These patterns may result from the interaction of various mechanisms related to the environmental context and the spatial and temporal scale of analysis. However, empirical evidence on diversity‐productivity patterns typically considers single temporal and spatial scales, and does not include the effect of environmental variables. In a metacommunity of macrophytes in ephemeral ponds, we analysed the diversity‐productivity relationship patterns in the field, the importance of the environmental variables of pond size and heterogeneity on such relationship, and the variation of these patterns at local (community level) and landscape scales (metacommunity level) across 52 ponds on twelve occasions, over five years (2005–2009). Combining all sampling dates, there were 377 ponds and 1954 sample‐unit observations. Vegetation biomass was used as a proxy for productivity, and biodiversity was represented by species richness, evenness, and their interaction. Environmental variables comprised pond area, depth and internal heterogeneity. Productivity and species richness were not directly related at the metacommunity level, and were positively related at the community level. Taking environmental variables into account revealed positive species richness‐productivity relationships at the metacommunity level and positive quadratic relationships at the community level. Productivity showed both positive and negative linear and nonlinear relationships with the size and heterogeneity of ponds. We found a weak relationship between productivity and evenness. The identity of variables associated with productivity changed between spatial scales and through time. The pattern of relationships between productivity and diversity depends on spatial scale and environmental context, and changes idiosyncratically through time within the same ecosystem. Thus, the diversity‐productivity relationship is not only a property of the study system, but also a consequence of environmental variations and the temporal and spatial scale of analysis.     

Connecting species richness, abundance and body size in deep-sea gastropods

Aim This paper examines species richness, abundance, and body size in deep‐sea gastropods and how they vary over depth, which is a strong correlate of nutrient input. Previous studies have documented the empirical relationships among these properties in terrestrial and coastal ecosystems, but a full understanding of how these patterns arise has yet to be obtained. Examining the relationships among macroecological variables is a logical progression in deep‐sea ecology, where patterns of body size, diversity, and abundance have been quantified separately but not linked together. Location 196–5042 m depth in the western North Atlantic. Method Individuals analysed represent all Vetigastropoda and Caenogastropoda (Class Gastropoda) with intact shells, excluding Ptenoglossa, collected by the Woods Hole Benthic Sampling Program (3424 individuals representing 80 species). Biovolume was measured for every individual separately (i.e. allowing the same species to occupy multiple size classes) and divided into log₂ body size bins. Analyses were conducted for all gastropods together and separated into orders and depth regions (representing different nutrient inputs). A kernel smoothing technique, Kolmogorov‐Smirnov test of fit, and OLS and RMA were used to characterize the patterns. Results Overall, the relationship between the number of individuals and species is right skewed. There is also a positive linear relationship between the number of individuals and the number of species, which is independent of body size. Variation among these relationships is seen among the three depth regions. At depths inferred to correspond with intermediate nutrient input levels, species are accumulated faster given the number of individuals and shift from a right‐skewed to a log‐normal distribution. Conclusion A strong link between body size, abundance, and species richness appears to be ubiquitous over a variety of taxa and environments, including the deep sea. However, the nature of these relationships is affected by the productivity regime and scale at which they are examined.     

Landscape‐ and small‐scale determinants of grassland species diversity: direct and indirect influences

Species richness is influenced both by mechanisms occurring at landscape scales, such as habitat availability, and local‐scale processes, that are related to abiotic conditions and plant–plant interactions. However, it is rarely tested to what extent local species richness can be explained by the combined effect of factors measured at multiple spatial scales. In this study, we quantified the simultaneous influence of historical landscape‐scale factors (past human population density, and past habitat availability – an index combining area and connectivity) and small‐scale environmental conditions (shrub cover, and heterogeneity of light, soil depth, and other soil environmental variables) on plant species richness in dry calcareous grasslands (alvars). By applying structural equation modelling (SEM) we found that both landscape conditions and local environmental factors had significant direct and indirect (i.e. through the modification of another factor), effects on species richness. At the landscape scale, we found a direct positive influence of historical habitat availability on species richness, and indirect positive influence of past human population (via its effects on historical habitat availability). At small scales, we found a positive direct influence of light heterogeneity and shrub cover on species richness. Conversely, we found that small‐scale soil environmental heterogeneity, which was mainly determined by soil depth heterogeneity, had a negative effect on species richness. Our study indicates that patterns of species richness in alvar grasslands are positively influenced by the anthropogenic management regime that maintained the landscape habitat conditions in the past. However, the abandonment of management, leading to shrub invasion and increased competition for light resources also influenced species richness. In contrast to the positive heterogeneity–diversity relationship we found that soil heterogeneity reduced species richness. Environmental heterogeneity, occurring at the plant neighbourhood scale (i.e. centimetres), can increase the isolation among suitable soil patches and thus hinder the normal functioning of populations. The combination of previous knowledge of the system with new ecological theories facilitates disentangling how species richness responds to complex relationships among factors operating at multiple scales.     

Ecological biogeography of cephalopod molluscs in the Atlantic Ocean: historical and contemporary causes of coastal diversity patterns

Aim One of the most recognized ecological paradigms on earth is the increase in species richness from the poles towards the equator. Here we undertake a comprehensive survey of the latitudinal gradients of species richness (LGSR) of coastal cephalopod fauna in the western (WA) and eastern margins (EA) of the Atlantic Ocean, and test climate and non‐climate theories to explain the variation in diversity. Location The coastal Atlantic Ocean. Methods The diversity and geographical ranges of coastal cephalopods were investigated by means of an exhaustive survey of the primary literature, reports and on‐line data bases. In order to test the productivity, ambient energy and area hypotheses, we investigated the relationship between diversity and net primary production (NPP), sea surface temperature (SST; measure of solar energy input) and continental shelf area, respectively. Results LGSR of cephalopod molluscs are present at both Atlantic coasts, but are quite distinct from each other. Historical processes (rise of the Central American Isthmus, formation of ‘Mare Lago’ and glaciations) explained much of the shape and the zenith of LGSR. Contemporary climate and non‐climate variables also each explained over 83% and 50% of the richness variation in WA and EA, respectively, and the best fitted models accounted for > 92% of the variance. By combining latitude with depth a strong Rapoport effect was observed in WA but not in EA. Main conclusions Besides the evolutionary history, we demonstrate that the contemporary environmental gradients (SST and NPP), shelf area and extent of coral habitat can predict many of the diversity patterns. The longitudinal difference in Rapoport's bathymetric rule is attributed to western fauna specialization to shallow coral reef habitats and greater ecological tolerance of eastern fauna to upwelling ecosystem dynamics. A combined approach of historical biogeography and species–area–energy theories was essential to fully understand broad‐scale variation in cephalopod biodiversity.