Random placement models predict species–area relationships in duck communities despite species aggregation

Species–area relationships are the product of many ecological processes and their interactions. Explanations for the species–area relationship (SAR) have focused on separating putative niche‐based mechanisms that correlate with area from sampling effects caused by patches with more individuals containing more species than patches with fewer individuals. We tested the hypothesis that SARs in breeding waterfowl communities are caused by sampling effects (i.e. random placement from the regional species pool). First, we described observed SARs and patterns of species associations for fourteen species of ducks on ponds in prairie Canada. Second, we used null models, which randomly allocated ducks to ponds, to test if observed SARs and patterns of species associations differed from those expected by chance. Consistent with the sampling effects hypothesis, observed SARs were accurately predicted by null models in three different years and for diving and dabbling duck guilds. This is the first demonstration that null models can predict SARs in waterbirds or any other aquatic organisms. Observed patterns of species association, however, were not well predicted by null models as in all years there was less observed segregation among species (i.e. more aggregation) than under the random expectation, suggesting that intraspecific competition could play a role in structuring duck communities. Taken together, our results indicate that when emergent properties of ecological communities such as the SAR appear to be caused by random processes, analyses of species associations can be critical in revealing the importance of niche‐based processes (e.g. competition) in structuring ecological communities.     

The theory of the nested species–area relationship: geometric foundations of biodiversity scaling

The relationship between sampled area and the number of species within that area, the species–area relationship (SAR), is a major biodiversity pattern and one of a few law‐like regularities in ecology. While the SAR for isolated units (islands or continents) is assumed to result from the dynamics of species colonization, speciation and extinction, the SAR for contiguous areas in which smaller plots are nested within larger sample areas can be attributed to spatial patterns in the distribution of individuals. The nested SAR is typically triphasic in logarithmic space, so that it increases steeply at smaller scales, decelerates at intermediate scales and increases steeply again at continental scales. I will review current theory for this pattern, showing that all three phases of the SAR can be derived from simple geometric considerations. The increase of species richness with area in logarithmic space is generally determined by overall species rarity, so that the rarer the species are on average, the higher is the local slope z. Rarity is scale‐dependent: species occupy only a minor proportion of area at broad spatial scales, leading to upward accelerating shape of the SAR at continental scales. Similarly, species are represented by only a few individuals at fine spatial scales, leading to high SAR slope also at small areas. Geometric considerations reveal links of the SAR to other macroecological patterns, namely patterns of β‐diversity, the species–abundance distribution, and the relationship between energy availability (or productivity) and species richness. Knowledge of the regularities concerning nested SARs may be used for standardizing unequal areas, upscaling species richness and estimating species loss due to area loss, but all these applications have their limits, which also follow from the geometric considerations.     

Estimates of species extinctions from species–area relationships strongly depend on ecological context

Species–area (SAR) and endemics–area (EAR) relationships are amongst the most common methods used to forecast species loss resulting from habitat loss. One critical, albeit often ignored, limitation of these area‐based estimates is their disregard of the ecological context that shapes species distributions. In this study, we estimate species loss using a spatially explicit mechanistic simulation model to evaluate three important aspects of ecological context: coexistence mechanisms (e.g. species sorting, competition–colonization tradeoffs and neutral dynamics), spatial distribution of environmental conditions, and spatial pattern of habitat loss. We found that 1) area‐based estimates of extinctions are sensitive to coexistence mechanisms as well as to the pattern of environmental heterogeneity; 2) there is a strong interaction between coexistence mechanisms and the pattern of habitat loss; 3) SARs always yield higher estimates of species loss than do EARs; and 4) SARs and EARs consistently underestimate the realized species loss. Our results highlight the need to integrate ecological mechanisms in area‐estimates of species loss.     

Species–area relationships of butterflies in Europe and species richness forecasting

Species–area relationships (SARs) of European butterfly species (Rhopalocera) appear to follow power functions with Mediterranean butterflies having a much higher slope value (z=0.49) compared to the slope for the northern and eastern European countries (z=0.10). A simulated process of species extinction by a stepwise density dependent random elimination of species affected species–area patterns differently. For Mediterranean countries SAR slopes decreased, for other European countries slopes increased during the extinction process. Comparisons of species numbers before and after extinction with those predicted by a classical SAR approach differed widely and revealed that SARs are not able to predict future species numbers at local scales. For Mediterranean countries the classical SAR approach underestimated the number of species remaining after simulated extinction, for all other European countries SARs highly overestimated species numbers. These contrasting patterns indicate that changes in SAR patterns do not unequivocally point to changes in species diversity or community structure as assumed by current theory. On the other hand, the results strongly indicate that simplified applications of SARs for forecasting might give misimpressions about species loss and future biodiversity if the initial community structure, especially relative densities and numbers of species with restricted range size, are not taken into account.     

Habitat structure determines spider diversity in highland ponds

Wetlands (e.g. ponds, meadows) can be found in many landscapes, playing an important role in maintaining regional biodiversity and supporting heterogeneous communities. Spiders are diversified predators that are highly influenced by changes in plant community structure, heterogeneous habitats sustain high spider diversity and abundance. We investigated the characteristics of spider biodiversity in ponds with different habitat structures, by examining patterns across habitats of ponds with different vegetation levels. Sampling took place in four occasions over a year. We compared spider abundance, species richness and composition among ponds including distinct vegetation variables, related to life form, type of leaves, coverage and height. Overall 1174 individuals (194 adults) of 11 families and 37 morphospecies were sampled. We found mostly expected differences in the manner that communities were structured between different habitats. Thus, higher variability of abundance was explained for higher habitat structure of ponds. We also found differences in species composition between ponds with low emergent vegetation and higher habitat structures. Additionaly, spiders were consistently structured more by turnover than nestedness components, with a greater beta diversity of web-builders. Our results suggest varying levels of habitat structures and species substitution shape pond spider communities, depending on habitat heterogeneity and spider guild. Those findings demonstrate the clear role of spatial habitat structure, with more spider species preferring to build webs or actively hunt at vegetated environments on ponds.     

The imprint of the geographical, evolutionary and ecological context on species-area relationships

Species-area relationships (SAR) are fundamental in the understanding of biodiversity patterns and of critical importance for predicting species extinction risk worldwide. Despite the enormous attention given to SAR in the form of many individual analyses, little attempt has been made to synthesize these studies. We conducted a quantitative meta-analysis of 794 SAR, comprising a wide span of organisms, habitats and locations. We identified factors reflecting both pattern-based and dynamic approaches to SAR and tested whether these factors leave significant imprints on the slope and strength of SAR. Our analysis revealed that SAR are significantly affected by variables characterizing the sampling scheme, the spatial scale, and the types of organisms or habitats involved. We found that steeper SAR are generated at lower latitudes and by larger organisms. SAR varied significantly between nested and independent sampling schemes and between major ecosystem types, but not generally between the terrestrial and the aquatic realm. Both the fit and the slope of the SAR were scale-dependent. We conclude that factors dynamically regulating species richness at different spatial scales strongly affect the shape of SAR. We highlight important consequences of this systematic variation in SAR for ecological theory, conservation management and extinction risk predictions.     

How to allow SAR collapse across local and continental scales: a resolution of the controversy between Storch et al. (2012) and Lazarina et al. (2013)

Up‐scaling species richness from local to continental scales is an unsolved problem of macroecology. Macroecologists hope that proper up‐scaling can uncover the hidden rules that underlie spatial patterns in species richness, but a machinery to up‐scale species richness also has a purely practical side at the scales and for the habitats where direct observations cannot be performed. The species–area relationship (SAR) could provide a tool for up‐scaling, but no valid method has yet been put forward. Such a method would have resulted from Storch et al.'s (2012) suggestion that there is a universal curve to which each rescaled SAR collapses, if Lazarina et al. (2013) had not shown that it does not: both arguments were supported by data analyses. Here we present an analytical model for mainland SAR and argue in favour of the latter authors. We identify 1) the variation in mean species‐range size, 2) the variation in forces that drive SAR at various scales, and 3) the finite‐area effect, as the reasons for the absence of collapse. Finally, we suggest a rescaling that might fix the problem. We conclude, however, that ecologists are still far from finding a practical, robust and easy‐to‐use solution for up‐scaling species richness from SARs.     

Agriculture rivals biomes in predicting global species richness

Species–area relationships (SARs) provide an avenue to model patterns of species richness and have recently been shown to vary substantially across regions of different climate, vegetation, and land cover. Given that a large proportion of the globe has been converted to agriculture, and considering the large variety in agricultural management practices, a key question is whether global SARs vary across gradients of agricultural intensity. We developed SARs for mammals that account for geographic variation in biomes, land cover and a range of land‐use intensity indicators representing inputs (e.g. fertilizer, irrigation), outputs (e.g. yields) and system‐level measures of intensity (e.g. human appropriation of net primary productivity – HANPP). We systematically compared the resulting SARs in terms of their predictive ability. Our global SAR with a universal slope was significantly improved by the inclusion of any one of the three variable types: biomes, land cover, and land‐use intensity. The latter, in the form of human appropriation of net primary productivity (HANPP), performed as well as biomes and land‐cover in predicting species richness. Other land‐use intensity indicators had a lower predictive ability. Our main finding that land‐use intensity performs as well as biomes and land cover in predicting species richness emphasizes that human factors are on a par with environmental factors in predicting global patterns of biodiversity. While our broad‐scale study cannot establish causality, human activity is known to drive species richness at a local scale, and our findings suggest that this may hold true at a global scale. The ability of land‐use intensity to explain variation in SARs at a global scale had not previously been assessed. Our study suggests that the inclusion of land‐use intensity in SAR models allows us to better predict and understand species richness patterns.     

Maladaptation and mass effects in a metacommunity: consequences for species coexistence

Metacommunity theories predict multispecies coexistence based on the interplay between local species interactions and regional migration. To date, most metacommunity models implicitly assume that evolution can be ignored. Yet empirical studies indicate a substantial potential for contemporary evolution. I evaluate how evolution alters species diversity in a simulated mass-effects (sink-source) metacommunity. Populations inhabiting source habitats became locally adapted, while subordinate competitors became maladapted because of assumed ecological and phenotypic trade-offs between habitats. This maladaptation decreased and leveled relative abundances among subordinate populations. These two effects produced two regions of departure from nonevolutionary predictions. Assuming low proportional migration, maladaptation reduced local species richness via an overall reduction in reproductive rates in sink populations. With intermediate proportional migration, a greater absolute reduction of reproductive rates in intermediate competitors leveled reproductive rates and thereby enhanced local species richness. Although maladaptation is usually viewed as a constraint on species coexistence, simulations suggest that its effects on diversity are manifold and dependent on interpatch migration and community context. Hence, metacommunity predictions often may profit from an evolutionary perspective. Results indicate that modifications of community connectivity, such as might occur during habitat fragmentation, could elicit rapid shifts in communities from regions of high to low biodiversity.     

A test of the mechanisms behind avian generalized individuals–area relationships

Aim Questions related to abundances of organisms are central to ecological research. A priori, a scale independent estimation of abundances would be expected. However, we find estimates of numbers of bird individuals from all over the world to increase less than proportionately with increasing plot size. At the whole assemblage level, the pattern holds across biogeographical regions and habitats. The slope of the interspecific and, for the majority of species, the intraspecific individuals–area relationship is also significantly shallower than 1. The question arises as to which mechanisms cause these patterns. Location Global. Methods At the assemblage, interspecific and intraspecific levels, we tested three mechanisms that could be responsible for these patterns by comparing the slope of the individuals–plot area relationship for subsets of a database compiled from the literature. Spatial autocorrelation was controlled for. Results There was no evidence for an influence of plot area choice in order to sample a constant number of individuals. Evidence for higher survey efficiency was available only with increasing number of visits at the intraspecific level. Evidence for influences of habitat heterogeneity was present at the assemblage, interspecific and intraspecific levels. This mechanism can work only if small plots are delimited non‐randomly in homogeneous habitat. Main conclusions Avian population size estimates without indication of the area over which they were obtained are of substantially less value than those coupled with that information. Ecologists planning to compare avian abundances between plots varying in some other factor of interest should minimize variations in their areas and/or account for them in data analyses. Population viability analyses, regional and global population size estimates, site prioritization and the scaling of ecosystem and species energy utilization need to address the plot area effect on assemblage and individual species abundances.     

Artificial ponds increase local dragonfly diversity in a global biodiversity hotspot

Human demands have led to an increased number of artificial ponds for irrigation of crops year-round. Certain insect species have established in these ponds, including dragonflies (Insecta: Odonata). There has been discussion around the value of artificial ponds for encouraging dragonfly diversity, with little work in biodiversity hotspots rich in rare and endemic species. We focus here on the Cape Floristic Region (CFR) global biodiversity hotspot, which has many endemic dragonfly species but has few natural ponds. Yet it has many artificial ponds mostly used for irrigation on local farms. This leads to an interesting question: to what extent do these artificial ponds provide habitats for dragonflies in this biologically rich, agriculturally fragmented landscape? To answer this, we recorded dragonfly species richness and abundances from 17 artificial ponds and 13 natural stream deposition pools as reference, in an area of the CFR where there are no local, natural, perennial ponds. Thirteen environmental and physical variables were recorded at the ponds and pools. We found that although ponds attracted no rare or threatened dragonfly species, they increased the area of occupancy and population sizes of many generalist species. These came from nearby natural deposition pools or from unknown sources elsewhere in the region, so providing refuges which otherwise would not be there. Interestingly, some CFR endemic species were also recorded at our artificial ponds. Overall dragonfly assemblages and those of true dragonflies (Anisoptera) and damselflies (Zygoptera) differed between artificial ponds and deposition pools, suggesting that artificial ponds are to some extent a novel ecosystem. Habitat type, elevation and temperature were significant drivers in structuring overall species assemblages. For the Anisoptera, riparian vegetation and level of landscape connectivity was important, while temperature was not. In contrast, Zygoptera species were most affected by river catchment, habitat type and temperature. In sum, these artificial ponds are stepping stone habitats across an increasingly fragmented landscape. Managing these ponds with perennial water, constant water levels, and maximum complexity and heterogeneity of habitats in terms of vegetation will conserve a wide range of generalists and some specialists.     

Species–area models to assess biodiversity change in multi-habitat landscapes: The importance of species habitat affinity

Species–area relationships (SARs) are a common tool to assess the impacts of habitat loss on species diversity. Species–area models that include habitat effects may better describe biodiversity patterns; also the shape of the SAR may be best described by other models than the classical power model. We compared the fit of 24 SAR models, i.e. eight basic models using three approaches: (i) single-habitat models, (ii) multi-habitat models which account for the effect of the habitat composition on total species diversity (= choros models) and (iii) multi-habitat models which also account for the differential use of habitats by different species groups (= countryside models). We use plant diversity data from a multi-habitat landscape in NW Portugal. Countryside models had the best fit both when predicting species–area patterns of species groups and of total species richness. Overall, choros models had a better fit than single-habitat models. We also tested the application of multi-habitat models to land-use change scenarios. Estimates of species richness using the choros model only depended on the number of habitats in the landscape. In contrast, for the countryside model, estimates of species richness varied continuously with the relative proportion of the different habitat types in the landscape, and projections suggest that land-use change impacts may be moderated by a species’ ability to use multiple habitats in the landscape. We argue that the countryside SAR is a better model to assess the impacts of land-use changes than the single-habitat SAR or the choros model, as species often face habitat change instead of real habitat loss, and species response to change is contingent on their differential use of habitats in the landscape.     

Global macroecology of bird assemblages in urbanized and semi‐natural ecosystems

Aim Despite the increasing pace of urbanization, little is known about how this process affects biodiversity globally. We investigate macroecological patterns of bird assemblages in urbanized areas relative to semi‐natural ecosystems. Location World‐wide. Methods We use a database of quantitative bird surveys to compare key assemblage structure parameters for plots in urbanized and semi‐natural ecosystems controlling for spatial autocorrelation and survey methodology. We use the term ‘urbanized’ instead of ‘urban’ ecosystems as many of the plots were not located in the centre of towns but in remnant habitat patches within conurbations. Results Some macroecological relationships were conserved in urbanized landscapes. Species–area, species–abundance and species–biomass relationships did not differ significantly between urbanized and non‐urbanized environments. However, there were differences in the relationships between productivity and assemblage structure. In forests, species richness increased with productivity; in both forests and open habitats, the evenness of species abundances declined as productivity increased. Among urbanized plots, instead, both species richness and the evenness of species abundances were independent of variation in productivity. Main conclusions Remnant habitats within urbanized areas are subject to many ecological alterations, yet key macroecological patterns differ remarkably little in urbanized versus non‐urbanized plots. Our results support the need for increased conservation activities in urbanized landscapes, particularly given the additional benefits of local experiences of biodiversity for the human population. With increasing urbanization world‐wide, broad‐scale efforts are needed to understand and manage the effects of this driver of change on biodiversity.     

Contrasting alien and native plant species–area relationships: the importance of spatial grain and extent

Aim The proportion of alien plant species in floras is increasingly being used to indicate the threat of invasions to native species and/or the homogenization of biodiversity. However, this indicator is only valuable if it is independent of the spatial extent and grain of observation. This study tested the equivalence of native and alien species–area relationships (SARs) in order to assess the support for scale invariance in the proportion of alien species in floras. Location England, UK. Methods Nested SARs were generated by assessing the richness of native and alien plant species drawn from the New atlas of the British and Irish flora for six areas comprising 100, 400, 900, 1600, 2500 and 3600 km² with each larger area containing all smaller areas. Five replicate sets of nested areas encompassing northern, southern, eastern, western and central regions were chosen. For each set of nested areas, the log‐transformed species richness was regressed on log‐transformed area to fit a power function to the SAR. Results Native and alien plant SARs reveal consistent differences in slope, highlighting that the proportion of alien species is a function of spatial grain. Aliens are more rare than natives and have higher spatial turnover leading to faster accumulation of species as area increases. However, equivalent samples drawn from a larger spatial extent reveal similar alien and native SARs. Main conclusions The significant differential scale dependence in native and alien species richness observed in this study reflects dissimilar influences of regional drivers such as habitat, but potentially also propagule pressure and introduction history, that leads to the relative rarity and high spatial turnover of alien species. Maps of invasion hotspots that identify areas where the proportion of the alien flora is particularly high should therefore be treated with considerable caution since patterns across most grains used for species monitoring will be scale dependent.     

Effects of community structure on the species–area relationship in China's forests

The species–area relationship (SAR) is the oldest and most frequently documented law in ecology. In a community, the SAR is regulated by the abiotic environment and biotic interactions and depends on the individual–spatial distribution of species (ISD) and the species–abundance distribution (SAD). In this study, we explored the effects of aggregation of ISDs and unevenness of SADs on SARs in forests of China by comparing the empirical and simulated SARs of 32 nested plots distributed along an extensive latitudinal gradient. Both aggregation and unevenness affected the shape of SARs significantly: ISDs accounted for 12.6 ± 4.0% of the incremental increase in species richness with area, and SADs accounted for 18.7 ± 3.8 and 23.5 ± 3.9% under the broken‐stick model and even abundance model, respectively. Effects of both aggregation and unevenness decreased as temperature increased, suggesting that individuals of a species were spatially more aggregated than random, and the individuals among species were more discrepant from the null distribution (broken‐stick model and even abundance model in this study), in the cold than in the warm areas. Taken together, our results demonstrate that ISDs and SADs within communities can shape SARs, but these effects vary along latitudinal gradients, and are likely mediated by temperature.     

Ecosystem services across the aquatic–terrestrial boundary: Linking ponds to pollination

Many small farmland ponds are built for nutrient retention, the conservation of biodiversity or both, yet they are relatively neglected habitats. For example, little is known about the potential for ponds to influence populations of beneficial terrestrial insects, deliver ecosystem services across the aquatic–terrestrial boundary and affect crop yield in insect-pollinated cash crops.We assessed whether the presence of a pond affects the abundance of pollinators and the quality and quantity of strawberry yield. We compared the abundance of pollinators and the quality and quantity of strawberries between habitats adjacent to the pond, semi-natural terrestrial habitat and field border without semi-natural vegetation (control habitat).We found significantly higher abundances of syrphids and bees next to ponds compared to control habitats. Also, syrphids were significantly more abundant at pond habitats compared to vegetation habitats and a similar tendency, although not significant, was found for the abundance of bees. The quantity and quality of strawberries was significantly higher near the vegetation and pond habitats compared to the control habitats.Our result supports the theory that the presence of semi-natural habitats, in the agricultural landscape benefits both public interest in biodiversity conservation and farmers’ interest in crop pollination. These benefits may also come from ponds as semi natural habitats. However, further studies are required to disentangle the effect of the pond per se and the effect of the associated terrestrial vegetation.     

Fishing‐down within populations harms seed dispersal mutualism

Large fish are often the most effective seed dispersers, but they are also the preferred target for fisheries. We recently started to comprehend the detrimental impacts of the extirpation of large frugivorous fish species on natural forest regeneration, but we lack a general understanding of how intraspecific size‐selective harvest affects fish–fruit mutualism. Our literature review demonstrated that large individuals within populations positively affect diverse aspects of seed dispersal, from consuming a higher diversity of seeds to enhancing germination. Furthermore, we filled a research gap by studying how individual size variations within two small frugivorous fish species (<16 cm) affect seed dispersal in flooded savannas. Even within small‐bodied species, large individuals swallow a higher number of intact seeds, but not necessarily a higher proportion. Overall, our results demonstrate the disproportional role of large‐bodied individuals as key seed dispersers in flooded habitats. Consequently, fishing‐down within both large‐ and small‐bodied species can negatively affect seed dispersal and natural regeneration in overfished wetlands.     

Species–area and species–sampling effort relationships: disentangling the effects

Species numbers tend to increase with both the area surveyed (species–area relationship, SAR) and the number of samples taken (species–sampling effort relationship, SSER). These two relationships differ in their nature and underlying mechanisms but are not clearly distinguished in field studies. To discriminate the effects of area (spatial extent) and sampling effort (SE) on species richness, several models explicitly involving both variables were proposed and tested against 13 datasets from marine micro‐, meio‐ and macrobenthos. A combination of power SSER and piecewise power SAR terms was found to have the best fit. The effects of area and SE were both significant, but the former one was noticeably weaker. The SSERs were roughly linear in log‐log space, whereas the SARs demonstrated scale‐dependent behavior with a noticeable threshold (slope breakpoint). Species richness was almost area‐independent below this threshold (the “small area effect”, SAE) but followed typical power‐law SAR beyond the threshold. This effect was similar to the “small island effect” but occurred for arbitrarily delineated areas within continuous habitats. Parameters of the SAR curves depended on organism size. The upper limit of the SAE increased from microorganisms to meiofauna to macrofauna. Also, SAR curves for unicellular groups had significantly lower slopes. SAE is supposed to indicate a spatial range of statistical homogeneity in species composition. Its upper limit corresponds to the characteristic size of a local community (a single habitat occupied by a common species pool). Interpretations of SAR and SSER parameters in terms of α‐ and β‐diversity are proposed. Both SAR and SSER slopes obtained from univariate regressions are overestimated. This upward bias depends on sampling design, decreasing for SAR but increasing for SSER with more unequally spaced samples. Both spatial extent and sampling effort should be taken into account to disentangle properly their effects on diversity.     

Spatial patterns in species–area relationships and species distribution in a West African forest–savanna mosaic

Aim To investigate the relationship between the slope z of the species–area relationship (SAR) and the intensity of spatial patterns in species number and dissimilarity for woody plants with different modes of seed dispersal. According to island theory we expect, for any given archipelago, steeper slopes and more pronounced spatial patterns for groups of less dispersive species. Location Ivory Coast, West Africa. Methods In a West African forest–savanna mosaic we collected presence–absence data for woody plant species in 49 forest islands. The parameters of the SARs were fitted by nonlinear regressions and then compared for plant species aggregated according to their mode of seed dispersal. We used the Mantel test to calculate the intensity of spatial patterns in species number, i.e. residual deviation from SAR, and species dissimilarity. Results The z‐value for bird‐dispersed species was lower (0.11) than that for wind‐dispersed species (0.27), with mammal‐dispersed species taking an intermediate value (0.16). This result suggests that, as a group, bird‐dispersed species are better colonizers. The spatial pattern in species number as well as species similarity was more pronounced for bird‐ compared with wind‐dispersed species. Main conclusions The standard interpretation of the theory of island biogeography claims that shallow slopes in the SAR imply low isolation of islands, i.e. good dispersal abilities of species. The results of our study appear to contradict this statement. The contradiction can eventually be resolved by a more detailed account of the colonization process, i.e. by distinguishing between dispersal and consecutive establishment of populations.