On the estimation of species richness based on the accumulation of previously unrecorded species

Estimation of species richness of local communities has become an important topic in community ecology and monitoring. Investigators can seldom enumerate all the species present in the area of interest during sampling sessions. If the location of interest is sampled repeatedly within a short time period, the number of new species recorded is typically largest in the initial sample and decreases as sampling proceeds, but new species may be detected if sampling sessions are added. The question is how to estimate the total number of species. The data collected by sampling the area of interest repeatedly can be used to build species accumulation curves: the cumulative number of species recorded as a function of the number of sampling sessions (which we refer to as “species accumulation data”). A classic approach used to compute total species richness is to fit curves to the data on species accumulation with sampling effort. This approach does not rest on direct estimation of the probability of detecting species during sampling sessions and has no underlying basis regarding the sampling process that gave rise to the data. Here we recommend a probabilistic, nonparametric estimator for species richness for use with species accumulation data. We use estimators of population size that were developed for capture‐recapture data, but that can be used to estimate the size of species assemblages using species accumulation data. Models of detection probability account for the underlying sampling process. They permit variation in detection probability among species. We illustrate this approach using data from the North American Breeding Bird Survey (BBS). We describe other situations where species accumulation data are collected under different designs (e.g., over longer periods of time, or over spatial replicates) and that lend themselves to of use capture‐recapture models for estimating the size of the community of interest. We discuss the assumptions and interpretations corresponding to each situation.     

Inferring species abundance distribution across spatial scales

A long-standing problem in ecology is to understand how the species–abundance distribution (SAD) varies with sampling scale. The problem was first characterized by Preston as the veil line problem. Although theoretical and empirical studies have now shown the nonexistence of the veil line, this problem has generated much interest in scaling biodiversity patterns. However, research on scaling SAD has so far exclusively focused on the relationship between the SAD in a smaller sampling area and a known SAD assumed for a larger area. An unsolved challenge is how one may predict species–abundance distribution in a large area from that of a smaller area. Although upscaling biodiversity patterns (e.g. species–area curve) is a major focus of macroecological research, upscaling of SAD across scale is, with few exceptions, ignored in the literature. Methods that directly predict SAD in a larger area from that of a smaller area have just started being developed. Here we propose a Bayesian method that directly answers this question. Examples using empirical data from tropical forests of Malaysia and Panama are employed to demonstrate the use of the method and to examine its performance with increasing sampling area. The results indicate that only 10-15% of the total census area is needed to adequately predict species abundance distribution of a region. In addition to species abundance distributions, the method also predicts well the regional species richness.     

Species-area relationships from a spatially explicit neutral model in an infinite landscape

We use recently developed technical methods to study species–area relationships from a spatially explicit extension of Hubbell's neutral model on an infinite landscape. Our model includes variable dispersal distances and exhibits qualitatively different behaviour from the cases of nearest-neighbour dispersal and finite periodic landscapes that have previously been studied. We show that different dispersal distances and even different dispersal kernels produce identical species–area curves up to rescaling of the two axes. This scaling property provides a straightforward method for fitting the model to empirical data. The species–area curves display all three phases observed empirically and enable the exponent describing the power law relationship for species–area curves to be identified as the gradient at the central phase. This exponent can take all values between 0 and 1 and is given by a simple function of the speciation rate, independent of all other model variables.     

Abundance and distribution within a guild of benthic stream fishes: local processes and regional patterns

1. Spatial patterns at regional and local scales were examined for evidence that species interactions can influence distribution and abundance within a guild of benthic fishes in upland streams of Oklahoma, U.S.A. Three groups of community patterns were examined: the species–area relationship, species–habitat associations, and interspecific associations.
2. The species–area relationship for riffle habitats was compared to a null species–area model based on random placement. The observed species–area curve was steeper resulting in less species per unit area in small streams than predicted by the null model.
3. Small, species-poor streams had summed fish densities at least as high as larger, species-rich streams, suggesting density compensation.
4. Several significant patterns of negative covariation were found among species at the regional scale, before and after statistically accounting for effects due to measured habitat variables.
5. For two of these negatively covarying taxa ( Cottus carolinae and Etheostoma spectabile ), the influence of each species on the distribution of the other was evaluated experimentally in field enclosures varying in depth and current velocity. The sculpin C . carolinae caused a shift in habitat use by the darter E . spectabile , but no reciprocal shift was found.
6. These results indicate an agreement between local and regional patterns of distribution for C . carolinae and E . spectabile and suggest that biotic interactions can influence regional patterns of distribution for species within this guild.     

Unprotecting the rare species: a niche‐based gap analysis for odonates in a core Cerrado area

Aim We evaluated Odonata distribution data and predicted the compositional resemblance based on niche‐based species distribution models to analyse the following questions: (1) How is estimated species richness distributed, and how can it be preserved under the actual network of conservation units (a gap analysis approach)? (2) How is the estimated odonate beta diversity distributed, and is there a better distribution of conservation units (a priority setting approach)? (3) Is the probability of being under protection a function of the potential species range size? and (4) Will the current conservation network proposals protect odonate taxa? Location Central Brazil in a core Cerrado area. Methods We generated odonate species distribution predictions based on MaxEnt and maps derived from estimated species richness, beta diversity and gap analysis for all species predicted to occur in the study area. Then, we compared these maps with current conservation units, land‐use patterns and proposals for the establishment of conservation units. Results Raw odonate species records provided limited utility for setting conservation priorities without the use of niche‐based models. However, area under the receiver operating curve (AUC) values were characterized by substantial variation that was related to the number of records. No current conservation units overlapped the areas with higher predicted richness and beta diversity, and in general, conservation units were not preserving restricted/rare species. There was a direct linear correlation between species range size and the proportion of its range protected in the current network of conservation units. Finally, we identified three areas with high odonate beta diversity where conservationist actions should be implemented. Main conclusions Current conservation units and future suggested areas do not overlap regions with high conservation values for odonates. Conservation units protect species at random, and the level of protection has a direct relationship with species range size; thus, wide‐range species are expected to be more protected than restricted or threatened species.     

Abundance and distribution within a guild of benthic stream fishes: local processes and regional patterns

1. Spatial patterns at regional and local scales were examined for evidence that species interactions can influence distribution and abundance within a guild of benthic fishes in upland streams of Oklahoma, U.S.A. Three groups of community patterns were examined: the species–area relationship, species–habitat associations, and interspecific associations.
2. The species–area relationship for riffle habitats was compared to a null species–area model based on random placement. The observed species–area curve was steeper resulting in less species per unit area in small streams than predicted by the null model.
3. Small, species-poor streams had summed fish densities at least as high as larger, species-rich streams, suggesting density compensation.
4. Several significant patterns of negative covariation were found among species at the regional scale, before and after statistically accounting for effects due to measured habitat variables.
5. For two of these negatively covarying taxa ( Cottus carolinae and Etheostoma spectabile ), the influence of each species on the distribution of the other was evaluated experimentally in field enclosures varying in depth and current velocity. The sculpin C . carolinae caused a shift in habitat use by the darter E . spectabile , but no reciprocal shift was found.
6. These results indicate an agreement between local and regional patterns of distribution for C . carolinae and E . spectabile and suggest that biotic interactions can influence regional patterns of distribution for species within this guild.     

The effect of habitat on the range expansion of a native and an introduced bird species

Aim Range expansion across a heterogeneous landscape may depend on the habitat selected and used by the expanding species. If habitat selection influences range expansion then localities colonized by a species should contain a greater proportion of favoured habitat (and less non‐habitat) than other nearby localities not colonized. White‐winged doves (Zenaida asiatica) and Eurasian collared doves (Streptopelia decaocto) are two bird species that provide an excellent opportunity to test this hypothesis, because the geographic ranges of both species have been expanding in North America for more than two decades. Location Continental USA. Methods We used distribution data from the North American Breeding Bird Survey to test whether the landscapes occupied by each species contained a greater proportion of favoured habitat (urban land, grassland/pasture, shrub land and cropland) and a lower proportion of non‐habitat (forest land) than landscapes where doves were not found. We tested each species separately in each of three broad expansion areas, namely East, Central and West. We also compared rates of spatial spread between expansion areas and between the two species. Results As predicted, both species tended to occupy landscapes with greater proportions of urban land, shrub land and cropland but with less forest land compared with landscapes without doves, in all three expansion areas. Contrary to prediction, occupied landscapes tended to have slightly less grassland/pasture than unoccupied landscapes. Rates of spread differed between the two species and among expansion areas. Main conclusions Range expansion and the extent to which a species fills or saturates its range are influenced by the habitat ecology of the expanding species. Species colonize localities based on the availability of suitable habitat. However, the role of habitat in a species’ range expansion does depend somewhat on the greater geographical setting. Over large regional and geographical scales, range expansion (rate of spread and saturation) may proceed unevenly, suggesting that range expansion is a very dynamic and context‐specific process.     

Anthropogenic impact on habitat connectivity: A multidimensional human footprint index evaluated in a highly biodiverse landscape of Mexico

Evaluating the cumulative effects of the human footprint on landscape connectivity is crucial for implementing policies for the appropriate management and conservation of landscapes. We present an adjusted multidimensional spatial human footprint index (SHFI) to analyze the effects of landscape transformation on the remnant habitat connectivity for 40 terrestrial mammal species representative of the Trans-Mexican Volcanic System in Michoacán (TMVS_Mich), in western central Mexico. We adjusted the SHFI by adding fragmentation and habitat loss to its original three components: land use intensity, time of human landscape intervention, and biophysical vulnerability. The adjusted SHFI was applied to four scenarios: one grouping all species and three grouping several species by habitat spatial requirements. Using the SHFI as a dispersal resistance surface and applying a circuit theory based approach, we analyzed the effects of cumulative human impact on habitat connectivity in the different scenarios. For evaluating the relationship between habitat loss and connectivity, we applied graph theory-based equivalent connected area (ECA) index. Results show over 60% of the TMVS_Mich has high SHFI values, considerably lowering current flow for all species. Nevertheless, the effect on connectivity of human impact is higher for species with limited dispersal capacity (100–500 m). Our approach provides a new form of evaluating human impact on habitat connectivity that can be applied to different scales and landscapes. Furthermore, the approach is useful for guiding discussions and implementing future biodiversity conservation initiatives that promote landscape connectivity as an adaptive strategy for climate change.     

Land use drivers of species re‐expansion: inferring colonization dynamics in Eurasian otters

Aim Land use intensity has been recognized as one of the major determinants of native species declines. The re‐expansion of species previously constrained by habitat degradation has been rarely investigated. Here, we use site occupancy models incorporating imperfect detection to identify the land use drivers of the re‐expansion of the Eurasian otter (Lutra lutra). Location Czech Republic. Methods We applied multi‐season occupancy models to otter presence–non‐detection data collected in three national surveys (1992, 2000, 2006) at 552 sites (11.2 × 12 km grid cells). Model parameters included site occupancy, colonization and extinction probabilities, and detection probability at a sub‐site level. We modelled changes in occupancy over time as a function of agricultural, urban and industrial land use and change in the extent of agricultural land use. Results Under the best fitting model, occupancy was estimated to be 34.6% in 1992, 51.3% in 2000 and 83.7% in 2006. Detection probability was neither perfect nor constant. Occupancy probability in 1992 was negatively related to land use gradients. Colonization was more likely to occur where a reduction in agricultural land was larger. Variation in extinction and colonization rates along land use gradients resulted in increased occupancy in industrial and especially urban landscapes. Conversely, occupancy remained almost unchanged along agricultural gradients. Main conclusions Dynamics of otter expansion were strongly associated with the two main patterns of the rapid environmental transition that has taken place in the Czech Republic since the early 1990s. Results show that a reduction in intensive agricultural land use led to an increase in otter distribution, providing evidence of the impact of agricultural land use on stream ecosystems. Moreover, otters recolonized urban and industrial landscapes, probably as a result of extensive reduction in water pollution from point sources. Our results suggest that active conservation of otter populations should focus on restoration of freshwater habitat at large scales, especially in agricultural landscapes.     

Assessing biodiversity with species accumulation curves; inventories of small reptiles by pit‐trapping in Western Australia

Abstract We examined 11 non‐linear regression models to determine which of them best fitted curvilinear species accumulation curves based on pit‐trapping data for reptiles in a range of heterogeneous and homogenous sites in mesic, semi‐arid and arid regions of Western Australia. A well‐defined plateau in a species accumulation curve is required for any of the models accurately to estimate species richness. Two different measures of effort (pit‐trapping days and number of individuals caught) were used to determine if the measure of effort influenced the choice of the best model(s). We used species accumulation curves to predict species richness, determined the trapping effort required to catch a nominated percentage (e.g. 95%) of the predicted number of species in an area, and examined the relationship between species accumulation curves with diversity and rarity. Species richness, diversity and the proportion of rare species in a community influenced the shape of species accumulation curves. The Beta‐P model provided the best overall fit (highest r²) for heterogeneous and homogeneous sites. For heterogeneous sites, Hill, Rational, Clench, Exponential and Weibull models were the next best. For homogeneous habitats, Hill, Weibull and Chapman–Richards were the next best models. There was very little difference between Beta‐P and Hill models in fitting the data to accumulation curves, although the Hill model generally over‐estimated species richness. Most models worked equally well for both measures of trapping effort. Because the number of individuals caught was influenced by both pit‐trapping effort and the abundance of individuals, both measures of effort must be considered if species accumulation curves are to be used as a planning tool. Trapping effort to catch a nominated percentage of the total predicted species in homogeneous and heterogeneous habitats varied among sites, but even for only 75% of the predicted number of species it was generally much higher than the typical effort currently being used for terrestrial vertebrate fauna surveys in Australia. It was not possible to provide a general indication of the effort required to predict species richness for a site, or to capture a nominated proportion of species at a site, because species accumulation curves are heavily influenced by the characteristics of particular sites.     

Influence of Land Mosaic Composition and Structure on Patchy Populations: The Case of the Water Vole (Arvicola sapidus) in Mediterranean Farmland

The ability of patchy populations to persist in human-dominated landscapes is often assessed using focal patch approaches, in which the local occurrence or abundance of a species is related to the properties of individual patches and the surrounding landscape context. However, useful additional insights could probably be gained through broader, mosaic-level approaches, whereby whole land mosaics with contrasting patch-network and matrix characteristics are the units of investigation. In this study we addressed this issue, analysing how the southern water vole (Arvicola sapidus) responds to variables describing patch-network and matrix properties within replicated Mediterranean farmland mosaics, across a gradient of agricultural intensification. Patch-network characteristics had a dominant effect, with the total amount of habitat positively influencing both the occurrence of water voles and the proportion of area occupied in land mosaics. The proportions of patches and area occupied by the species were positively influenced by mean patch size, and negatively so by patch isolation. Matrix effects were weak, although there was a tendency for a higher proportion of occupied patches in more intensive, irrigated agricultural landscapes, particularly during the dry season. In terms of conservation, results suggest that water voles may be able to cope well with, or even be favoured by, the on-going expansion of irrigated agriculture in Mediterranean dry-lands, provided that a number of patches of wet herbaceous vegetation are maintained within the farmland mosaic. Overall, our study suggests that the mosaic-level approach may provide a useful framework to understand the responses of patchy populations to land use change.     

Levels of endemism are not necessarily biased by the co-presence of species with different range sizes: a case study of Vilenkin and Chikatunov's models

Aim  To illustrate problems in the methods proposed by B. Vilenkin and V. Chikatunov to study levels of endemism and species–area relationships.
Location  The study used data on the distribution of tenebrionid beetles (Coleoptera, Tenebrionidae) on the Aegean Islands (Greece).
Methods  A total of 32 islands and 170 taxa (species and subspecies) were included in this study. Levels of endemism were evaluated both as the proportion of endemic taxa, and according to the methods proposed by Vilenkin and Chikatunov, which are based on the number of non-endemic taxa and various relationships with area. A model of the species–area relationship proposed by these authors was also analysed.
Results  The number of endemic taxa was positively correlated with the number of taxa with different distribution types, but this positive correlation did not influence the estimation of the level of endemism. In fact, the commonly used estimate of endemicity as a percentage was strongly correlated with the endemism values calculated according to the method of Vilenkin and Chikatunov. The usual power function fitted the species–area relationship as well as the most complicated method of Vilenkin and Chikatunov.
Main conclusions  As hypothesized by Vilenkin and Chikatunov, the number of endemic taxa was influenced both by the number of taxa of other biogeographical ranks, and by an island's area. However, explanations for the positive relationship between the number of endemic taxa and taxa of different biogeographical ranks are equivocal. Importantly, this relationship did not necessarily influence the level of endemism, which could be expressed adequately by percentages. The method proposed by Vilenkin and Chikatunov to estimate the species–area relationship cannot be clearly justified on theoretical grounds and is of questionable practical utility.     

The wealth of species: ecological communities, complex systems and the legacy of Frank Preston

General statistical patterns in community ecology have attracted considerable recent debate. Difficulties in discriminating among mathematical models and the ecological mechanisms underlying them are likely related to a phenomenon first described by Frank Preston. He noted that the frequency distribution of abundances among species was uncannily similar to the Boltzmann distribution of kinetic energies among gas molecules and the Pareto distribution of incomes among wage earners. We provide additional examples to show that four different 'distributions of wealth' (species abundance distributions, species–area and species–time relations, and distance decay of compositional similarity) are not unique to ecology, but have analogues in other physical, geological, economic and cultural systems. Because these appear to be general statistical patterns characteristic of many complex dynamical systems they are likely not generated by uniquely ecological mechanistic processes.     

Hydrological and land use determinants of Eucalyptus camaldulensis occurrence in floodplain wetlands

Hydrological and land use changes can affect species in human altered landscapes. Typically the impacts of hydrological and land use changes are examined separately, with hydrological determinants used to explain the distribution of species in water dependent and aquatic habitats and land use factors used to examine terrestrial species. However, given the connectedness of aquatic and terrestrial habitats, stressors originating in one domain may be important in the other. To explore the importance of integrating both hydrological and land use factors, we tested the relative contribution of hydrological factors and land use context as determinants of the dominant riparian tree species, Eucalyptus camaldulensis Dehn. throughout wetlands of the Condamine catchment, southern Queensland, Australia. The occurrence of E. camaldulensis was modelled against hydrological and land use factors using generalized linear models (GLMs) and validated using internal bootstrapping procedures. Validated models which included both hydrological (distance from weir, wetland–river connectivity and groundwater depth) and land use factors (agricultural land cover and grazing intensity) performed better than those developed using only hydrological factors. The study results highlight the importance of an integrated perspective which considers both hydrological and land use factors in order to understand occurrence patterns of riparian and floodplain tree species in a range of settings. This approach could be especially important when assessing changes to hydrology and land use which may be triggered by climatic changes.     

The 29°N latitudinal line: an important division in the Hengduan Mountains, a biodiversity hotspot in southwest China

This paper aimed to explore the division of the southern and northern Hengduan Mountains based on gradients in species similarity and richness, and to analyze species richness in each sub-region. The Hengduan Mountain region was divided into nine latitudinal belts using one degree of latitude to define the belt after which distribution of seed plants within each latitudinal belt was recorded. Latitudinal patterns of species similarity were measured using the Jaccard similarity index for each pair of adjacent latitudinal belts. Non-metric multidimentional scaling (NMDS) was also used to analyze the similarity in species composition among the nine latitudinal belts. The latitudinal pattern of species similarity and the NMDS ordination both showed a great change in species composition across the 29°N latitudinal line, essentially dividing the Hengduan Mountain region into southern and northern sub-regions. Species richness, shown by the c-value of the species–area power function, and species–area ratio along a latitudinal gradient both showed a sharp decrease across the latitudinal belt from 29°0' to 29°59'N. The southern sub-region occupied 40% of the total area of the Hengduan Mountain region, but contained more than 80% of all the seed plants in the region. The higher species richness and endemism in the southern sub-region showed it to be the core of the Hengduan biodiversity hotspot, a result not unexpected because of the greater extremes of topography and wider diversity of habitats in the southern portion.     

Shapes and functions of species–area curves (II): a review of new models and parameterizations

This paper has extended and updated my earlier list and analysis of candidate models used in theoretical modelling and empirical examination of species–area relationships (SARs). I have also reviewed trivariate models that can be applied to include a second independent variable (in addition to area) and discussed extensively the justifications for fitting curves to SARs and the choice of model. There is also a summary of the characteristics of several new candidate models, especially extended power models, logarithmic models and parameterizations of the negative-exponential family and the logistic family. I have, moreover, examined the characteristics and shapes of trivariate linear, logarithmic and power models, including combination variables and interaction terms. The choice of models according to best fit may conflict with problems of non-normality or heteroscedasticity. The need to compare parameter estimates between data sets should also affect model choice. With few data points and large scatter, models with few parameters are often preferable. With narrow-scale windows, even inflexible models such as the power model and the logarithmic model may produce good fits, whereas with wider-scale windows where inflexible models do not fit well, more flexible models such as the second persistence (P2) model and the cumulative Weibull distribution may be preferable. When extrapolations and expected shapes are important, one should consider models with expected shapes, e.g. the power model for sample area curves and the P2 model for isolate curves. The choice of trivariate models poses special challenges, which one can more effectively evaluate by inspecting graphical plots.     

Reserves in Context: Planning for Leakage from Protected Areas

When protected areas reduce threats within their boundaries, they often displace a portion of these threats into adjacent areas through a process known as ‘leakage’, undermining conservation objectives. Using theoretical models and a case study of terrestrial mammals in Indonesia, we develop the first theoretical explanation of how leakage impacts conservation actions, and highlight conservation strategies that mitigate these impacts. Although leakage is a socio-economic process, we demonstrate that its negative impacts are also affected by the distribution of species, with leakage having larger impacts in landscapes with homogeneous distribution of species richness. Moreover, leakage has a greater negative effect when conservation strategies are implemented opportunistically, even creating the potential for perversely negative consequences from protected area establishment. Leakage thereby increases the relative benefits of systematic conservation planning over opportunism, especially in areas with high leakage and heterogeneously distributed species. Although leakage has the potential to undermine conservation actions, conservation planning can minimize this risk.     

Functional connectivity in replicated urban landscapes in the land snail (Cornu aspersum)

Urban areas are highly fragmented and thereby exert strong constraints on individual dispersal. Despite this, some species manage to persist in urban areas, such as the garden snail, Cornu aspersum, which is common in cityscapes despite its low mobility. Using landscape genetic approaches, we combined study area replication and multiscale analysis to determine how landscape composition, configuration and connectivity influence snail dispersal across urban areas. At the overall landscape scale, areas with a high percentage of roads decreased genetic differentiation between populations. At the population scale, genetic differentiation was positively linked with building surface, the proportion of borders where wooded patches and roads appeared side by side and the proportion of borders combining wooded patches and other impervious areas. Analyses based on pairwise genetic distances validated the isolation‐by‐distance and isolation‐by‐resistance models for this land snail, with an equal fit to least‐cost paths and circuit‐theory‐based models. Each of the 12 landscapes analysed separately yielded specific relations to environmental features, whereas analyses integrating all replicates highlighted general common effects. Our results suggest that urban transport infrastructures facilitate passive snail dispersal. At a local scale, corresponding to active dispersal, unfavourable habitats (wooded and impervious areas) isolate populations. This work upholds the use of replicated landscapes to increase the generalizability of landscape genetics results and shows how multiscale analyses provide insight into scale‐dependent processes.     

Differential responses of cryptic bat species to the urban landscape

Urbanization is a key global driver in the modification of land use and has been linked to population declines even in widespread and relatively common species. Cities comprise a complex assortment of habitat types yet we know relatively little about the effects of their composition and spatial configuration on species distribution. Although many bat species exploit human resources, the majority of species are negatively impacted by urbanization. Here, we use data from the National Bat Monitoring Programme, a long‐running citizen science scheme, to assess how two cryptic European bat species respond to the urban landscape. A total of 124 × 1 km² sites throughout Britain were surveyed. The landscape surrounding each site was mapped and classified into discrete biotope types (e.g., woodland). Generalized linear models were used to assess differences in the response to the urban environment between the two species, and which landscape factors were associated with the distributions of P. pipistrellus and P. pygmaeus. The relative prevalence of P. pygmaeus compared to P. pipistrellus was greater in urban landscapes with a higher density of rivers and lakes, whereas P. pipistrellus was frequently detected in landscapes comprising a high proportion of green space (e.g., parklands). Although P. pipistrellus is thought to be well adapted to the urban landscape, we found a strong negative response to urbanization at a relatively local scale (1 km), whilst P. pygmaeus was detected more regularly in wooded urban landscapes containing freshwater. These results show differential habitat use at a landscape scale of two morphologically similar species, indicating that cryptic species may respond differently to anthropogenic disturbance. Even species considered relatively common and well adapted to the urban landscape may respond negatively to the built environment highlighting the future challenges involved in maintaining biodiversity within an increasingly urbanized world.     

Human impacts on the species-area relationship in reef fish assemblages

The relationship between species richness and area is one of the oldest, most recognized patterns in ecology. Here we provide empirical evidence for strong impacts of fisheries exploitation on the slope of the species–area relationship (SAR). Using comparative field surveys of fish on protected and exploited reefs in three oceans and the Mediterranean Sea, we show that exploitation consistently depresses the slope of the SAR for both power-law and exponential models. The magnitude of change appears to be proportional to fishing intensity. Results are independent of taxonomic resolution and robust across coral and rocky reefs, sampling protocols and statistical methods. Changes in species richness, relative abundance and patch occupancy all appear to contribute to this pattern. We conclude that exploitation pressure impacts the fundamental scaling of biodiversity as well as the species richness and spatial distribution patterns of reef fish. We propose that species–area curves can be sensitive indicators of community-level changes in biodiversity, and may be useful in quantifying the human imprint on reef biodiversity, and potentially elsewhere.