Biodiversity sampling is labor intensive, and a substantial fraction of a biota is often represented by species of very low abundance, which typically remain undetected by biodiversity surveys. Statistical methods are widely used to estimate the asymptotic number of species present, including species not yet detected. Additional sampling is required to detect and identify these species, but richness estimators do not indicate how much sampling effort (additional individuals or samples) would be necessary to reach the asymptote of the species accumulation curve. Here we develop the first statistically rigorous nonparametric method for estimating the minimum number of additional individuals, samples, or sampling area required to detect any arbitrary proportion (including 100%) of the estimated asymptotic species richness. The method uses the Chao1 and Chao2 nonparametric estimators of asymptotic richness, which are based on the frequencies of rare species in the original sampling data. To evaluate the performance of the proposed method, we randomly subsampled individuals or quadrats from two large biodiversity inventories (light trap captures of Lepidoptera in Great Britain and censuses of woody plants on Barro Colorado Island [BCI], Panama). The simulation results suggest that the method performs well but is slightly conservative for small sample sizes. Analyses of the BCI results suggest that the method is robust to nonindependence arising from small‐scale spatial aggregation of species occurrences. When the method was applied to seven published biodiversity data sets, the additional sampling effort necessary to capture all the estimated species ranged from 1.05 to 10.67 times the original sample (median ≈ 2.23). Substantially less effort is needed to detect 90% of the species (0.33–1.10 times the original effort; median ≈ 0.80). An Excel spreadsheet tool is provided for calculating necessary sampling effort for either abundance data or replicated incidence data.

The causes of global variation in species richness have been debated for nearly two centuries with no clear resolution in sight. Competing hypotheses have typically been evaluated with correlative models that do not explicitly incorporate the mechanisms responsible for biotic diversity gradients. Here, we employ a fundamentally different approach that uses spatially explicit Monte Carlo models of the placement of cohesive geographical ranges in an environmentally heterogeneous landscape. These models predict species richness of endemic South American birds (2248 species) measured at a continental scale. We demonstrate that the principal single-factor and composite (species-energy, water-energy and temperature-kinetics) models proposed thus far fail to predict (r²⩽0.05) the richness of species with small to moderately large geographical ranges (first three range-size quartiles). These species constitute the bulk of the avifauna and are primary targets for conservation. Climate-driven models performed reasonably well only for species with the largest geographical ranges (fourth quartile) when range cohesion was enforced. Our analyses suggest that present models inadequately explain the extraordinary diversity of avian species in the montane tropics, the most species-rich region on Earth. Our findings imply that correlative climatic models substantially underestimate the importance of historical factors and small-scale niche-driven assembly processes in shaping contemporary species-richness patterns.

The role of intraspecific and interspecific interactions in structuring biotic communities at fine spatial scales is well documented, but the signature of species interactions at coarser spatial scales is unclear. We present evidence that species interactions may be a significant factor in mediating the regional assembly of the Danish avifauna. Because >95% of breeding species ( n = 197) are migratory, we hypothesized that dispersal limitation would not be important and that breeding distributions would largely reflect resource availability and autecological habitat preferences. Instead, we detected a striking pattern of spatial segregation between ecologically similar species at two spatial scales with a suite of null models that factored in the spatial distribution of habitats in Denmark as well as population size and biomass of each species. Habitat utilization analyses indicated that community-wide patterns of spatial segregation could not be attributed to the patchy distribution of habitat or to gross differences in habitat utilization among ecologically similar species. We hypothesize that, when habitat patch size is limited, conspecific attraction in concert with interspecific territoriality may result in spatially segregated distributions of ecologically similar species at larger spatial scales. In the Danish avifauna, the effects of species interactions on community assembly appear pervasive and can be discerned at grain sizes up to four orders of magnitude larger than those of individual territories. These results suggest that species interactions should be incorporated into species distribution modeling algorithms designed to predict species occupancy patterns based on environmental variables.

The search for traits associated with plant invasiveness has yielded contradictory results, in part because most previous studies have failed to recognize that different traits are important at different stages along the introduction–naturalization–invasion continuum. Here we show that across six different habitat types in temperate Central Europe, naturalized non-invasive species are functionally similar to native species occurring in the same habitat type, but invasive species are different as they occupy the edge of the plant functional trait space represented in each habitat. This pattern was driven mainly by the greater average height of invasive species. These results suggest that the primary determinant of successful establishment of alien species in resident plant communities is environmental filtering, which is expressed in similar trait distributions. However, to become invasive, established alien species need to be different enough to occupy novel niche space, i.e. the edge of trait space.