Environmental correlates of body size distributions of European springtails (Hexapoda: Collembola)

Aim Species–body size distributions (SBDs) are plots of species richness across body size classes. They have been linked to energetic constraints, speciation–extinction dynamics and to evolutionary trends. However, little is known about the spatial variation of size distributions. Here we study SBDs of European springtails (Collembola) at a continental scale and test whether minimum, average and maximum body size and the shapes of size distributions change across latitudinal and longitudinal gradients and whether SBDs of islands and mainlands differ. We also test whether the island rule and the positive body size–range size relationship of vertebrates also holds for Collembola. Location Europe. Methods We use a unique data set on the spatial distributions of 2102 species of European springtails across 52 countries and larger islands together with associated data on body size, area, climate variables, longitude and latitude. Differences in the central moments of SBDs are inferred from simultaneous spatial autoregression models. Results The SBD of the European Collembola and its largest suborder Entomobryomorpha is unimodal and symmetrical. Average, minimum and maximum body weight and the skewness of the mainland/island SBDs peaked at intermediate latitudes. We could not find simple latitudinal gradients in minimum and maximum body weight. Average and maximum body size increased with country/island area in accordance with the island rule in vertebrates, while minimum body size did not significantly differ between islands and mainlands. Finally, we found a weak but statistically significant positive correlation of range size and body size. Main conclusions We provide evidence for differences in body size distributions between islands and mainlands that are in part in line with the island rule in invertebrates. We also find evidence for an interspecific body size–range size relationship similar to that of vertebrates although the vertebrate pattern is much stronger than the springtail pattern. Our results on latitudinal gradients of maximum and average body size imply the need to account for species richness and area effects in the study of latitudinal gradients in body size. We recommend implementing sample size and area effects in the study of body size distributions on islands and mainlands.     

Area effect of distribution of Silphids (Coleoptera,Silphidae) on Korean islands

We investigated the spatial patterns in the distribution of carrion beetles on 26 islands to test the area and distance effects on species diversity. The relationships between species richness and assemblage and island area, distance from the mainland, and shortest distance from the largest island within 2?km were analyzed using linear regression, Canonical Correspondence Analysis (CCA) and Multi-Response Permutation Procedures (MRPP). We hypothesized that both area and distance from the mainland played a significant role in species richness, while geographic location in the sea influenced aggregation of carrion beetles on islands. Area was the only factor influencing species richness of carrion beetles on islands. Moreover, distance from the mainland and close islands did not significantly influence to carrion beetles. In addition, geographic location in the same Sea did not influence beetle assemblages. Taken together, the results of this study indicated that carrion beetles on Korean islands were largely affected by island size and that dispersal was stochastic.     

Spatial patterns in the distribution of European springtails (Hexapoda: Collembola)

Using a large database on the spatial distribution of European springtails (Collembola) we investigated how range sizes and range distribution across European countries and major islands vary. Irrespective of ecological guild, islands tended to contain more endemic species than mainland countries. Nestedness and species co‐occurrence analysis based on country species lists revealed latitudinal and longitudinal gradients of species occurrences across Europe. Species range sizes were much more coherent and had fewer isolated occurrences than expected from a null model based on random colonization. We did not detect clear postglacial colonization trajectories that shaped the faunal composition across Europe. Our results are consistent with a multiregional postglacial colonization. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 498–506.     

Effects of plant diversity on Collembola in an experimental grassland ecosystem

The response of species numbers and density of Collembola to manipulation of plant species richness (1, 2, 4, 8, 32 species) and number of plant functional groups (grasses, legumes and non-legume herbs) was studied in an experimental grassland at the Swiss BIODEPTH site (Lupsingen, Switzerland) in October 1997. Plant species richness or number of plant functional groups did not affect total diversity of Collembola, however, the number of Collembola species increased in the presence of legumes and the grass Trisetum flavescens . The abundance of Protaphorura armata increased but that of Hypogastruridae/Neanuridae significantly decreased with increasing number of plant functional groups. Other groups including the herbivorous Symphypleona did not respond to plant species richness and plant functional groups. Possibly, Hypogastruridae/Neanuridae species are weak competitors declining in density if the density of other Collembola groups increase. In general, the effect of the number of plant functional groups on the densities of collembolan taxa was stronger than that of plant species richness. Changes in Collembola density and diversity in part was likely caused by increased soil microbial and fine root biomass in treatments with higher plant functional group diversity. The presence of legumes resulted in an increase in the densities of total Collembola, Symphypleona/Neelipleona and Isotomidae indicating that they benefited from the high litter quality and the increased microbial biomass in the rhizosphere of legumes. The results suggest that microbivorous soil invertebrates are controlled by food quality rather than quantity. Furthermore, they indicate that presence of certain plant species and functional groups may be more important for collembolan community structure than the diversity of plant species and functional groups per se.     

Plant species diversity and polyploidy in islands of natural vegetation isolated in extensive cultivated lands

Natural polyploidy is often related to a longer life span, vegetative reproduction and higher competitive ability. In this paper, we test the possibility that these characteristics may favour the survival of polyploid taxa under conditions of long-term habitat fragmentation. In islands of natural vegetation isolated in extensive vineyards located in the South of France and in a large neighbouring area of natural vegetation, plant species richness and the relative abundance of polyploid taxa were assessed according to island size, isolation and vegetation structure. High species richness was observed, with numerous species restricted to the islands, suggesting that these may constitute refugia. However, species richness was not related to island size or to degree of isolation except for the flora of the woody areas. A very positive effect of area fragmentation on plant richness was observed, which is probably attributable to relatively low species overlap among the islands. Particularly high species richness was observed in open areas, provided that these were not extensively colonized by shrubs which seem to be responsible for local extinction of many annual taxa. Polyploids, which comprised mostly perennial herbs and woody species, were predominant in all the islands and in the large reference area. In open habitats invaded by shrubby species, a higher relative frequency of polyploids was observed in islands than in the reference area. Moreover, polyploid taxa were present in a larger number of islands than the diploid taxa, which were often restricted to a single island, suggesting that, after a long period of isolation, the polyploids may still have a lower probability of extinction. Evidence was obtained from vegetation structure analysis that diploid and polyploid annual herbs were restricted to open habitats and were both eliminated by shrubby species. Conversely, the diploid perennial herbs were also significantly affected by shrub colonization whereas the polyploids were mostly present in shrubby areas. This suggests that the higher competitive ability of polyploid perennial herbs may constitute a critical factor responsible for their wider distribution over the islands. We report the implications of our findings on conservation strategies, more particularly for a Mediterranean flora.     

Collembola as bioindicators of restoration in mined sand dunes of Northeastern Brazil

Opencast mining causes severe environmental impacts by removing the vegetation cover and depleting the fauna. Reforestation methods using native species and diverse pre- and post-disturbance approaches aim to recover the original richness and diversity of species found before the impact. Bioindicators are powerful tools to evaluate the restoration of the original environmental conditions in disturbed areas. We used species richness, endemism and diversity measurements of Collembola to compare successional stages in reforested sites of different ages compared with a control undisturbed area. Richness and abundance of Collembola were subjected to correlation analysis with age of plots and vegetational variables. Areas that were reforested for up to 16 years supported a much lower Collembola species richness than undisturbed areas. Both the age of reforestation plots and vegetation variables (number of trees, diameter of crowns, depth of leaf litter and tree species richness) were positively and significantly correlated to collembolan abundance and richness. The results showed that the diversity of the 16-year-old plot was significantly higher than that of younger areas, but significantly less diverse than that of the control area. Endemic species were more sensitive to disturbance than non-endemics. Thus, species richness and diversity of soil Collembola can be only partially restored with appropriate reforestation methods, and although it takes many years, to some extent even endemic species can be gradually restored. Nevertheless, the maintenance of undisturbed diversity reservoirs linked by ecological corridors to reforested plots is imperative, as only undisturbed areas can support most of the endemic species able to re-colonize reforested sites.     

Mammals of Australian islands: factors influencing species richness

Distribution patterns of indigenous non-volant terrestrial mammals on 257 Australian islands were examined in relation to environmental parameters and the effects of human-induced disturbance during prehistoric and historic times on island species numbers. Species occurrence for individual species, for taxonomic and trophic groups, and for all species together was related to environmental parameters using regression analysis and the extreme-value function model. Patterns of occurrence were examined separately within three major biogeographic regions derived by pattern analysis. The number of species known to have occurred on these islands during historic times was adequately predicted from area alone. No statistically significant improvement in predicted species number was gained by including island elevation, mean annual rainfall, isolation from the mainland or the number of potentially competing species present on the island. Similarly, no single factor other than area was found to influence consistently the presence of individual species. We conclude that the occurrence of indigenous non-volant terrestrial mammal species on these islands indicates a relictual rather than equilibrial fauna. Visitation by Aboriginal people during prehistoric times did not significantly increase mammal extinctions on islands. Examination of patterns of species richness for a given area on a regional basis showed that islands in and around Bass Strait and Tasmania (Bass Region) were the most species-rich, islands off the northern coasts were slightly less rich, and islands off the south western coasts had fewest species. This is in contrast to the usual latitudinal gradient in species richness patterns. However, islands off the northern and eastern coasts had an overall greater number of different species. When considered in relation to the number of different species of mammals occurring within each region, islands of a given size in Bass Region typically bore a higher proportion of this species pool than other regions. The Bass Region was found to be particularly rich in macropoid herbivores and dasyurid carnivores and insectivores. Analyses indicated that there is a very strong relationship between the presence of exotics as a whole and the local extinction of native mammals. Many mammal species formerly widespread on the Australian mainland are now restricted totally to islands (nine species) or are threatened with extinction on the mainland and have island populations of conservation significance (ten species). In all, thirty-five islands protect eighteen taxa of Australian threatened mammals. The land-use and management of these islands is of considerable importance to nature conservation. The introduction of exotic mammals to these islands should be prevented; any introductions that occur should be eradicated immediately.     

Large-scale species-richness gradients in the Atlantic Ocean

The increase in species richness from the poles to the Equator has been observed in numerous terrestrial and aquatic taxa. A number of different hypotheses have been put forward as explanations for this trend, e.g. area and energy availability. However, whether these hypotheses apply to large spatial scales in marine environments remains unclear. The present study shows a clear latitudinal gradient from high to low latitude (from 80 degrees N to 70 degrees S) in marine species richness for 6643 species (fishes and invertebrates) in 10 different taxa dwelling in benthic and pelagic habitats on both sides of the Atlantic. The patterns in benthic taxa are strongly influenced by coastal hydrographic processes, with marked peaks and troughs, and consequently the gradients are not symmetric along both Atlantic sides. Pelagic taxa show a plateau-shaped distribution and the influence from coastal events on gradients could not be demonstrated. The relationships between species richness and different environmental factors indicate that area size does not explain the latitudinal pattern in benthic species richness on a large spatial scale. Sea-surface temperature (positive relationship) is the best predictor of this pattern for benthic species, and nitrate concentration (negative relationship) is the best predictor for pelagic species. The results call into question the existence of a single primary cause that would explain the pattern in marine species richness on a large spatial scale.     

Human population, grasshopper and plant species richness in European countries

Surprisingly, several studies over large scales have reported a positive spatial correlation of people and biodiversity. This pattern has important implications for conservation and has been documented for well studied taxa such as plants, amphibians, reptiles, birds and mammals. However, it is unknown whether the pattern applies also to invertebrates other than butterflies and more work is needed to establish whether the species–people relationship is explained by both variables correlating with other environmental factors. We studied whether grasshopper species richness (Orthoptera, suborder Caelifera) is related to human population size in European countries. As expected, the number of Caelifera species increases significantly with increasing human population size. But this is not the case when controlling for country area, latitude and number of plant species. Variations in Caelifera species richness are primarily associated with variations in plant species richness. Caelifera species richness also increases with decreasing mean annual precipitation, Gross Domestic Product per capita (used as an indicator for economic development) and net fertility rate of the human population. Our analysis confirms the hypothesis that the broad-scale human population–biodiversity correlations can be explained by concurrent variations in factors other than human population size such as plant species richness, environmental productivity, or habitat heterogeneity. Nonetheless, more populated countries in Europe still have more Caelifera species than less populated countries and this poses a particular challenge for conservation.     

Species–area relationships in the Andaman and Nicobar Islands emerge because rarer species are disproportionately favored on larger islands

The island species–area relationship (ISAR) describes how the number of species increases with increasing size of an island (or island‐like habitat), and is of fundamental importance in island biogeography and conservation. Here, we use a framework based on individual‐based rarefaction to infer whether ISARs result from passive sampling, or whether some processes are acting beyond sampling (e.g., disproportionate effects and/or habitat heterogeneity). Using data on total and relative abundances of four taxa (birds, butterflies, amphibians, and reptiles) from multiple islands in the Andaman and Nicobar archipelago, we examine how different metrics of biodiversity (total species richness, rarefied species richness, and abundance‐weighted effective numbers of species emphasizing common species) vary with island area. Total species richness increased for all taxa, as did rarefied species richness controlling for a given sampling effort. This indicates that the ISAR did not result because of passive sampling, but that instead, some species were disproportionately favored on larger islands. For birds, frogs, and lizards, this disproportionate effect was only associated with species that were rarer in the samples, but for butterflies, both more common and rarer species were affected. Furthermore, for the two taxa for which we had plot‐level data (reptiles and amphibians), within‐island β‐diversity did not increase with island size, suggesting that within‐island compositional effects were unlikely to be driving these ISARs. Overall, our results indicate that the ISARs of these taxa are most likely driven by disproportionate effects, that is, where larger islands are important sources of biodiversity beyond a simple sampling expectation, especially through their influence on rarer species, thus emphasizing their role in the preservation and conservation of species.     

Springtail community (Hexapoda,Collembola) in arable chernozems of Kursk oblast

The composition of springtail communities (the ratio of taxa, species richness, morphoecological spectrum, and spatial organization) in typical deep chernozems of Kursk oblast has been studied in crop fields, forest strips, and an old fallow. Approximately 40 springtail species have been recorded, with the Onychiuridae being dominant with respect to species diversity and contribution to the total abundance. Springtails are distributed over the soil profile to a depth of no less than 50 cm. The pattern of their vertical distribution in areas with tree and shrub vegetation differs from that in arable land. The response of springtails to weakly expressed sheet erosion of chernozem proved to be insufficient for using them as bioindicators of the initial stages of soil erosion.     

Ecological biogeography of southern ocean islands: the importance of considering spatial issues

Understanding patterns of among-island variation in species richness has long been an important question in ecology and biogeography. However, despite the clear spatial nature of the data used for such investigations, the spatial distribution of the different sampled locations is rarely explicitly considered, which may be critical for statistical and biological reasons. In a recent study, Chown et al. (1998) investigated the relationships between species richness of different indigenous and introduced taxonomic groups and a variety of variables characterizing Southern Ocean islands, and here, we use these data to address spatial issues. As predicted, we found spatial autocorrelation in species richness for terrestrial taxa with high dispersal ability or for terrestrial taxa that had time to disperse locally (introduced land birds and indigenous taxa) but not for taxa that had low opportunity to disperse to nearby islands (introduced plants, insects, and mammals), which suggests that colonization from nearby islands has played an important role in shaping present-day patterns of among-island variation in species richness. Interestingly, in several cases, the estimated effect of variables changed when spatial covariance was incorporated. Moreover, the absence of autocorrelation of some variables allowed us to confirm some important results of Chown et al. (1998), notably those involving the potential impact of human presence on the biodiversity of these islands. Overall, our results illustrate the importance of considering spatial structures in ecological studies. This is notably the case when dispersal processes can be expected to explain some of the observed patterns.     

Determinants of the diversity of plants, birds and mammals of coastal islands of the Humboldt current systems: implications for conservation

Sound conservation plans for islands require understanding the processes underlying to the patterns of species richness and composition. Larger islands are often the targets of conservation assuming that the island area mainly determines species richness, and that species composition is nested across islands. However, in small-island these patterns could be altered because of stochastic processes, and species assemblages could be disharmonious. In addition, human impact could further modify the distribution pattern and diversity. Here we use the case of seven islands from the coastal system of Coquimbo as a model to address the role of environmental variables and human impacts on species richness and assembly rules of plants, birds, and mammals. We hypothesize (a) the existence of a small-island effect, and the prevalence of habitat diversity and anthropogenic impacts as main drivers of species richness, and (b) the existence of disharmonious assemblages, characterized by a low degree of nestedness and random patterns of species co-occurrence. Our results showed that (a) species richness is mainly correlated with habitat diversity, and only weakly related to island area supporting the ‘small-island effect’ and (b) species composition is highly structured, but that such structure may be the result of anthropogenic activities. Nestedness was observed in plants and landbirds, while co-occurrence patterns were only detected in plants. Assemblages in small-islands departed from the nestedness pattern and maintain rare species. Currently, only three of the seven islands are protected by national regulations, excluding the smaller ones that are subjected to human disturbance and invasive mammals. Our study suggests that it necessary to include all the islands in a major protected area to preserve both richness and species composition of a number of representative islands of the Humboldt current systems. We showed that conservation plans solely based on island area might not be robust.     

Deep phylogeographic divisions and long‐term persistence of forest invertebrates (Hexapoda: Collembola) in the North‐Western Mediterranean basin

The North‐Western Mediterranean basin is well known for its high number of relictual endemic taxa, and has been indicated as one of the world’s major biodiversity hotspots at the species level. A possible contributing factor may be long‐term persistence of populations and their prolonged stability. This study was designed to investigate the phylogeographic structure of three common species of the genus Lepidocyrtus (Hexapoda: Collembola), soil‐dwelling arthropods characterized by limited dispersal capabilities and generally associated with forest habitats. We observed a remarkable geographic structure, with numerous deeply divergent genetic lineages occupying islands as well as mainland sites with no apparent gene flow among most sites, even across distances of only tens of kilometres. The reconstructed time frame for the evolution of these lineages suggests divergence between 5 and 15 Ma. This indicates a remarkably ancient origin and long‐term persistence of individual lineages over a fine geographic scale despite the occurrence of abrupt sea level and climatic fluctuations in the area. This further suggests that currently recognized morphological species might be a serious underestimation of the true springtail biodiversity within this region.     

Species Richness-Environment Relationships of European Arthropods at Two Spatial Grains: Habitats and Countries

We study how species richness of arthropods relates to theories concerning net primary productivity, ambient energy, water-energy dynamics and spatial environmental heterogeneity. We use two datasets of arthropod richness with similar spatial extents (Scandinavia to Mediterranean), but contrasting spatial grain (local habitat and country). Samples of ground-dwelling spiders, beetles, bugs and ants were collected from 32 paired habitats at 16 locations across Europe. Species richness of these taxonomic groups was also determined for 25 European countries based on the Fauna Europaea database. We tested effects of net primary productivity (NPP), annual mean temperature (T), annual rainfall (R) and potential evapotranspiration of the coldest month (PET_min) on species richness and turnover. Spatial environmental heterogeneity within countries was considered by including the ranges of NPP, T, R and PET_min. At the local habitat grain, relationships between species richness and environmental variables differed strongly between taxa and trophic groups. However, species turnover across locations was strongly correlated with differences in T. At the country grain, species richness was significantly correlated with environmental variables from all four theories. In particular, species richness within countries increased strongly with spatial heterogeneity in T. The importance of spatial heterogeneity in T for both species turnover across locations and for species richness within countries suggests that the temperature niche is an important determinant of arthropod diversity. We suggest that, unless climatic heterogeneity is constant across sampling units, coarse-grained studies should always account for environmental heterogeneity as a predictor of arthropod species richness, just as studies with variable area of sampling units routinely consider area.     

Global patterns and predictors of tropical reef fish species richness

In the marine realm, the tropics host an extraordinary diversity of taxa but the drivers underlying the global distribution of marine organisms are still under scrutiny and we still lack an accurate global predictive model. Using a spatial database for 6336 tropical reef fishes, we attempted to predict species richness according to geometric, biogeographical and environmental explanatory variables. In particular, we aimed to evaluate and disentangle the predictive performances of temperature, habitat area, connectivity, mid‐domain effect and biogeographical region on reef fish species richness. We used boosted regression trees, a flexible machine‐learning technique, to build our predictive model and structural equation modeling to test for potential ‘mediation effects’ among predictors. Our model proved to be accurate, explaining 80% of the total deviance in fish richness using a cross‐validated procedure. Coral reef area and biogeographical region were the primary predictors of reef fish species richness, followed by coast length, connectivity, mid‐domain effect and sea surface temperature, with interactions between the region and other predictors. Important indirect effects of water temperature on reef fish richness, mediated by coral reef area, were also identified. The relationship between environmental predictors and species richness varied markedly among biogeographical regions. Our analysis revealed that a few easily accessible variables can accurately predict reef fish species richness. They also highlight concerns regarding ongoing environmental declines, with region‐specific responses to variation in environmental conditions predicting a variable response to anthropogenic impacts.     

Dispersal ability modulates the strength of the latitudinal richness gradient in European beetles

Aim A debate exists as to whether present‐day diversity gradients are governed by current environmental conditions or by changes in environmental conditions through time. Recent studies have shown that latitudinal richness gradients might be partially caused by incomplete post‐glacial recolonization of high‐latitude regions; this leads to the prediction that less mobile taxa should have steeper gradients than more mobile taxa. The aim of this study is to test this prediction. Location Europe. Methods We first assessed whether spatial turnover in species composition is a good surrogate for dispersal ability by measuring the proportion of wingless species in 19 European beetle clades and relating this value to spatial turnover (β_sim) of the clade. We then linearly regressed β_sim values of 21 taxa against the slope of their respective diversity gradients. Results A strong relationship exists between the proportion of wingless species and β_sim, and β_sim was found to be a good predictor of latitudinal richness gradients. Main conclusions Results are consistent with the prediction that poor dispersers have steeper richness gradients than good dispersers, supporting the view that current beetle diversity gradients in Europe are affected by post‐glacial dispersal lags.     

Belowground carabid beetle diversity in the western Palaearctic - effects of history and climate on range-restricted taxa (Coleoptera, Carabidae)

Broad-scale patterns of subterranean diversity are a fascinating but neglected part of biodiversity research. Carabid beetles adapted to belowground habitats form a particularly species-rich part of the subterranean fauna. We studied large-scale diversity patterns of these belowground carabids across the western Palaearctic and evaluated potential impacts of historical and contemporary environmental conditions on the distribution of these taxa, using available species richness and environmental data at country level. Regression modelling and variation partitioning showed a strong relationship between species richness and range in elevation. Potential effects of climatic variables, mainly those related to ambient energy input, were much weaker. We discuss the implications of this combination of effects, which suggests, concordant with the absence of subterranean carabids in northern and highest richness in southern Europe, a strong prevailing influence of historical processes on current richness distributions of these taxa. Previous studies did not provide clear indications for such an influence. In contrast to more mobile and widespread carabid beetles, dispersal limitation due to high adaptation of belowground carabids to subterranean habitats has probably hindered their re-colonization of former permafrost and glaciated regions. Hotspots of highest belowground diversity are located in regions with an assumed long-term stability of environmental conditions, correlating with patterns of other dispersal-limited taxa such as many endemic plants. Our study provides important new information in the discussion of potential determinants of the distinct geographic patterns of belowground diversity. Moreover, it contributes to a better understanding of range size related differences previously found in the distribution of diversity and environmental dependencies of widespread and range-restricted species within the highly diverse carabid beetles.     

Adaptive management as a tool to improve the conservation of endemic floras: the case of Sicily, Malta and their satellite islands

The presence of endemic species is among the fundamental criteria for characterizing the biodiversity of a territory. Analyzing species richness, extinction level and distribution drivers is an important preliminary step to set conservation priorities and test environmental policies. This study applied the concept of adaptive management to develop strategies for the conservation of endemic floras by considering, as a case study, Sicily, Malta and their neighboring small islands. Adaptive management can be defined as the systematic acquisition and application of reliable information to improve management over time. The development of adaptive conservation strategies aimed preliminary: (1) to quantify endemic plant diversity; (2) to assess the current IUCN knowledge; (3) to analyze the spatial patterns of species distribution in relation to number of colonized habitats, preferential habitats, altitudinal range, and bedrock; (4) to assess whether Natura 2000 network contributed significantly to increase the overlap between endemic distributional areas and protected surface. Strictly Sicilian endemics were 202 taxa amounting to 7.0 % of the whole native flora (c. 2900 taxa). The current picture of extinction risk is still incomplete because over 50 % of endemics were never assessed or assessed with old IUCN criteria. The spatial range size of endemics depended by 40 % on bedrock, and altitudinal and niche breadth. Habitat type did not influence the range size of endemics. The overlap between endemic distributional areas and protected surface increased from 41.3 to 63.3 % with Natura 2000. Adaptive management needs measurable goals to test the progressive improvement of conservation strategies over time, and the reduction in threatened species may be considered as an indicator of successful conservation outcomes. Feedback plays an important role in the periodic revision of biodiversity assessment, distribution modeling, and environmental management, which are fundamental to predict conservation outcomes in the face of extreme uncertainty. In particular, the exhaustive knowledge of the IUCN status is a primary step to implement adaptive measures of conservation, especially as regards endemic floras that are potentially more vulnerable to large-scale or unpredictable and stochastic threats.     

Plants on small islands revisited: the effects of spatial scale and habitat quality on the species–area relationship

Understanding how species diversity is related to sampling area and spatial scale is central to ecology and biogeography. Small islands and small sampling units support fewer species than larger ones. However, the factors influencing species richness may not be consistent across scales. Richness at local scales is primarily affected by small‐scale environmental factors, stochasticity and the richness at the island scale. Richness at whole‐island scale, however, is usually strongly related to island area, isolation and habitat diversity. Despite these contrasting drivers at local and island scales, island species–area relationships (SARs) are often constructed based on richness sampled at the local scale. Whether local scale samples adequately predict richness at the island scale and how local scale samples influence the island SAR remains poorly understood. We investigated the effects of different sampling scales on the SAR of trees on 60 small islands in the Raja Ampat archipelago (Indonesia) using standardised transects and a hierarchically nested sampling design. We compared species richness at different grain sizes ranging from single (sub)transects to whole islands and tested whether the shape of the SAR changed with sampling scale. We then determined the importance of island area, isolation, shape and habitat quality at each scale on species richness. We found strong support for scale dependency of the SAR. The SAR changed from exponential shape at local sampling scales to sigmoidal shape at the island scale indicating variation of species richness independent of area for small islands and hence the presence of a small‐island effect. Island area was the most important variable explaining species richness at all scales, but habitat quality was also important at local scales. We conclude that the SAR and drivers of species richness are influenced by sampling scale, and that the sampling design for assessing the island SARs therefore requires careful consideration.