Human land use promotes the abundance and diversity of exotic species on Caribbean islands

Human land use causes major changes in species abundance and composition, yet native and exotic species can exhibit different responses to land use change. Native populations generally decline in human‐impacted habitats while exotic species often benefit. In this study, we assessed the effects of human land use on exotic and native reptile diversity, including functional diversity, which relates to the range of habitat use strategies in biotic communities. We surveyed 114 reptile communities from localities that varied in habitat structure and human impact level on two Caribbean islands, and calculated species richness, overall abundance, and evenness for every plot. Functional diversity indices were calculated using published trait data, which enabled us to detect signs of trait filtering associated with impacted habitats. Our results show that environmental variation among sampling plots was explained by two Principal Component Analysis (PCA) ordination axes related to habitat structure (i.e., forest or nonforest) and human impact level (i.e., addition of man‐made constructions such as roads and buildings). Several diversity indices were significantly correlated with the two PCA axes, but exotic and native species showed opposing responses. Native species reached the highest abundance in forests, while exotic species were absent in this habitat. Human impact was associated with an increase in exotic abundance and species richness, while native species showed no significant associations. Functional diversity was highest in nonforested environments on both islands, and further increased on St. Martin with the establishment of functionally unique exotic species in nonforested habitat. Habitat structure, rather than human impact, proved to be an important agent for environmental filtering of traits, causing divergent functional trait values across forested and nonforested environments. Our results illustrate the importance of considering various elements of land use when studying its impact on species diversity and the establishment and spread of exotic species.     

Scale-dependent relationships between native richness, resource stability and exotic cover in dock fouling communities of Washington, USA

Aim In terrestrial plant communities, the relationship between native species diversity and exotic success is typically scale‐dependent. It is often proposed that within local neighbourhoods, high native diversity limits resources, thereby inhibiting exotic success. However, environmental variation that manifests over space or time can create positive correlations between native diversity and exotic success at larger scales. In marine habitats, there have been few multi‐scale surveys of this pattern, so it is unclear how diversity, resource limitation and the environment influence the success of exotic species in these systems. Location Washington, USA. Methods I analysed nested spatial and temporal surveys of fouling communities, which are assemblages of sessile marine invertebrates, to test whether the relationships between native richness, resource availability and exotic cover supported the diversity‐stability and diversity‐resistance theories, to test whether these relationships changed with spatio‐temporal scale, and to explore the temperature preferences of native and exotic fouling species. Results Survey data failed to support diversity‐stability theory: space availability actually increased with native richness at the local neighbourhood scale, and neither space availability nor variability decreased with native richness across larger spatio‐temporal scales. I did find support for diversity‐resistance theory, as richness negatively correlated with exotic cover in local neighbourhoods. Unexpectedly, this negative correlation disappeared at intermediate scales, but emerged again at the regional scale. This scale‐dependent pattern could be partially explained by contrasting water temperature preferences of native and exotic species. Main conclusions Within local neighbourhoods, native diversity may inhibit exotic abundance, but the mechanism is unlikely related to resource limitation. At the largest scale, correlations suggest that native richness is higher in cooler environments, whereas exotic richness is higher in warmer environments. This large‐scale pattern contrasts with the typical plant community pattern, and has important implications for coastal management in the face of global climate change.     

Life history of peracarid species in South‐western Atlantic: Comparison of population traits between native and exotic species

Life‐history traits of exotic species are important to understand the process involved in their settlement and their potential impact on native biodiversity. In this context, the seasonal density, the population structure and the reproductive patterns of exotic and native peracarid species of two natural marine environments of Southwestern Atlantic were studied in order to determine the traits that favour the invasion success of exotic species. Five samples, consisted of algal patches (0.20 × 0.20 m quadrants), were collected seasonally from 2016 to 2017 in two intertidal environments (La Estafeta and Cerro Avanzado). Both environments presented high richness of cryptogenic and exotic species (Tanais dulongii, Monocorophium insidiosum, Ampithoe valida, Melita palmata and Jassa marmorata), and only two native species were recorded (Apohyale grandicornis and Exosphaeroma lanceolatum). The comparison of life‐history traits suggested that the distribution, dominance and the highest densities of some exotic species are closely related to their continuous reproductive and recruitment periods, their capability to adapt their life‐history strategies to different environmental conditions and to a more efficient distribution of resources during reproduction; however, in native species, only A. grandicornis registered similar life‐history traits than exotic populations, suggesting that their distribution could be limited by a latitudinal gradient. We expect that these results provide essential information to understand the invasion pattern of exotic species and their potential impact on native biodiversity in Southwestern Atlantic.     

Comparison of native and exotic distribution and richness models across scales reveals essential conservation lessons

Comparing native and exotic plant species distribution and richness models can help to reveal the causes of invasive exotic species proliferation and provide recommendations for preserving native‐dominated ecosystems. However, models may have limited applicability if potentially divergent patterns across scales, spatial autocorrelation and correspondence with community‐wide patterns such as species richness are not considered. I modeled the distributions of 20 dominant native and 20 dominant exotic species among and within patches in a heavily‐invaded and threatened ecosystem in western North America, examining the roles of scale and species origin on variable selection, spatial autocorrelation and model accuracy to determine conditions that favour native over exotic dominants, and derive recommendations for effective management. I also compared distribution models with native and exotic species richness models, to determine the extent to which dominant native and exotic species were representative of synoptic community patterns. Predictability was lower for exotic dominants, possibly because they are environmental generalists, and was lower within than among patches. Predictors were generally shared between distribution and richness models; however, species‐specific differences were common within both native and exotic species groups. Predictors for individual species across scales were frequently different and sometimes opposing. Distribution and richness models suggest that management assuming environmental affiliation at one scale may be ineffective at another; that site prioritization to maximize native versus exotic richness may not preserve the habitat of some common native species; and that intensive management to reduce exotics may be difficult due to low predictability and shared affiliations with natives. Comparing native and exotic distribution and richness models at two scales enabled scale‐specific conservation recommendations and elucidated trade‐offs between management for richness and representation that distribution models at an individual scale would not have allowed.     

Functional redundancy in heterogeneous environments: implications for conservation

It has been argued that one of the best ways to conserve biological diversity is to maintain the integrity of functional processes within communities, and this can be accomplished by assessing how much ecological redundancy exists in communities. Evidence suggests, however, that the functional roles species play are subject to the influences of local environmental conditions. Species may appear to perform the same function (i.e. be redundant) under a restricted set of conditions, yet their functional roles may vary in naturally heterogeneous environments. Incorporating the environmental context into ecological experiments would provide a critical perspective for examining functional redundancy among species.     

Contrasting patterns and mechanisms of spatial turnover for native and exotic freshwater fish in Europe

Aim We compare the distribution patterns of native and exotic freshwater fish in Europe, and test whether the same mechanisms (environmental filtering and/or dispersal limitation) govern patterns of decrease in similarity of native and exotic species composition over geographical distance (spatial species turnover). Locations Major river basins of Europe. Methods Data related to geography, habitat diversity, regional climate and species composition of native and exotic freshwater fish were collated for 26 major European river basins. We explored the degree of nestedness in native and exotic species composition, and quantified compositional similarity between river basins according to the beta‐sim (independent of richness gradient) and Jaccard (dependent of richness gradient) indices of similarity. Multiple regression on distance matrices and variation‐partitioning approaches were used to quantify the relative roles of environmental filtering and dispersal limitation in shaping patterns of decreasing compositional similarity over geographical distance. Results Native and exotic species exhibited significant nested patterns of species composition, indicating that differences in fish species composition between river basins are primarily the result of species loss, rather than species replacement. Both native and exotic compositional similarity decreased significantly with increasing geographical distance between river basins. However, gradual changes in species composition with geographical distance were found only for exotic species. In addition, exotic species displayed a higher rate of similarity decay (higher species turnover rate) with geographical distance, compared with native species. Lastly, the majority of explained variation in exotic compositional similarity was uniquely related to geography, whereas native compositional similarity was either uniquely explained by geography or jointly explained by environment and geography. Main conclusions Our study suggests that large‐scale patterns of spatial turnover for exotic freshwater fish in Europe are generated by human‐mediated dispersal limitation, whereas patterns of spatial turnover for native fish result from both dispersal limitation relative to historical events (isolation by mountain ranges, glacial history) and environmental filtering.     

Deconstructing the native–exotic richness relationship in plants

Aim Classic theory suggests that species‐rich communities should be more resistant to the establishment of exotic species than species‐poor communities. Although this theory predicts that exotic species should be less diverse in regions that contain more native species, macroecological analyses often find that the correlation between exotic and native species richness is positive rather than negative. To reconcile results with theory, we explore to what extent climatic conditions, landscape heterogeneity and anthropogenic disturbance may explain the positive relationship between native and exotic plant richness. Location Catalonia (western Mediterranean region). Methods We integrated floristic records and GIS‐based environmental measures to make spatially explicit 10‐km grid cells. We asked whether the observed positive relationship between native and exotic plant richness (R²= 0.11) resulted from the addition of several negative correlations corresponding to different environmental conditions identified with cluster analysis. Moreover, we directly quantified the importance of common causal effects with a structural equation modelling framework. Results We found no evidence that the relationship between native and exotic plant richness was negative when the comparison was made within environmentally homogeneous groups. Although there were common factors explaining both native and exotic richness, mainly associated with landscape heterogeneity and human pressure, these factors only explained 17.2% of the total correlation. Nevertheless, when the comparison was restricted to native plants associated with human‐disturbed (i.e. ruderal) ecosystems, the relationship was stronger (R²= 0.52) and the fraction explained by common factors increased substantially (58.3%). Main conclusions While our results confirm that the positive correlation between exotic and native plant richness is in part explained by common extrinsic factors, they also highlight the great importance of anthropic factors that – by reducing biotic resistance – facilitate the establishment and spread of both exotic and native plants that tolerate disturbed environments.     

Predicting to new environments: tools for visualizing model behaviour and impacts on mapped distributions

Data limitations can lead to unrealistic fits of predictive species distribution models (SDMs) and spurious extrapolation to novel environments. Here, we want to draw attention to novel combinations of environmental predictors that are within the sampled range of individual predictors but are nevertheless outside the sample space. These tend to be overlooked when visualizing model behaviour. They may be a cause of differing model transferability and environmental change predictions between methods, a problem described in some studies but generally not well understood. We here use a simple simulated data example to illustrate the problem and provide new and complementary visualization techniques to explore model behaviour and predictions to novel environments. We then apply these in a more complex real‐world example. Our results underscore the necessity of scrutinizing model fits, ecological theory and environmental novelty.     

Identifying Native Vegetation for Reducing Exotic Species during the Restoration of Desert Ecosystems

There is currently much interest in restoration ecology in identifying native vegetation that can decrease the invasibility by exotic species of environments undergoing restoration. However, uncertainty remains about restoration's ability to limit exotic species, particularly in deserts where facilitative interactions between plants are prevalent. Using candidate native species for restoration in the Mojave Desert of the southwestern U.S.A., we experimentally assembled a range of plant communities from early successional forbs to late‐successional shrubs and assessed which vegetation types reduced the establishment of the priority invasive annuals Bromus rubens (red brome) and Schismus spp. (Mediterranean grass) in control and N‐enriched soils. Compared to early successional grass and shrub and late‐successional shrub communities, an early forb community best resisted invasion, reducing exotic species biomass by 88% (N added) and 97% (no N added) relative to controls (no native plants). In native species monocultures, Sphaeralcea ambigua (desert globemallow), an early successional forb, was the least invasible, reducing exotic biomass by 91%. However, the least‐invaded vegetation types did not reduce soil N or P relative to other vegetation types nor was native plant cover linked to invasibility, suggesting that other traits influenced native‐exotic species interactions. This study provides experimental field evidence that native vegetation types exist that may reduce exotic grass establishment in the Mojave Desert, and that these candidates for restoration are not necessarily late‐successional communities. More generally, results indicate the importance of careful native species selection when exotic species invasions must be constrained for restoration to be successful.     

Invasibility and species richness of island endemic arthropods: a general model of endemic vs. exotic species

Aim This paper has two objectives. First, we examine how a variety of biotic, abiotic and anthropogenic factors influence the endemic and introduced arthropod richness on an oceanic island. Second, we look at the relationship between the endemic and introduced arthropod richness, to ask whether areas with high levels of endemic species richness deter invasions. Location The work was carried out on a young volcanic island, Terceira, in the Azores. Methods We used standard techniques to collect data on arthropod species richness. Environmental data were obtained from the CIELO climatic model and using GIS. The explanatory value of environmental variables on a small‐scale gradient of endemic and exotic arthropod species richness was examined with generalized linear models (GLMs). In addition, the impact of both endemic and exotic species richness in the communities was assessed by entering them after the environmental variable(s) to see if they contributed significantly to the final model (the hierarchical method). Results Abiotic (climatic and geomorphological) variables gave a better explanation of the variation in endemic species richness, whereas anthropogenic variables explained most of the variation in introduced species richness. Furthermore, after accounting for all environmental variables, part of the unexplained variance in the endemic species richness is explained by the introduced species richness and vice‐versa. That is, areas with high levels of endemic species richness had many introduced species. There is evidence of a somewhat inverse spatial distribution between a group of oceanic‐type, forest‐dwelling, endemic, relict arthropods and a group of more generalist endemic arthropods that are able to survive in disturbed marginal sites particularly rich in non‐indigenous species. Main conclusions Richness of endemic species is mainly driven by abiotic factors such as a climatic axis (oceanic‐type localities with lower temperatures and summer precipitations) and a binary variable CALD (location of sites in caldeiras or ravines), whereas richness of introduced species depends on disturbance related factors. However, after factoring out these major influences, there is a correlation between endemic and introduced richness, suggesting that – independent of the environmental and geographical factors that affect the distribution of endemic or introduced species – the richest endemic assemblages are more prone to invasion, due probably to a facilitation process. Inconclusive evidence suggests that non‐indigenous species are limited to those sites under anthropogenic influence located mainly near forest edges, but the rate of expansion of those species to high‐altitude, core pristine sites has still to be tested.     

Stress relief may promote the evolution of greater phenotypic plasticity in exotic invasive species: a hypothesis

Invasion ecologists have often found that exotic invaders evolve to be more plastic than conspecific populations from their native range. However, an open question is why some exotic invaders can even evolve to be more plastic given that there may be costs to being plastic. Investigation into the benefits and costs of plasticity suggests that stress may constrain the expression of plasticity (thereby reducing the benefits of plasticity) and exacerbate the costs of plasticity (although this possibility might not be generally applicable). Therefore, evolution of adaptive plasticity is more likely to be constrained in stressful environments. Upon introduction to a new range, exotic species may experience more favorable growth conditions (e.g., because of release from natural enemies). Therefore, we hypothesize that any factors mitigating stress in the introduced range may promote exotic invaders to evolve increased adaptive plasticity by reducing the costs and increasing the benefits of plasticity. Empirical evidence is largely consistent with this hypothesis. This hypothesis contributes to our understanding of why invasive species are often found to be more competitive in a subset of environments. Tests of this hypothesis may not only help us understand what caused increased plasticity in some exotic invaders, but could also tell us if costs (unless very small) are more likely to inhibit the evolution of adaptive plasticity in stressful environments in general.     

Eutrophication,biodiversity loss,and species invasions modify the relationship between host and parasite richness during host community assembly

Host and parasite richness are generally positively correlated, but the stability of this relationship in response to global change remains poorly understood. Rapidly changing biotic and abiotic conditions can alter host community assembly, which in turn, can alter parasite transmission. Consequently, if the relationship between host and parasite richness is sensitive to parasite transmission, then changes in host composition under various global change scenarios could strengthen or weaken the relationship between host and parasite richness. To test the hypothesis that host community assembly can alter the relationship between host and parasite richness in response to global change, we experimentally crossed host diversity (biodiversity loss) and resource supply to hosts (eutrophication), then allowed communities to assemble. As previously shown, initial host diversity and resource supply determined the trajectory of host community assembly, altering post‐assembly host species richness, richness‐independent host phylogenetic diversity, and colonization by exotic host species. Overall, host richness predicted parasite richness, and as predicted, this effect was moderated by exotic abundance—communities dominated by exotic species exhibited a stronger positive relationship between post‐assembly host and parasite richness. Ultimately, these results suggest that, by modulating parasite transmission, community assembly can modify the relationship between host and parasite richness. These results thus provide a novel mechanism to explain how global environmental change can generate contingencies in a fundamental ecological relationship—the positive relationship between host and parasite richness.     

A radiotelemetry study of habitat use by the exotic Ring‐necked Parakeet Psittacula krameri in Belgium

Little is known about the foraging ecology of invasive bird species in Europe. We used radio‐telemetry to assess home‐ranges of breeding male Ring‐necked Parakeets in Brussels. Results indicate that parakeets primarily forage in parks and gardens while avoiding forests. This can probably be explained by the higher food availability in anthropogenic habitats and fits a general pattern that invasive species generally select heavily altered environments.     

The paradox of invasion in birds: competitive superiority or ecological opportunism?

Why can alien species succeed in environments to which they have had no opportunity to adapt and even become more abundant than many native species? Ecological theory suggests two main possible answers for this paradox: competitive superiority of exotic species over native species and opportunistic use of ecological opportunities derived from human activities. We tested these hypotheses in birds combining field observations and experiments along gradients of urbanization in New South Wales (Australia). Five exotic species attained densities in the study area comparable to those of the most abundant native species, and hence provided a case for the invasion paradox. The success of these alien birds was not primarily associated with a competitive superiority over native species: the most successful invaders were smaller and less aggressive than their main native competitors, and were generally excluded from artificially created food patches where competition was high. More importantly, exotic birds were primarily restricted to urban environments, where the diversity and abundance of native species were low. This finding agrees with previous studies and indicates that exotic and native species rarely interact in nature. Observations and experiments in the field revealed that the few native species that exploit the most urbanized environments tended to be opportunistic foragers, adaptations that should facilitate survival in places where disturbances by humans are frequent and natural vegetation has been replaced by man-made structures. Successful invaders also shared these features, suggesting that their success is not a paradox but can be explained by their capacity to exploit ecological opportunities that most native species rarely use.     

Using multi-scale species distribution data to infer drivers of biological invasion in riparian wetlands

Aim Biological invasion is a major conservation problem that is of interest to ecological science. Understanding mechanisms of invasion is a high priority, heightened by the management imperative of acting quickly after species introduction. While information about invading species’ ecology is often unavailable, species distribution data can be collected near the onset of invasion. By examining distribution patterns of exotic and native plant species at multiple spatial scales, we aim to identify the scale (of those studied) that accounts for most variability in exotic species abundance, and infer likely drivers of invasion. Location River Murray wetlands, south‐eastern Australia. Methods A nested, crossed survey design was used to determine the extent of variation in wetland plant abundance, grazing intensity and water depth at four spatial scales (reaches, wetland clumps, wetlands, wetland sections), and among three Depth‐strata. We examined responses of exotic and native species groups (grouped into terrestrial and amphibious taxa), native weeds and 10 individual species using hierarchical ANOVA. Results As a group dominated by terrestrial taxa, exotic species cover varied at reach‐, wetland‐ and section‐scales. This likely reflects differences in abiotic characteristics and propagule pressure at these scales. Groups based on native species did not vary at any scale examined. Cover of 10 species mostly varied among and within wetlands (patterns unrelated to species’ origin or functional group), but species’ responses differed, despite individual plants being similar in size. While flora mostly varied among wetlands, exotic cover varied most among reaches (26%), which was attributed to hydrological modification and human activities. Main conclusions Multi‐scale surveys can rapidly identify factors likely to affect species’ distributions and can indicate where future research should be directed. By highlighting disproportionate variation in exotic cover among reaches, this study suggests that flow regulation and human‐mediated dispersal facilitate exotic plant invasion in River Murray wetlands.     

The inherent multidimensionality of temporal variability: how common and rare species shape stability patterns

Empirical knowledge of diversity–stability relationships is mostly based on the analysis of temporal variability. Variability, however, often depends on external factors that act as disturbances, which makes comparisons across systems difficult to interpret. Here, we show how variability can reveal inherent stability properties of ecological communities. This requires that we abandon one‐dimensional representations, in which a single variability measurement is taken as a proxy for how stable a system is, and instead consider the whole set of variability values generated by all possible stochastic perturbations. Despite this complexity, in species‐rich systems, a generic pattern emerges from community assembly, relating variability to the abundance of perturbed species. Strikingly, the contrasting contributions of different species abundance classes to variability, driven by different types of perturbations, can lead to opposite diversity–stability patterns. We conclude that a multidimensional perspective on variability helps reveal the dynamical richness of ecological systems and the underlying meaning of their stability patterns.     

A latitudinal gradient of beta diversity for exotic vascular plant species in North America

Determining relationships between the ranges of introduced species and geographical and environmental factors is an important step in understanding the mechanisms and processes of the spread of introduced species. In this study, I examined the beta diversity and latitude relationship for all naturalized exotic species of vascular plants in North America at a continental scale. Beta diversity was calculated as the absolute value of the slope of the relationship between the natural logarithm of the Simpson index of similarity (lnS) and spatial distance between pairs of state‐level exotic floras within four latitudinal zones examined. Relative contributions of spatial distance and environmental difference to species turnover between exotic floras were examined. I found that beta diversity decreased monotonically from low to high latitudes: beta diversity for the southernmost zone was shallower than that for the northernmost zone by a factor of 2.6. Regression models of lnS in relation to spatial distance and environmental (climatic and topographical) difference for each latitudinal zone demonstrated that the explanatory power of these variables diminishes monotonically with latitude: the explained variance in lnS is 70.4%, 62.1%, 53.9%, and 33.9%, respectively, for the four latitudinal zones from south to north. For the southernmost zone, 58.3% of the variance in lnS is explained by climate variables and topography, and spatial distance explains only 2.3% of the variance. In contrast, for the northernmost zone, more than half the amount (22.5%) of the explained variance in lnS is attributable to spatial distance, and the remaining (18.9%) of the explained variance is attributable to climate variables and topography.     

Environmental factors determining invasibility of urban waters for exotic macroinvertebrates

Aim Urbanization usually leads to biotic homogenization with a decrease in native species and increase in exotic species. We investigated whether local environmental factors in urban water bodies, such as water quality, habitat structure and biotic interactions, influenced the invasion of these systems by exotic macroinvertebrate species. Location Urban surface water systems in lowlands of the Rhine‐Meuse delta. Methods Presence and abundance of native and exotic macroinvertebrate species were compared between different urban water types and related to environmental variables with multivariate analysis and spearman’s correlations. Moreover, co‐existence of related native and exotic species was studied. Results In total nine exotic species were found in the following taxa: Tricladida (1), Crustacea (5), Bivalvia (1) and Gastropoda (2). Taxonomically related native and exotic crustacean species did not seem to be influenced by competition in nutrient‐rich urban waters; most species showed high abundances. Nevertheless, two exotic crustacean species were much more abundant in waters where other crustacean species were absent, possibly filling empty niches. Native species richness and abundance was positively related to environmental heterogeneity in the form of submerged vegetation. The occurrence and abundance of most exotic species were positively related to several eutrophication indicators, such as nitrate, sludge layer and lemnid vegetation. Main conclusions Exotic species in urban waters were mostly detritivorous or omnivorous and therefore dependent on leaf breakdown. In nutrient‐rich water systems, where food availability was high, exotic crustacean species co‐existed with native crustacean species, while in nutrient‐poor, richly vegetated systems, native Asellidae dominated exotic Asellidae. In the turbid water bodies with very little vegetation, native species richness was low and two exotic crustacean species were relatively abundant in these water systems. Invasibility of urban water systems could be reduced by stimulating the development of submerged and nymphaeid vegetation and decreasing nutrient levels.     

Spatial nonstationarity and scale-dependency in the relationship between species richness and environmental determinants for the sub-Saharan endemic avifauna

Aim This article aims to test for and explore spatial nonstationarity in the relationship between avian species richness and a set of explanatory variables to further the understanding of species diversity variation. Location Sub‐Saharan Africa. Methods Geographically weighted regression was used to study the relationship between species richness of the endemic avifauna of sub‐Saharan Africa and a set of perceived environmental determinants, comprising the variables of temperature, precipitation and normalized difference vegetation index. Results The relationships between species richness and the explanatory variables were found to be significantly spatially variable and scale‐dependent. At local scales > 90% of the variation was explained, but this declined at coarser scales, with the greatest sensitivity to scale variation evident for narrow ranging species. The complex spatial pattern in regression model parameter estimates also gave rise to a spatial variation in scale effects. Main conclusions Relationships between environmental variables are generally assumed to be spatially stationary and conventional, global, regression techniques are therefore used in their modelling. This assumption was not satisfied in this study, with the relationships varying significantly in space. In such circumstances the average impression provided by a global model may not accurately represent conditions locally. Spatial nonstationarity in the relationship has important implications, especially for studies of species diversity patterns and their scaling.     

Species pool size and invasibility of island communities: a null model of sampling effects

The success of alien species on oceanic islands is considered to be one of the classic observed patterns in ecology. Explanations for this pattern are based on lower species richness on islands and the lower resistance of species‐poor communities to invaders, but this argument needs re‐examination. The important difference between islands and mainland is in the size of species pools, not in local species richness; invasibility of islands should therefore be addressed in terms of differences in species pools. Here I examine whether differences in species pools can affect invasibility in a lottery model with pools of identical native and exotic species. While in a neutral model with all species identical, invasibility does not depend on the species pool, a model with non‐zero variation in population growth rates predicts higher invasibility of communities of smaller pools. This is because of species sampling; drawing species from larger pools increases the probability that an assemblage will include fast growing species. Such assemblages are more likely to exclude random invaders. This constitutes a mechanism through which smaller species pools (such as those of isolated islands) can directly underlie differences in invasibility.