Linking biotic homogenization to habitat type, invasiveness and growth form of naturalized alien plants in North America

Aim Biotic homogenization is a growing phenomenon and has recently attracted much attention. Here, we analyse a large dataset of native and alien plants in North America to examine whether biotic homogenization is related to several ecological and biological attributes. Location North America (north of Mexico). Methods We assembled species lists of native and alien vascular plants for each of the 64 state‐ and province‐level geographical units in North America. Each alien species was characterized with respect to habitat (wetland versus upland), invasiveness (invasive versus non‐invasive), life cycle (annual/biennial versus perennial) and habit (herbaceous versus woody). We calculated a Jaccard similarity index separately for native, for alien, and for native and alien species. We used the average of Jaccard dissimilarity index (1 ? Jaccard index) of all paired localities as a measure of the mean beta diversity of alien species for each set of localities examined in an analysis. We used a homogenization index to quantify the effect of homogenization or differentiation. Results We found that (1) wetland, invasive, annual/biennial and herbaceous alien plants markedly homogenized the state‐level floras whereas non‐invasive and woody alien plants tended to differentiate the floras; (2) beta diversity was significantly lower for wetland, invasive, annual/biennial and herbaceous alien plants than their counterparts (i.e. upland, non‐invasive, perennial and woody alien plants, respectively); and (3) upland and perennial alien plants each played an equal role in homogenizing and differentiating the state‐level floras. Main conclusions Our study shows that biotic homogenization is clearly related to habitat type (e.g. wetland versus uplands), species invasiveness and life‐history traits such as life cycle (e.g. annual/biennial and herbaceous versus woody species) at the spatial scale examined. These observations help to understand the process of biotic homogenization resulting from alien vascular plants in North America.     

Integrating the Study of Non-native Plant Invasions across Spatial Scales

Non-native (alien, exotic) plant invasions are affecting ecological processes and threatening biodiversity worldwide. Patterns of plant invasions, and the ecological processes which generate these patterns, vary across spatial scales. Thus, consideration of spatial scale may help to illuminate the mechanisms driving biological invasions, and offer insight into potential management strategies. We review the processes driving movement of non-native plants to new locations, and the patterns and processes at the new locations, as they are variously affected by spatial scale. Dispersal is greatly influenced by scale, with different mechanisms controlling global, regional and local dispersal. Patterns of invasion are rarely documented across multiple spatial scales, but research using multi-scale approaches has generated interesting new insights into the invasion process. The ecological effects of plant invasions are also scale-dependent, ranging from altered local community diversity and homogenization of the global flora, to modified biogeochemical cycles and disturbance regimes at regional or global scales. Therefore, the study and control of invasions would benefit from documenting invasion processes at multiple scales.     

Phylogenetic beta diversity of native and alien species in European urban floras

Aim Human activities have weakened biogeographical barriers to dispersal, increasing the rate of introduction of alien plants. However, their impact on beta diversity and floristic homogenization is poorly understood. Our goal is to compare the phylogenetic beta diversity of native species with that of two groups of alien species, archaeophytes and neophytes (introduced before and after ad 1500, respectively), across European urban floras to explore how biological invasions affect phylogenetic turnover at a continental scale. Location Twenty European cities located in six countries between 49 and 53° N latitude in continental Europe and the British Isles. Methods To compare the phylogenetic beta diversity of native and alien species we use the average phylogenetic dissimilarity of individual floras from their group centroid in multivariate space. Differences in phylogenetic beta diversity among different species groups are then assessed using a randomization test for homogeneity of multivariate dispersions. Results Across European urban floras, and when contrasted with natives, archaeophytes are usually associated with lower levels of phylogenetic beta diversity while neophytes tend to increase phylogenetic differentiation. Main conclusions While archaeophytes tend to promote limited homogenization in phylogenetic beta diversity, because of their diverse geographical origin together with short residence times in the invaded regions, neophytes are not promoting biotic homogenization of urban floras across Europe. Therefore, in spite of the increasing rate of alien invasion, an intense phylogenetic homogenization of urban cities is not to be expected soon.     

Searching for phylogenetic pattern in biological invasions

It has been suggested that alien species with close indigenous relatives in the introduced range may have reduced chances of successful establishment and invasion (Darwin's naturalization hypothesis). Studies trying to test this have in fact been addressing four different hypotheses, and the same data can support some while rejecting others. In this paper, we argue that the phylogenetic pattern will change depending on the spatial and phylogenetic scales considered. Expectations and observations from invasion biology and the study of natural communities are that at the spatial scale relevant to competitive interactions, closely related species will be spatially separated, whereas at the regional scale, species in the same genera or families will tend to co-occur more often than by chance. We also argue that patterns in the relatedness of indigenous and naturalized plants are dependent on the continental/island setting, spatial occupancy levels, and on the group of organisms under scrutiny. Understanding how these factors create a phylogenetic pattern in invasions will help us predict which groups are more likely to invade where, and should contribute to general ecological theory.     

Do climatically similar regions contain similar alien floras? A comparison between the mediterranean areas of central Chile and California

Aim Taxonomic comparisons of alien floras across climatically similar regions have been proposed as a powerful approach for increasing our understanding of plant invasions across scales. However, detailed comparisons between the alien biotas of climatically similar regions are scarce. This study aims to compare the taxonomic patterns of alien species richness in mediterranean‐type climate areas of central Chile and California, in order to better understand how climatically similar regions converge or diverge in terms of their alien flora. Location Central Chile and California, United States. Methods We compared the alien floras of the state of California in the United States and central Chile, considering within‐region variation and taxonomic composition up to the species level. To test for within‐region variation, administrative units and counties were grouped within seven latitudinal bands for each region. We tested for differences in the relative contributions of the various origins of the naturalized species to each region. We used a family naturalization index to establish which families had relatively higher numbers of naturalized species in each region. We evaluated the similarity, using cluster analyses with Jaccard’s similarity index, of alien taxa between regions and latitudinal bands using presence–absence matrices at the species, genus and family levels. We used principal components analysis to determine the presence of a compositional gradient including all latitudinal bands. Results We recorded 1212 alien plant species in California and 593 in central Chile, of which 491 are shared between the two regions. These figures include 25 species that are native to California and 37 that are native to Chile. A comparison between the alien floras of central Chile and California reveals three major trends: (1) higher naturalized species diversity for California than for Chile, at all taxonomic levels; (2) differences in the proportion of species according to origin, with America, Africa, Asia and Australia providing a larger number of species in California than in Chile; (3) segregation between regions in terms of taxonomic composition of their alien flora, and a rather weak differentiation within regions; and (4) a trend towards higher similarity between the alien floras of latitudinal bands associated with higher levels of human disturbances. Main conclusions The alien floras of central Chile and California are significantly different, but this difference diminishes in highly disturbed areas. Thus, the current high levels of species movement caused by globalization, together with increasing levels of anthropogenic disturbances, should reduce the differentiation of the alien floras in these regions, increasing overall biotic homogenization.     

Patterns of alien plant distribution at multiple spatial scales in a large national park: implications for ecology, management and monitoring

Aim Spatial scale is critical for understanding and managing biological invasions. In providing direction to managing alien plant invasions, much emphasis is placed on collecting spatially explicit data. However, insufficient thought is often given to how the data are to be used, frequently resulting in the incompatibility of the data for different uses. This paper explores the role of spatial scale in interpreting, managing and monitoring alien plant invasions in a large protected area. Location Kruger National Park, South Africa. Methods Using 27,000 spatially‐explicit records of invasive alien plants for the Kruger National Park (> 20,000 km²) we assessed alien plant species richness per cell at nine different scales of resolution. Results When assessing the patterns of alien plants at the various scales of resolution, almost identical results are obtained when working at scales of quarter‐degree grids and quaternary watersheds (the fourth level category in South Africa's river basin classification system). Likewise, insights gained from working at resolutions of 0.1–0.5 km and 1–5 km are similar. At a scale of 0.1 × 0.1 km cells, only 0.4% of the Kruger National Park is invaded, whereas > 90% of the park is invaded when mapped at the quarter‐degree cell resolution. Main conclusions Selecting the appropriate scale of resolution is crucial when evaluating the distribution and abundance of alien plant invasions, understanding ecological processes, and operationalizing management applications and monitoring strategies. Quarter‐degree grids and quaternary watersheds are most useful at a regional or national scale. Grid cells of 1 to 25 km² are generally useful for establishing priorities for and planning management interventions. Fine‐scale data are useful for informing management in areas which are small in extent; they also provide the detail appropriate for assessing patterns and rates of invasion.     

Biotic homogenization: a new research agenda for conservation biogeography

Aim Biotic homogenization describes the process by which species invasions and extinctions increase the genetic, taxonomic or functional similarity of two or more biotas over a specified time interval. The study of biotic homogenization is a young and rapidly emerging research area in the budding field of conservation biogeography, and this paper aims to synthesize our current knowledge of this process and advocate a more systematic approach to its investigation. Methods Based on a comprehensive examination of the primary literature this paper reviews the process of biotic homogenization, including its definition, quantification, underlying ecological mechanisms, environmental drivers, the empirical evidence for different taxonomic groups, and the potential ecological and evolutionary implications. Important gaps in our knowledge are then identified, and areas of new research that show the greatest promise for advancing our current thinking on biotic homogenization are highlighted. Results Current knowledge of the patterns, mechanisms and implications of biotic homogenization is highly variable across taxonomic groups, but in general is incomplete. Quantitative estimates are almost exclusively limited to freshwater fishes and plants in the United States, and the principal mechanisms and drivers of homogenization remain elusive. To date research has focused on taxonomic homogenization, and genetic and functional homogenization has received inadequate attention. Trends over the past decade, however, suggest that biotic homogenization is emerging as a topic of greater research interest. Main conclusions My investigation revealed a number of important knowledge gaps and priority research needs in the science of biotic homogenization. Future studies should examine the homogenization process for different community properties (species occurrence and abundance) at multiple spatial and temporal scales, with careful attention paid to the various biological mechanisms (invasions vs. extinctions) and environmental drivers (environmental alteration vs. biotic interactions) involved. Perhaps most importantly, this research should recognize that there are multiple possible outcomes resulting from the accumulation of species invasions and extinctions, including biotic differentiation whereby genetic, taxonomic or functional similarity of biotas decreases over time.     

Does functional homogenization accompany taxonomic homogenization of British birds and how do biotic factors and climate affect these processes?

Environmental change has reshuffled communities often causing taxonomic homogenization rather than differentiation. Some studies suggest that this increasing similarity of species composition between communities is accompanied by an increase in similarity of trait composition—functional homogenization—although different methodologies have failed to come to any consistent conclusions. Functional homogenization could have a large effect on ecosystem functioning and stability. Here, we use the general definition of homogenization as “reduced spatial turnover over time” to compare changes in Simpson's beta diversity (taxonomic turnover) with changes in Rao's quadratic entropy beta diversity (functional turnover) in British breeding birds at three spatial scales. Using biotic and climatic variables, we identify which factors may predispose a site to homogenization. The change in turnover measures between two time periods, 20 years apart, was calculated. A null model approach was taken to identify occurrences of functional homogenization and differentiation independent of changes in taxonomic turnover. We used conditional autoregressive models fitted using integrated nested Laplace approximations to determine how environmental drivers and factors relating to species distributions affect changes in spatial turnover of species and functional diversity. The measurement of functional homogenization affects the chance of rejection of the null models, with many sites showing taxonomic homogenization unaccompanied by functional homogenization, although occurrence varies with spatial scale. At the smallest scale, while temperature‐related variables drive changes in taxonomic turnover, changes in functional turnover are associated with variation in growing degree days; however, changes in functional turnover become more difficult to predict at larger spatial scales. Our results highlight the multifactorial processes underlying taxonomic and functional homogenization and that redundancy in species traits may allow ecosystem functioning to be maintained in some areas despite changes in species composition.     

Spread of common species results in local‐scale floristic homogenization in grassland of Switzerland

Aim We assess changes in plant species richness and changes in species dissimilarity at local scale in Swiss grassland between the time periods 2001–2004 and 2006–2009. Further, we provide an ecological interpretation of the observed taxonomic homogenization of vascular plants. Location Switzerland. Methods Changes in species richness and changes in Simpson dissimilarity index of vascular plants in grassland (meadows and pastures) were examined. The analyses were based on species lists recorded on 339 10‐m² sample plots from a systematic sample covering the entire Switzerland. Each sample plot had been surveyed once in 2001–2004 and once in 2006–2009 with 5 years between the first and the second survey. Changes in species dissimilarity were interpreted by comparing the relative contribution of several indicator species groups. Results Mean species richness of vascular plants in grassland increased during the study period. In contrast, species dissimilarity of plants decreased, suggesting local‐scale floristic homogenization of grassland in Switzerland. It was mostly because of the spread of common species, namely the species that are tolerant to high nutrient levels, the species of low conservation value and the species adapted to moderate temperature levels that led to taxonomic homogenization. Target species for conservation did only marginally affect taxonomic homogenization. In contrast to the predictions from studies of taxonomic homogenization on larger scales, the taxonomic homogenization of grassland at local scale was not explained by the spread of neophytic species. Main conclusions The biotic diversity of grassland in Switzerland changed considerably between 2001–2004 and 2006–2009. The observed taxonomic homogenization was merely because of the spread of common species. Local‐scale changes in land use regimes implemented by agri‐environmental schemes and other conservation efforts on parts of the entire grassland area were, apparently, not enough to prevent the total grassland from recent taxonomic homogenization.     

Biotic homogenization of oceanic islands depends on taxon,spatial scale and the quantification approach

Biotic homogenization reduces the regional distinctiveness of biotas with significant ecological and evolutionary consequences. The outcome of this process may depend on the spatial scale of inquiry (both resolution and extent), the selected taxon and dissimilarity index as well as on the contribution of species extinctions and introductions. In the present research, we try to disentangle the effects of these factors on homogenization patterns comparing six taxonomic groups (pteridophytes, spermatophytes, breeding birds, mammals, reptiles and non-marine molluscs) within and between five Atlantic archipelagos of the Macaronesian Region. Taxonomic homogenization was analyzed by partitioning β-diversity into spatial turnover of species composition and nestedness. Total compositional change was divided into changes related to extinctions/extirpations of native and to introductions of alien species. Analyses were carried out at two different spatial resolutions (island versus archipelago unit) and geographic extents (within each archipelago and across the whole Macaronesian Region). Pteridophytes and reptiles tended to taxonomic differentiation, while mammals and molluscs showed homogenization regardless of scale and resolution. For spermatophytes, the most species-rich group, taxonomic heterogenization traded off with homogenization from the local to regional extent. Birds revealed heterogenization at the island, but not at the archipelago resolution. Extirpations of native species generally led to homogenization at the local extent, whereas the effect of alien introductions varied according to taxon and spatial scale. Furthermore, overall changes in species pool similarities were driven both by spatial turnover and nestedness. We demonstrate that biotic homogenization after human colonization within Macaronesia clearly depended on taxon, spatial scale and the dissimilarity measure. We suggest that homogenization of island biotas is first conditioned by initial dissimilarity related to taxon characteristics, such as dispersal capacity or endemicity, evolutionary processes, archipelago configurations and environmental variation along spatial scales. Thus, similarity change is the outcome of the impacts of number, proportion and distribution type of lost and gained species. Rare extirpated and common introduced species homogenize, while common extirpated and rare introduced species differentiate island biotas. Partitioning of beta diversity helps to improve our understanding of the homogenization process.     

Taxonomic homogenization and differentiation across Southern Ocean Islands differ among insects and vascular plants

Aim To investigate taxonomic homogenization and/or differentiation of insect and vascular plant assemblages across the Southern Ocean Islands (SOI), and how they differ with changing spatial extent and taxonomic resolution. Location Twenty‐two islands located across the Southern Ocean, further subdivided into five island biogeographical provinces. These islands are used because comprehensive data on both indigenous and non‐indigenous insect and plant species are available. Methods An existing database was updated, using newly published species records, identifying the indigenous and non‐indigenous insect and vascular plant species recorded for each island. Homogenization and differentiation were measured using Jaccard’s index (JI) of similarity for assemblages across all islands on a pairwise basis, and for island pairs within each of the biogeographical provinces. The effects of taxonomic resolution (species, genus, family) and distance on levels of homogenization or differentiation were examined. To explore further the patterns of similarity among islands for each of the taxa and groupings (indigenous and non‐indigenous), islands were clustered based on JI similarity matrices and using group averaging. Results Across the SOI, insect assemblages have become homogenized (0.7% increase in similarity at species level) while plant assemblages have become differentiated at genus and species levels. Homogenization was recorded only when pairwise distances among islands exceeded 3000 km for insect assemblages, but distances had to exceed 10,000 km for plant assemblages. Widely distributed non‐indigenous plant species tend to have wider distributions across the SOI than do their insect counterparts, and this is also true of the indigenous species. Main conclusions Insect assemblages across the SOI have become homogenized as a consequence of the establishment of non‐indigenous species, while plant assemblages have become more differentiated. The likely reason is that indigenous plant assemblages are more similar across the SOI than are insect assemblages, which show greater regionalization. Thus, although a suite of widespread, typically European, weedy, non‐indigenous plant species has established on many islands, the outcome has largely been differentiation. Because further introductions of insects and vascular plants are probable as climates warm across the region, the patterns documented here are likely to change through time.     

Native and alien plant species richness in relation to spatial heterogeneity on a regional scale in Germany

Aim The aim of our study was to reveal relationships between richness patterns of native vs. alien plant species and spatial heterogeneity across varying landscape patterns at a regional scale. Location The study was carried out in the administrative district of Dessau (Germany), covering around 4000 km². Methods Data on plant distribution of the German vascular flora available in grid cells covering 5′ longitude and 3′ latitude (c. 32 km²) were divided into three status groups: native plants, archaeophytes (pre 1500 AD aliens) and neophytes (post 1500 AD aliens). Land use and abiotic data layers were intersected with 125 grid cells comprising the selected area. Using novel landscape ecological methods, we calculated 38 indices of landscape composition and configuration for each grid cell. Principal components analysis (PCA) with a set of 29 selected, low correlated landscape indices was followed by multiple linear regression analysis. Results PCA reduced 29 indices to eight principal components (PCs) that explained 80% cumulative variance. Multiple linear regression analysis was highly significant and explained 41% to 60% variance in plant species distribution (adjusted R²) with three significant PCs (tested for spatial autocorrelation) expressing moderate to high disturbance levels and high spatial heterogeneity. Comparing the significance of the PCs for the species groups, native plant species richness is most strongly associated with riverine ecosystems, followed by urban ecosystems, and then small‐scale rural ecosystems. Archaeophyte and neophyte richness are most strongly associated with urban ecosystems, followed by small‐scale rural ecosystems and riverine ecosystems for archaeophytes, and riverine ecosystems and small‐scale rural ecosystems for neophytes. Main conclusions Our overall results suggest that species richness of native and alien plants increases with moderate levels of natural and/or anthropogenic disturbances, coupled with high levels of habitat and structural heterogeneity in urban, riverine, and small‐scale rural ecosystems. Despite differences in the order of relevance of PCs for the three plant groups, we conclude that at the regional scale species richness patterns of native plants as well as alien plants are promoted by similar factors.     

Differences in the trait compositions of non-indigenous and native plants across Germany

This paper explores the differences in the trait compositions of non-indigenous (neophytic) and native plant species for selected traits in Germany. Our set of functional traits addresses species’ reproductive biology, life history, morphology and ecophysiology. To take account of broad-scale heterogeneity across the country we compared the relative frequencies of neophytes and natives with particular trait attributes at the scale of grid cells (c. 130 km² each). Subsequently, we compared the differences at the grid cell scale to the differences in the corresponding comparisons at the scale of the entire country. Finally, we explored how variation in the trait compositions of the non-indigenous species across the country relates to variation in the trait compositions of the natives. We found remarkable differences in the trait compositions of neophytes and natives at the grid cell scale. Neophytes were over-represented in insect- and self-pollinated species and in species with a later and longer flowering season. Furthermore, the proportions of species with mesomorphic or hygromorphic leaf anatomy, of annual herbs and of trees as well as of non-clonals and polyploids were significantly higher in neophytes than in natives. These differences at the grid cell scale could vary distinctly from the corresponding differences observed at the country scale. This result highlights the complexity of the invasion process and suggests an importance of spatial scale for the comparisons. Correlation analysis indicated, that for traits relating to plant morphology and ecophysiology, the relative frequencies of the non-indigenous species increased with those of the natives. This suggests that favourable environments for natives with particular attributes constitute an increased suitability for neophytes with these attributes as well. Our study provides a step forward towards an integrated understanding of traits in plant invasions across spatial scales and broad-scale heterogeneity and underlines the necessity to understand the role of functional traits in plant invasions with reference to spatial scale and in the context of the environment.     

Scale dependence of plant species richness in a network of protected areas

Despite the widely recognised importance of reserve networks, their effectiveness in encompassing and maintaining biodiversity is still debated. Species diversity is one of the most affordable measures of biodiversity, but it is difficult to survey such data over large scales. This research aimed to perform a sample-based assessment of species richness of groups of plants with different conservation value (alien species, protected species, and all species) within a reserve network, testing the use of partitioning as a tool for assessing diversity at different spatial scales, from the plot to the entire network. Plant diversity patterns differed for the groups of species for most of the investigated spatial scales. Despite these patterns assumed divergent tendencies when different species groups were considered, most of the species richness within the network was given by larger scale β-diversity for both alien and protected species, as well for all species. Diversity partitioning proved an effective tool to quantify the role of spatial scales in structuring the total species richness of the network, and is helpful in planning reserve networks.     

Do non‐native species invasions lead to biotic homogenization at small scales? The similarity and functional diversity of habitats compared for alien and native components of Mediterranean floras

Although a number of recent studies have demonstrated biotic homogenization, these have mainly focused on larger spatial scales. Homogenizing effects are equally important at finer resolutions, e.g. through increasing similarity between habitats, which may result in a simplification of ecosystem structure and function. One major cause of homogenization is the expanding ranges of alien species, although it is not clear whether they are inherently homogenizing at smaller scales. We therefore assessed whether the alien flora is less complex across habitats than the resident native flora of Mediterranean Islands. From a regional data base, we examined floristic lists for between‐habitat taxonomic and functional similarity, and within‐habitat functional diversity, using resampled data sets to control for sample size biases. Aliens and natives showed equivalent complexity in most respects. At the taxonomic level, between‐island and between‐habitat similarities were almost identical, and when ecosystem function was measured by a functional group classification system, this was also true of between‐habitat similarities and within‐habitat diversities. When ecosystem function was measured using Grime's CSR classification, aliens were found to be more functionally homogenous between‐habitats and less functionally diverse within habitats. However, since the CSR profiles of aliens and natives differed, simplification is not inevitable due to ecological segregation of the two floras (aliens tend to be recruited to disturbed habitats rather than displacing natives). One deficiency is a lack of large scale species abundance data. A simple simulation exercise indicated that this is likely to lead to substantial overestimation of true levels of similarity, although would only influence the comparison between aliens and natives if they have different abundance distribution curves. The results indicate that alien floras are not intrinsically more simple than natives, but a higher proportion of competitive strategists among aliens may still cause small‐scale homogenization as these include many strong competitors that are likely to dominate communities.     

Changes in taxonomy and species distributions and their influence on estimates of faunal homogenization and differentiation in freshwater fishes

Aim To assess how changing taxonomy and distribution data affect estimates of faunal homogenization and differentiation as agents of global change in freshwater fishes. Location Provinces and territories of Canada. Methods Species presence–absence data were collated in 2000 and 2005 from regional and national lists, and faunal homogenization and differentiation were calculated using Jaccard’s faunal similarity index. Differences between time periods and areas were summarized using principal coordinate analysis. Differences in faunal assemblages between native and total faunas were assessed via Whittaker’s (1960) beta diversity (β_w) index and tests of differences in multivariate dispersion of fish species compositions. Results Among aquatic ecoregions in one province (British Columbia) there were four taxonomic changes and 18 distributional changes between the 2000 and 2005 databases. Pairwise Jaccard’s faunal similarity index between the eight aquatic ecoregions declined by an average of 4.8% from 35.9% in native faunas to 31.1% in total faunas (introductions – extinctions/extirpations) indicating overall faunal differentiation. Average pairwise similarity declined by 0.9% between 2000 and 2005. Across thirteen provinces and territories of Canada, there were five taxonomic changes and 61 distributional changes between the 2000 and 2005 databases. Generally, faunal homogenization increased; pairwise Jaccard’s increased by an average of 1.8% from 27.1% in native faunas to 28.9% in total faunas or an average of 0.6% per comparison. Main conclusions Despite changing taxonomy and fish distribution information, comparative analysis of 2000 and 2005 databases consistently show overall faunal differentiation at the smallest (provincial) spatial scale and homogenization at the largest scale (across Canada) and that these trends continued between time periods. Homogenization and differentiation followed expectations from conceptual models based on the relative prevalence of species invasions and extinctions within communities. General conclusions of the onset and extent of homogenization and differentiation were relatively insensitive to our changing understanding of taxonomy and distribution.     

Human Agency in Biological Invasions: Secondary Releases Foster Naturalisation and Population Expansion of Alien Plant Species

The human mediation of biological invasions is still an underestimated phenomenon. This paper attempts to show that introductions on varying spatial scales may strongly foster invasions throughout the whole invasion process. As shown by data from central Europe, invasions frequently result from an interplay of biological and anthropogenic mechanisms. The latter, however, cannot be explained nor predicted by ecological rules. This may be an important reason for the limited predictability of invasions. Initial introductions from a donor to a new range are here distinguished from following secondary releases within the new range. The rate of naturalisation is higher in deliberately introduced plants as compared to accidental introductions. Due to higher numbers of accidental introductions, such species contribute significantly to the pool of naturalised species. Secondary releases of alien species are frequently made over long periods subsequent to the initial introduction. They may mimic demographic and dispersal processes that lead to population growth and range expansion. They also offer a pathway to overcome spatial isolation in species whose propagules are not naturally moved long distances. This even holds for most of Germany's noxious alien plant species. Secondary releases may thus promote invasions even beyond the threshold of naturalisation. In consequence, attempts at prevention should focus on secondary releases as well as on initial introductions. In the last section of the paper, the final invasion stage subsequent to naturalisation is shown as a multi-scale phenomenon. In consequence, the classification of a species as 'invasive' depends on the perspective chosen. Using different biologically or anthropocentrically based approaches leads to sub-sets of alien species that overlap only partially. In conclusion, the term `invasive' should preferably be used in a broader sense to describe the entire invasion process. This revised version was published online in July 2006 with corrections to the Cover Date.     

Taxonomic homogenization in ungulates: patterns and mechanisms at local and global scales

Aim The aim of this paper is to examine taxonomic homogenization in ungulates globally and at the local scale in South Africa. Specifically, we aim to examine the roles of distance, scale, time, extinctions vs. introductions, and extralimital vs. extraregional introductions in the homogenization of ungulate biotas, and to determine pathways of introduction of ungulate species globally and the proximate explanatory variables of ungulate introductions in South Africa. Location Forty‐one countries globally and three spatial resolutions in South Africa. Methods Indigenous, extirpated and established introduced ungulate species data were obtained for countries globally, and at a quarter‐degree grid‐cell resolution in South Africa. Homogenization was calculated using Jaccard’s index of similarity (JI) for countries globally and for three spatial resolutions in South Africa. Zoo holdings and transfer data from the International Species Information System database were used to investigate the relationship between non‐indigenous ungulate species introductions and the number of non‐indigenous ungulate species in zoos. Relationships between JI and species richness, and between numbers of introductions and several environmental and social factors were examined using generalized linear models. Results Homogenization in ungulates was 2% for countries globally and 8% at the coarsest resolution in South Africa. Homogenization increased with increasing resolution and with time, but it decreased with increasing percentage change in species richness. Globally, introductions contributed more to homogenization than did extinctions. Within South Africa, extralimital introductions contributed more to the homogenization of ungulate assemblages than did extraregional ones, and ungulates were typically introduced to high‐income areas with high human population and livestock densities. The same was not true in the past, when ungulates were introduced to ungulate species‐poor areas. The number of non‐indigenous ungulate species established in a country is significantly related to the number of non‐indigenous ungulate species in zoos in the country, possibly owing to sales of surplus animals from zoos. Main conclusions Ungulate faunas are homogenized at both the global scale and in South Africa, with extralimital introductions being of considerable significance regionally. In consequence, increasing attention will have to be given to the conservation consequences of ungulate translocations, both within particular geopolitical regions and across the globe.     

Null model of biotic homogenization: a test with the European freshwater fish fauna

In recent years, there has been growing concern about how species invasions and extinctions could change the distinctiveness of formerly disparate fauna and flora, a process called biotic homogenization. In the present study, a null model of biotic of homogenization was developed and applied to the European freshwater fish fauna. We found that non-native fish species led to the greatest homogenization in south-western Europe and greatest differentiation in north-eastern Europe. Comparing these observed patterns to those expected by our null model empirically demonstrated that biotic homogenization is a non-random ecological pattern, providing evidence for previous assumptions. The place of origin of non-native species was also considered by distinguishing between exotic (originating from outside Europe) and translocated species (originating from within Europe). We showed that exotic and translocated species generated distinct geographical patterns of biotic homogenization across Europe because of their contrasting effects on the changes in community similarity among river basins. Translocated species promoted homogenization among basins, whereas exotic species tended to decrease their compositional similarity. Quantifying the individual effect of exotic and translocated species is therefore an absolute prerequisite to accurately assess the spatial dynamics of biotic homogenization.     

BIODIVERSITY RESEARCH: Native‐exotic species richness relationships across spatial scales and biotic homogenization in wetland plant communities of Illinois,USA

Aim To examine native‐exotic species richness relationships across spatial scales and corresponding biotic homogenization in wetland plant communities. Location Illinois, USA. Methods We analysed the native‐exotic species richness relationship for vascular plants at three spatial scales (small, 0.25 m² of sample area; medium, 1 m² of sample area; large, 5 m² of sample area) in 103 wetlands across Illinois. At each scale, Spearman’s correlation coefficient between native and exotic richness was calculated. We also investigated the potential for biotic homogenization by comparing all species surveyed in a wetland community (from the large sample area) with the species composition in all other wetlands using paired comparisons of their Jaccard’s and Simpson’s similarity indices. Results At large and medium scales, native richness was positively correlated with exotic richness, with the strength of the correlation decreasing from the large to the medium scale; at the smallest scale, the native‐exotic richness correlation was negative. The average value for homogenization indices was 0.096 and 0.168, using Jaccard’s and Simpson’s indices, respectively, indicating that these wetland plant communities have been homogenized because of invasion by exotic species. Main Conclusions Our study demonstrated a clear shift from a positive to a negative native‐exotic species richness relationship from larger to smaller spatial scales. The negative native‐exotic richness relationship that we found is suggested to result from direct biotic interactions (competitive exclusion) between native and exotic species, whereas positive correlations likely reflect the more prominent influence of habitat heterogeneity on richness at larger scales. Our finding of homogenization at the community level extends conclusions from previous studies having found this pattern at much larger spatial scales. Furthermore, these results suggest that even while exhibiting a positive native‐exotic richness relationship, community level biotas can/are still being homogenized because of exotic species invasion.