Invasiveness and homogenization: synergism of wide dispersal and high local abundance

Aim To determine whether invasive and locally abundant non‐native species have a more homogenizing effect on plant communities than non‐invasive and less abundant non‐native species. Location California and Florida counties, conservation areas in the USA, and eight US cities. Methods Species lists among counties, conservation areas and cities were compared to see whether invasive and abundant non‐native species increased the Jaccard index of similarity between localities beyond any increases caused by non‐invasive and less abundant non‐native species. Results For all comparisons, we found that invasive non‐native species have a significantly greater homogenizing effect than non‐invasive non‐native species. For the US conservation areas, we found that locally abundant invasive species tend to be more widespread and more widely shared than less abundant invasive species. There is also a positive relationship between homogenization by invasive species and the magnitude of human disturbance. Main conclusions Invasive non‐native species tend to be disproportionately shared among communities relative to non‐invasive non‐native species. This effect is enhanced by human disturbance, as measured by the ratio of non‐native to native species. There is a synergism between abundance and geographical range which enhances the homogenizing effects of abundant species. Invasive species, with wide ecological niches, are more widely shared among communities and more locally abundant. Abundant invasive species are thus more spatially homogenizing, and more ecologically dominant (functionally homogenizing). Also, ‘perceived homogenization’ is probably greater than homogenization measured by the increase in shared species. The abundant species typically seen by the casual observer in a biological community are probably more commonly shared between communities than less common species. Studies that lack abundance data and measure homogenization only on the basis of shared species, which includes most homogenization studies to date, probably underestimate the homogenizing impacts of non‐native species as perceived by people.     

Do Exotics Homogenize or Differentiate Communities? Roles of Sampling and Exotic Species Richness

It is commonly asserted that exotic species promote the homogenization of biological communities. However, theoretical evidence implies that exotic species may often have the opposite effect, of differentiating biological communities where different exotic species become established in different areas. Furthermore, few empirical studies have directly measured the homogenizing effects of exotics. In this study, I used the plant inventories of 20 localities in the United States to measure whether exotic plant increased the similarity of those localities. I calculated Jaccard's index of similarity (JI) for exotic species and then for native species to see if the exotic JI was consistently higher when comparing two localities. I found that JI for both exotic and native species decline exponentially with increasing distance and latitude separation between localities so that localities that share many native species also tend to share many exotic species. More importantly, in nearly half of the pairwise locality comparisons (87 of 190), the (JI_exotic/JI_native) ratio is less than one, indicating a slight tendency for exotic species to differentiate rather than homogenize the localities analyzed. Also, the pattern of differentiation versus homogenization is strongly related to exotic species richness. When both localities have relatively few exotics, there is a clear tendency for exotics to produce biotic differentiation (JI_exotic/JI_native < 1). Much of this pattern seems related to the right-skewed frequency distribution of the geographic ranges of exotic species. As with native species, most exotics occur in few localities so there is a high probability that localities with low numbers of exotic species will share very few, if any, of those exotics. As exotic richness increases, the homogenizing effects become increasingly pronounced.     

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.     

Floristic homogenization by native ruderal and alien plants in north-east Spain: the effect of environmental differences on a regional scale

Aim To evaluate the relative potential contribution of native ruderals and aliens to plant homogenization at a regional scale, after taking into account the effect of diverse environmental distances. Location Catalonia (north‐east Spain) Methods We have used the flora module of the BDBC project (Catalonian Database of Biodiversity), which provides information on plant species distribution per 10 × 10 km Universal Transverse Mercator (UTM) cell. Pairwise floristic similarities of: (1) total, (2) native non‐ruderal, (3) native ruderal, and (4) alien vascular plant species have been calculated for a particularly well‐sampled subset of UTM cells, using a modified version of the Simpson index. These similarities have been compared per UTM pair using Mantel tests, before and after considering their relative association with geographical, climatic and landscape distances from linear regression models. The floristic similarity of both total and native non‐ruderal species was also correlated with the proportion of alien and native ruderal species after discounting the effects of environmental distances. Results The proportion of variance explained by environmental correlates was highest for the floristic similarity of native non‐ruderal plants and lowest for that of aliens. In all plant groups, climatic distance was the main significant variable of species similarity. Geographical distance was only significant for total and native non‐ruderal species and was of secondary importance in both cases. Landscape distance was not significant in any case. Similarities among both aliens and native ruderals were significantly higher than among native non‐ruderals, but these differences disappeared after removing the effect of environmental distances. Main conclusions Species similarity between sites may depend on differences in environmental factors other than geographical distance. This has to be taken into account when exploring the implications for biotic homogenization. In the case of Catalonian flora, the potentially homogenizing effect of native ruderal and alien species seems to be associated with their lower dependence on geographical distance and climatic factors compared with those of native, non‐ruderal species.     

Processes at multiple scales affect richness and similarity of non‐native plant species in mountains around the world

Aim To investigate how species richness and similarity of non‐native plants varies along gradients of elevation and human disturbance. Location Eight mountain regions on four continents and two oceanic islands. Methods We compared the distribution of non‐native plant species along roads in eight mountainous regions. Within each region, abundance of plant species was recorded at 41–84 sites along elevational gradients using 100‐m² plots located 0, 25 and 75 m from roadsides. We used mixed‐effects models to examine how local variation in species richness and similarity were affected by processes at three scales: among regions (global), along elevational gradients (regional) and with distance from the road (local). We used model selection and information criteria to choose best‐fit models of species richness along elevational gradients. We performed a hierarchical clustering of similarity to investigate human‐related factors and environmental filtering as potential drivers at the global scale. Results Species richness and similarity of non‐native plant species along elevational gradients were strongly influenced by factors operating at scales ranging from 100 m to 1000s of km. Non‐native species richness was highest in the New World regions, reflecting the effects of colonization from Europe. Similarity among regions was low and due mainly to certain Eurasian species, mostly native to temperate Europe, occurring in all New World regions. Elevation and distance from the road explained little of the variation in similarity. The elevational distribution of non‐native species richness varied, but was always greatest in the lower third of the range. In all regions, non‐native species richness declined away from roadsides. In three regions, this decline was steeper at higher elevations, and there was an interaction between distance and elevation. Main conclusions Because non‐native plant species are affected by processes operating at global, regional and local scales, a multi‐scale perspective is needed to understand their patterns of distribution. The processes involved include global dispersal, filtering along elevational gradients and differential establishment with distance from roadsides.     

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.     

Non‐native plants have greater impacts because of differing per‐capita effects and nonlinear abundance–impact curves

Invasive, non‐native species can have tremendous impacts on biotic communities, where they reduce the abundance and diversity of local species. However, it remains unclear whether impacts of non‐native species arise from their high abundance or whether each non‐native individual has a disproportionate impact – that is, a higher per‐capita effect – on co‐occurring species compared to impacts by native species. Using a long‐term study of wetlands, we asked how temporal variation in dominant native and non‐native plants impacted the abundance and richness of other plants in the recipient community. Non‐native plants reached higher abundances than natives and had greater per‐capita effects. The abundance–impact relationship between plant abundance and richness was nonlinear. Compared with increasing native abundance, increasing non‐native abundance was associated with steeper declines in richness because of greater per‐capita effects and nonlinearities in the abundance–impact relationship. Our study supports eco‐evolutionary novelty of non‐natives as a driver of their outsized impacts on communities.     

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.     

Long‐term dynamics of bird diversity in forest and suburb: decay,turnover or homogenization?

Aim Urbanization and deforestation are important drivers of biodiversity change. However, long‐term changes in faunal communities within urbanizing regions are poorly understood. We investigated how well observed community changes in both space and time agree with expectations based on current paradigms in urban ecology. Location Greater Brisbane region, Australia. Methods We compared bird assemblages in two time‐periods 15 years apart, at multiple sites in remnant forest and residential suburbs across an urbanizing landscape. Differences in assemblage composition, species abundances and functional groupings were assessed within and between habitats. Results Compared with forest, suburbs in both time‐periods had over twice the total bird abundance, a different species composition, greater between‐site community similarity, a greater proportion of non‐native species and greater dominance by large‐bodied species. These differences corresponded with changes in sites whose habitat was converted from forest to suburb. Between time‐periods, abundances of 58% of suburban species changed significantly compared with those of 11% in forest. Increaser species outnumbered decreasers in suburbs, with the reverse in forest. Abundance of small‐bodied birds decreased 70% in suburbs and 20% in forest. Broad‐spectrum competitors and nest predators were common among suburban increasers. Among invasive species, the number of increasers was counterbalanced by decreasers. Both site‐scale species richness and between‐site community similarity increased to a small extent in both habitats. Main conclusions Species composition and ecological function of suburban bird communities were very dynamic. Suburban assemblages were neither a subset of forest species nor an increasingly non‐native compilation. Communities in large forest patches were comparatively stable. The notion of habitat‐specific species turnover better characterizes the nature of most changes than either species decline or homogenization, even though both of these were evident. There is considerable scope for careful urban planning, focused on both among‐ and within‐habitat variety, to sustain bird diversity in urbanizing landscapes.     

Compositional similarity among urban floras within and across continents: biogeographical consequences of human-mediated biotic interchange

Anthropogenic activities have weakened biogeographical barriers to dispersal resulting in the global spread and establishment of an increasing number of non‐native species. We examine the broad‐scale consequences of this phenomenon based on an analysis of compositional similarity across urban floras in the northeastern United States and Europe. We test the prediction that homogenization of species composition is uniquely defined within vs. between continents based on the time and place of origin of non‐native species. In this case, for archaeophytes and neophytes in Europe (introduced before and after ad 1500, respectively) and non‐native species originating from within and outside the United States. More species in urban floras were shared within than between continents. Within Europe, archaeophytes shared more species across urban floras compared with neophytes; strong associations were not observed for non‐native species across US urban floras. Between the two continents, non‐native species in the United States that originated from outside the United States shared species primarily with archaeophytes but also with European natives and neophytes. These results suggest that the direction of biotic interchange was unidirectional with species moving primarily from Europe to the United States with archaeophytes playing a primary and non‐native species originating from outside the two continents a secondary role as a homogenizing source. Archaeophytes, based on combination of biogeographical, evolutionary, and ecological factors in association with a long history of anthropogenic influence, appear to have played a prominent role in the continental and intercontinental homogenization of species composition. This suggests that the uniform homogenization of the Earth's biota is not imminent and is presently directed by a combination of biogeographically defined anthropogenic and historical factors.     

Species composition,functional and phylogenetic distances correlate with success of invasive Chromolaena odorata in an experimental test

Biotic resistance may influence invasion success; however, the relative roles of species richness, functional or phylogenetic distance in predicting invasion success are not fully understood. We used biomass fraction of Chromolaena odorata, an invasive species in tropical and subtropical areas, as a measure of ‘invasion success’ in a series of artificial communities varying in species richness. Communities were constructed using species from Mexico (native range) or China (non‐native range). We found strong evidence of biotic resistance: species richness and community biomass were negatively related with invasion success; invader biomass was greater in plant communities from China than from Mexico. Harvesting time had a greater effect on invasion success in plant communities from China than on those from Mexico. Functional and phylogenetic distances both correlated with invasion success and more functionally distant communities were more easily invaded. The effects of plant‐soil fungi and plant allelochemical interactions on invasion success were species‐specific.     

The role of exotic species in homogenizing the North American flora

Exotic species have begun to homogenize the global biota, yet few data are available to assess the extent of this process or factors that constrain its advance at global or continental scales. We evaluate homogenization of vascular plants across America north of Mexico by comparing similarity in the complete native and exotic floras between states and provinces of the USA and Canada. Compared with native species, exotic plants are distributed haphazardly among areas but spread more widely, producing differentiation of floras among neighbouring areas but homogenization at greater distance. The number of exotic species is more closely associated with the size of the human population than with ecological conditions, as in the case of native species, and their distributions are less influenced by climate than those of native species.     

Effects of introduced species on floristic similarity: Comparing two US states

This study aims to examine the effects of introduced species on increasing (homogenizing) or decreasing (differentiating) floristic similarity of plant composition. We calculated the Jaccard index for each pair of counties within two states of USA, California and Florida. We computed the Jaccard index separately for all (native plus exotic) species, for native species, and for exotic species. We further calculated a homogenization index between all species and native species for each pair of counties by subtracting similarity index for native species from that for all species. We correlated the Jaccard and homogenization indices to geographic distance, latitude separation, and longitude separation between pairs of counties and to average human population density and average land area of the two counties. We find a very strong pattern of differentiation for introduced species among nearly all Florida counties. In California, introduced species have a differentiating effect in about half the comparisons. We also find that introduced species tend to have a more homogenizing (or less differentiating) effect with increasing distances between counties. Our results do not show a clear relationship between human population density and the homogenization index.     

Biotic homogenization of three insect groups due to urbanization

Cities are growing rapidly, thereby expected to cause a large‐scale global biotic homogenization. Evidence for the homogenization hypothesis is mostly derived from plants and birds, whereas arthropods have so far been neglected. Here, I tested the homogenization hypothesis with three insect indicator groups, namely true bugs, leafhoppers, and beetles. In particular, I was interested whether insect species community composition differs between urban and rural areas, whether they are more similar between cities than between rural areas, and whether the found pattern is explained by true species turnover, species diversity gradients and geographic distance, by non‐native or specialist species, respectively. I analyzed insect species communities sampled on birch trees in a total of six Swiss cities and six rural areas nearby. In all indicator groups, urban and rural community composition was significantly dissimilar due to native species turnover. Further, for bug and leafhopper communities, I found evidence for large‐scale homogenization due to urbanization, which was driven by reduced species turnover of specialist species in cities. Species turnover of beetle communities was similar between cities and rural areas. Interestingly, when specialist species of beetles were excluded from the analyses, cities were more dissimilar than rural areas, suggesting biotic differentiation of beetle communities in cities. Non‐native species did not affect species turnover of the insect groups. However, given non‐native arthropod species are increasing rapidly, their homogenizing effect might be detected more often in future. Overall, the results show that urbanization has a negative large‐scale impact on the diversity specialist species of the investigated insect groups. Specific measures in cities targeted at increasing the persistence of specialist species typical for the respective biogeographic region could help to stop the loss of biodiversity.     

Impacts of a simulated heat wave on composition of a marine community

Extreme events, such as heat waves, are predicted to increase in frequency, duration, and severity as a consequence of climate change. However, global change research generally focuses on increases in mean temperatures and fails to address the potential impacts of increasingly severe heat waves. In addition, climate change may interact with another primary threat to biodiversity, non‐native species invasions. We assessed the impacts of a short‐term heat wave on the marine epibenthic fouling community of Bodega Harbor, California, USA, by exposing experimental mesocosms to a simulated heat wave in the laboratory and then monitoring community development in the field. We hypothesized that (1) juveniles would be more susceptible to heat waves than adults, (2) native species would be more susceptible than non‐native species, and (3) non‐native species would recover more quickly than native species. We observed no effect of the heat wave on juvenile species richness, either initially or during the recovery period, relative to communities at ambient seawater temperatures. In contrast, total adult species richness initially declined in response to the heat wave. Adult community composition also changed in heat‐wave treatments, with non‐natives representing the majority of species and occupying more cover than native species. The reduction in native richness associated with the heat wave persisted through the recovery period, whereas invasive richness was actually higher on heat‐wave versus ambient plates at 95 days. Heat waves have the potential to alter the composition of this community because of species‐, taxon‐, and/or origin‐specific responses; for example, non‐native bryozoans displayed greater resistance than native and non‐native tunicates. Recovery from the heat wave occurred via growth of resistant individuals and larval recruitment. Our study highlights the importance of considering species’ and community responses to heat waves, and not just mean predicted temperature increases, to evaluate the consequences of climate change.     

Phylogenetic patterns differ for native and exotic plant communities across a richness gradient in Northern California

Aim Increasingly, ecologists are using evolutionary relationships to infer the mechanisms of community assembly. However, modern communities are being invaded by non‐indigenous species. Since natives have been associated with one another through evolutionary time, the forces promoting character and niche divergence should be high. On the other hand, exotics have evolved elsewhere, meaning that conserved traits may be more important in their new ranges. Thus, co‐occurrence over sufficient time‐scales for reciprocal evolution may alter how phylogenetic relationships influence assembly. Here, we examined the phylogenetic structure of native and exotic plant communities across a large‐scale gradient in species richness and asked whether local assemblages are composed of more or less closely related natives and exotics and whether phylogenetic turnover among plots and among sites across this gradient is driven by turnover in close or distant relatives differentially for natives and exotics. Location Central and northern California, USA. Methods We used data from 30 to 50 replicate plots at four sites and constructed a maximum likelihood molecular phylogeny using the genes: matK, rbcl, ITS1 and 5.8s. We compared community‐level measures of native and exotic phylogenetic diversity and among‐plot phylobetadiversity. Results There were few exotic clades, but they tended to be widespread. Exotic species were phylogenetically clustered within communities and showed low phylogenetic turnover among communities. In contrast, the more species‐rich native communities showed higher phylogenetic dispersion and turnover among sites. Main conclusions The assembly of native and exotic subcommunities appears to reflect the evolutionary histories of these species and suggests that shared traits drive exotic patterns while evolutionary differentiation drives native assembly. Current invasions appear to be causing phylogenetic homogenization at regional scales.     

Toward a mechanistic understanding and prediction of biotic homogenization

The widespread replacement of native species with cosmopolitan, nonnative species is homogenizing the global fauna and flora. While the empirical study of biotic homogenization is substantial and growing, theoretical aspects have yet to be explored. Consequently, the breadth of possible ecological mechanisms that can shape current and future patterns and rates of homogenization remain largely unknown. Here, we develop a conceptual model that describes 14 potential scenarios by which species invasions and/or extinctions can lead to various trajectories of biotic homogenization (increased community similarity) or differentiation (decreased community similarity); we then use a simulation approach to explore the model's predictions. We found changes in community similarity to vary with the type and number of nonnative and native species, the historical degree of similarity among the communities, and, to a lesser degree, the richness of the recipient communities. Homogenization is greatest when similar species invade communities, causing either no extinction or differential extinction of native species. The model predictions are consistent with current empirical data for fish, bird, and plant communities and therefore may represent the dominant mechanisms of contemporary homogenization. We present a unifying model illustrating how the balance between invading and extinct species dictates the outcome of biotic homogenization. We conclude by discussing a number of critical but largely unrecognized issues that bear on the empirical study of biotic homogenization, including the importance of spatial scale, temporal scale, and data resolution. We argue that the study of biotic homogenization needs to be placed in a more mechanistic and predictive framework in order for studies to provide adequate guidance in conservation efforts to maintain regional distinctness of the global biota.     

Reservoirs promote the taxonomic homogenization of fish communities within river basins

Most studies analyzing patterns in biotic homogenization of fish communities have used large-scale approaches, while the community-level effects of species introductions and local extinctions within river basins have been sparsely analyzed. In this article, we examine patterns in freshwater fish α- and β-diversity in relation to the presence of reservoirs in a Mediterranean river (Guadiana river; Iberian Peninsula). We used fish samples from 182 river localities and 59 reservoir ones to address two main questions: (i) do reservoirs favor the establishment of invasive fish species?; and (ii) do reservoirs bear taxonomically homogenized fish communities? Although total species richness was not different between rivers and reservoirs, the latter had more invasive species and less native ones. Fish species found in reservoirs tended to be larger ones, but invasive species of any size showed higher preferences for reservoirs. Native species that were rare or absent in reservoirs were those that showed higher sensitivity to invasive species in rivers. Reservoir fish communities were taxonomically homogenized in relation to river ones, both when considering all fish species and using only natives or only invasive ones. Our results suggest that invasive species occupying reservoirs constitute an ecological filter excluding most native species from such systems. Invasive species in the study area are often widely introduced elsewhere, while native species found in reservoirs are congeneric and ecologically similar to those found in other Iberian studies. Thus, we conclude that reservoirs promote taxonomic homogenization at multiple spatial scales, while could also be promoting the functional homogenization of Iberian fish communities.     

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.