Distance decay of similarity among European urban floras: the impact of anthropogenic activities on β diversity

Aim We examine how two categories of non‐native species (archaeophyte and neophyte, introduced before and after ad 1500, respectively) have had different impacts on β diversity across European urban floras. Our goal is to use the unique biological perspective provided by urban areas, and the contrasting historical and geographical perspectives provided by archaeophytes and neophytes, to infer how non‐native species will impact upon β diversity in the future. Location Twenty‐two urban areas located in seven European countries. Methods We used the β‐sim dissimilarity index to estimate the level of β diversity for 231 unique pair‐wise combinations of 22 urban floras. We examined bivariate plots of dissimilarity by geographical separation of city centres to evaluate distance decay of similarity for native species, archaeophytes and neophytes. Results Based on average percentages, 52.8% (SD = 8.2%) of species in the urban floras were identified as non‐native with 28.3% (SD = 6.9%) classified as neophytes and 24.5% (SD = 4.9%) as archaeophytes. Relative to native species, across urban floras, archaeophytes were associated with higher compositional similarity and weaker distance decay patterns, whereas neophytes were associated with lower compositional similarity and stronger distance decay patterns. Main conclusions Across European urban floras, archaeophytes and neophytes occurred in similar numbers but archaeophytes were consistently associated with lower β diversity and neophytes with higher β diversity. Thus, the impact of non‐native species on β diversity can be determined, at least in part, through their historical and geographical associations with anthropogenic activities. If archaeophytes represent the long‐term biogeographical outcome for human commensal species, neophytes could develop similar patterns. The consequences, however, are likely to be more substantial ecologically and geographically due to the increasing numbers of neophytes and their global anthropogenic associations. Nevertheless, at present, our findings suggest that, based on occurrence information, neophytes have not achieved this state with European urban floras retaining regionally distinct assemblages of neophytes.     

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.     

Native and alien floras in urban habitats: a comparison across 32 cities of central Europe

Aim To determine relative effects of habitat type, climate and spatial pattern on species richness and composition of native and alien plant assemblages in central European cities. Location Central Europe, Belgium and the Netherlands. Methods The diversity of native and alien flora was analysed in 32 cities. In each city, plant species were recorded in seven 1‐ha plots that represented seven urban habitat types with specific disturbance regimes. Plants were classified into native species, archaeophytes (introduced before ad 1500) and neophytes (introduced later). Two sets of explanatory variables were obtained for each city: climatic data and all‐scale spatial variables generated by analysis of principal coordinates of neighbour matrices. For each group of species, the effect of habitat type, climate and spatial variables on variation in species composition was determined by variation partitioning. Responses of individual plant species to climatic variables were tested using a set of binomial regression models. Effects of climatic variables on the proportion of alien species were determined by linear regression. Results In all cities, 562 native plant species, 188 archaeophytes and 386 neophytes were recorded. Proportions of alien species varied among urban habitats. The proportion of native species decreased with increasing range and mean annual temperature, and increased with increasing precipitation. In contrast, proportions of archaeophytes and neophytes increased with mean annual temperature. However, spatial pattern explained a larger proportion of variation in species composition of the urban flora than climate. Archaeophytes were more uniformly distributed across the studied cities than the native species and neophytes. Urban habitats rich in native species also tended to be rich in archaeophytes and neophytes. Main conclusions Species richness and composition of central European urban floras are significantly affected by urban habitat types, climate and spatial pattern. Native species, archaeophytes and neophytes differ in their response to these factors.     

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.     

Species richness and phylogenetic diversity of native and non‐native species respond differently to area and environmental factors

Aim

To test whether native and non‐native species have similar diversity–area relationships (species–area relationships [SARs] and phylogenetic diversity–area relationships [PDARs]) and whether they respond similarly to environmental variables.

Location

United States.

Methods

Using lists of native and non‐native species as well as environmental variables for >250 US national parks, we compared SARs and PDARs of native and non‐native species to test whether they respond similarly to environmental conditions. We then used multiple regressions involving climate, land cover and anthropogenic variables to further explore underlying predictors of diversity for plants and birds in US national parks.

Results

Native and non‐native species had different slopes for SARs and PDARs, with significantly higher slopes for native species. Corroborating this pattern, multiple regressions showed that native and non‐native diversity of plants and birds responded differently to a greater number of environmental variables than expected by chance. For native species richness, park area and longitude were the most important variables while the number of park visitors, temperature and the percentage of natural area were among the most important ones for non‐native species richness. Interestingly, the most important predictor of native and non‐native plant phylogenetic diversity, temperature, had positive effects on non‐native plants but negative effects on natives.

Main conclusions

SARs, PDARs and multiple regressions all suggest that native and non‐native plants and birds responded differently to environmental factors that influence their diversity. The agreement between diversity–area relationships and multiple regressions with environmental variables suggests that SARs and PDARs can be both used as quick proxies of overall responses of species to environmental conditions. However, more importantly, our results suggest that global change will have different effects on native and non‐native species, making it inappropriate to apply the large body of knowledge on native species to understand patterns of community assembly of non‐native species.

     

Compositional similarity and the distribution of geographical range size for assemblages of native and non-native species in urban floras

There is evidence that, within a region, non-native species introduced from nearby sources (extralimital native) promote homogenization and non-native species introduced from distant sources (exotic) promote differentiation of species composition. A possible explanation for these associations is that they are related to differences in the distribution of geographical range size. We test this by examining geographical ranges, delineated within a defined region, for assemblages of vascular plants in eight urban floras in the USA. Across floras, native species had the largest, least variable ranges and the greatest proportion of shared species. Exotic species had the most variable ranges with concentrations of species with small and large ranges and the lowest proportion of shared species. Extralimital natives had concentrations of species with intermediate-sized ranges and intermediate proportions of shared species. These results suggest that patterns of compositional similarity were associated with the relative strength and equality of two opposing patterns within species range size distributions: species with small vs. large ranges. In general, concentrations of species with small ranges promoted low levels and concentrations of species with large ranges promoted high levels of compositional similarity. However, patterns documented for exotic species will likely continue to develop, possibly taking on new forms, depending on how geographical distributions and the rate of introductions of exotic species change over time. Our findings also suggest that processes underlying these patterns have operated at two spatiotemporal scales. The first scale reflects historical consequences of anthropogenic activities occurring within regional extents that have promoted the introduction of extralimital natives; the second scale reflects modern consequences of anthropogenic activities operating at an increasingly global extent that have promoted the introduction of exotic species.     

The role of non‐native plants and vertebrates in defining patterns of compositional dissimilarity within and across continents

Aim Human activities have led to the spread and establishment of increasing numbers of non‐native species. Here we assess whether non‐native plant and vertebrate species have affected species compositions within and across Europe and North America. We also assess the effects of intra‐continental species exchange using the example of vertebrates. Location European countries and North America (states in the contiguous United States and provinces of Canada). Methods We measured compositional dissimilarity of native and non‐native assemblages of vascular plants and vertebrates and related these patterns to climatic dissimilarity and geographical distance. We considered three categories of non‐native species (introduced after ad 1500), namely: those (1) originating outside of both continents, (2) native to one continent and non‐native to the other, and (3) native in a particular region of a continent but non‐native in another region. Results The presence of non‐native plants and vertebrates led to more homogeneous species compositions between continents and to less homogeneous species composition within Europe compared with the native assemblages. In North America, the presence of non‐native plants led to more homogeneous species compositions and the presence of non‐native vertebrates had no effect. Species compositions being more homogeneous than the native composition were found for the three categories of non‐native vertebrate species for both continents. Between continents, climate was a better predictor of compositional dissimilarity for non‐native plants, whereas for vertebrates the explanatory power of climate and geographical distance were comparable. By contrast, within continents, climate was a better predictor of compositional dissimilarity of both plants and vertebrates. Conclusions We found clear evidence for biotic homogenization as a consequence of species displacement. However, in relation to overall species richness this effect was rather small, indicating that floras and faunas are still quite distinct. Therefore, claiming that we already face homogeneous biotas might be premature, although clear indications are visible which should raise a note of caution, especially in the light of increasing globalization.     

Predicting and explaining plant invasions through analysis of source area floras: some critical considerations

Comparing species that become invasive with others from the same regional species pool that do not invade raises several issues about the accuracy of analyses attempting to define the determinants of invasiveness. The delimitation of the source area and deciding which species group(s) to include are especially relevant in analyses focusing on species originating in Europe. Historical patterns of immigration of alien species into Europe must be considered since European floras comprise a complex mix of native species, historical introductions (archaeophytes) and relative newcomers (neophytes). We make three main points: (1) Archaeophytes (species introduced to Europe before the discovery of America) differ from natives in a number of traits and in historical association with people; it is misleading to lump archaeophytes with native taxa. (2) Taxa from climatically and geographically different regions, representing distinct floristic geoelements, need to be treated separately, and not as a homogenous pool of potential invaders. Restricting the source species pool to native taxa with comparable phytogeographical characteristics reduces the variation associated with chance of dispersal by humans from the source area. (3) For prediction, a clear distinction should be made between accuracy (the proportion of those found to be alien that were also predicted to be there) and reliability (or predictive value, the proportion of those predicted to become aliens that do so). Information accumulated over centuries by botanists in Central Europe provides an excellent opportunity to deal with these issues and avoid spurious results. To illustrate these issues, we revisit a recently published study of Central‐European plant species as invaders in two Argentinean provinces ( Prinzing et al., 2002 ) to explore and demonstrate the implications of the above points. We hope that future studies will build on these points to achieve more reliable predictions.     

The distribution of range sizes of native and alien plants in four European countries and the effects of residence time

Aim Do the statistical distributions of range sizes of native and alien species differ? If so, is this because of residence time effects? And can such effects indicate an average time to a maximum? Location Ireland, Britain, Germany and the Czech Republic. Methods The data are presence or absence of higher plants in mapping units of 100 km² (Ireland and Britain) or c. 130 km² (Germany and the Czech Republic) in areas varying from 79 to 357 thousand km². Logit transforms of range sizes so defined were tested for normality, and examined by ANOVA, and by loess, ordinary least square (OLS) and reduced major axis regressions. Results Current range sizes, in logits, are near normally distributed. Those of native plants are larger than those of naturalized neophytes (plants introduced since 1500 ad ) and much larger than those of casual neophytes. Archaeophytes (introduced earlier) have range sizes slightly larger than natives, except in Ireland. Residence time, the time since an invasive species arrived in the wild at a certain place, affects range sizes. The relationships of the range of naturalized neophytes to residence time are effectively straight in all four places, showing no significant curvature or asymptote back to 1500, though there are few records between 1500 and 1800. The relationships have an r² of only about 10%. Both OLS regressions and reduced major axes can be used to estimate the time it takes for the range of a naturalized neophyte to reach a maximum. Main conclusions Established neophytes have smaller range size distributions than natives probably because many have not yet reached their maximum. We estimate it takes at least 150 years, possibly twice that, on average, for the maximum to be reached in areas of the order of 10⁵ km². Policy needs to allow for the variation in rates of spread and particularly the long time needed to fill ranges. Most naturalized neophytes are still expanding their ranges in Europe.     

Co‐varying impacts of land use and non‐native brown trout on fish communities in small streams

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Retention of gene diversity during the spread of a non‐native plant species

Spatial expansion, which is a crucial stage in the process to successful biological invasion, is anticipated to profoundly affect the magnitude and spatial distribution of genetic diversity in novel colonized areas. Here, we show that, contrasting common expectations, Pyrenean rocket (Sisymbrium austriacum), retained SNP diversity as this introduced plant species descended in the Meuse River Basin. Allele frequencies did not mirror between‐population distances along the predominant expansion axis. Reconstruction of invasion history based on the genotypes of historical herbarium specimens indicated no influence of additional introductions or multiple points of entry on this nongradual pattern. Assignment analysis suggested the admixture of distant upstream sources in recently founded downstream populations. River dynamics seem to have facilitated occasional long‐distance dispersal which brought diversity to the expansion front and so maintained evolutionary potential. Our findings highlight the merit of a historical framework in interpreting extant patterns of genetic diversity in introduced species and underscore the need to integrate long‐distance dispersal events in theoretical work on the genetic consequences of range expansion.