Neophyte Invasion in Moldavia (Eastern Romania) in Different Habitat Types

The actual state of neophyte invasion in Moldavia (Eastern Romania) is described in this paper on the basis of 11,055 phytosociological relevés. We analyzed the i) proportion of relevés with neophytes, ii) mean proportion of neophytes per relevés, and iii) mean coverage of neophytes per relevé for 36 EUNIS habitat types to identify general plant invasion patterns. The level of invasion differed considerably between habitats. The invasion of neophytes especially affected habitats strongly determined or influenced by man, such as anthropogenic woodlands, ruderal habitats, arable lands or trampled areas. Most natural habitats are either slightly invaded or entirely free of neophytes. Only riverine willow stands and wet tall-herb stands are relatively highly invaded. However, the absence of neophytes in some natural habitats less represented in the phytosociological dataset could be of artifactual nature. No significant relationship between the number of neophytes and non-neophytes was found in the analysis across different habitats. When the analyses were made within-habitats, both negative and positive relationships were found, which confirm that the relationship between alien and native species richness depends on the habitats. A total number of 105 neophyte species were recorded in the phytosociological relevés used in this study. Among these, 13 species that are currently considered invasive in Moldavia occur in at least 10 types of habitats.     

Altitudinal distribution of native and alien plant species in roadside communities from central Argentina

Roadsides may homogenize the distribution of native species and act as corridors for the spread of alien taxa. We examined the variation in native and alien plant species richness and composition at two spatial scales defined by altitude and habitat type (edges and fill slopes), as well as the relationship between native and exotic species richness in roadside plant communities in mountains from central Argentina. Following a gradient from 1100 to 2200 m a.s.l. along a mountain road, plant species cover was recorded within sample plots of 30 m × 10 m systematically located at 100‐m altitude intervals on both roadside habitats. Although native species richness decreased with altitude and composition changed accordingly, the number of alien species peaked at both extremes of the elevation gradient and did not reflect an altitudinal replacement of chorological groups. The number of both native and alien species was higher in roadside edges, but a negative association between the richness of native and alien species occurred only on fill slopes, suggesting that roadside habitats differ in their susceptibility to plant species colonization and in the mechanisms driving native and alien species richness. Our results highlight the importance of altitude and roadside habitat as factors controlling plant species richness and composition along roadside communities in central Argentina. Although altitude acts as a filter for native plants, it apparently did not constrain the establishment of alien species along the studied roadsides, indicating that the influence of this road as a plant species corridor may increase with time, promoting the opportunities for aliens to expand their current distribution.     

Landscape context modulates alien plant invasion in Mediterranean forest edges

Natural habitats in human-altered landscapes are especially vulnerable to biological invasions, especially in their edges. We aim to understand the influence of landscape and local characteristics on biological invasions by exploring the level of plant invasion and alien species traits in forest edges in highly urbanized landscapes. We identified all plant species in 73 paired plots in the edge and 50 m towards the interior of the forest. We explored the association between alien species richness and similarity in species composition between edge and interior plots with landscape and local variables, using generalized linear models and variance partitioning techniques. Then, we performed Fourth-corner analyses to explore the association between alien plant traits and local and landscape variables. In contrast to native species richness, alien species richness was more affected by the surrounding landscape than by the local characteristics of the edge. Road proximity was positively associated with alien species richness and proportion and was its most important correlate, whereas disturbance was negatively associated with native species richness and was its most influential factor. Alien plant traits were also primarily associated with landscape characteristics. For instance, species of Mediterranean origin and introduced for agriculture were associated with higher agriculture use in the landscape. Our findings suggest that risk analyses of habitat vulnerability to invasion must consider the landscape context in order to successfully predict highly invaded areas and identify potentially successful invaders.     

Observed and dark diversity of alien plant species in Europe: estimating future invasion risk

Invasion by alien species is a growing concern for nature conservation. We estimated the level of invasion by alien plant species and future invasion risks at the European scale. We used a pan-European atlas and eight regional plant atlases to determine the distribution of alien and native plant richness. In addition, we estimated alien and native dark diversity (species currently absent from a site but present in the surrounding region and able to colonize the site). We used relative diversity metrics to indicate current and future risks by alien species: relative alien richness (compared to native species), alien and native completeness (log-ratio of observed to dark diversity) and completeness difference between alien and native species. Observed and relative richness of alien species were greatest in NW Europe; this suggests that sites in NW Europe could be more disturbed. Observed alien and native species richness show clear regional hotspots; the distribution of completeness values is dispersed, indicating local hotspots. Northern Europe has relatively lower alien completeness, likely because potential invaders inhabit the region but have not yet reached many localities, thereby suggesting a risk of future invasion. A greater number of potential alien species in the region increases the probability that some alien species could have detrimental impacts. Both alien richness and completeness are positively correlated with native richness and completeness, respectively, indicating that both groups share similar distribution patterns. Alien species diversity metrics in Europe are related positively to human population density and agricultural land-use. We suggest that the dark diversity concept can broaden our understanding of alien species diversity and future invasion risks.     

Mixed messages from multiple information sources on invasive species: a case of too much of a good thing?

Aim The increasing number and availability of online databases of alien species beg a question of their comparability given most do not adopt standard criteria in the definition of species status or taxonomic treatment and vary in their comprehensiveness. In this study, we compare the consistency of two major European databases for the regions they have in common. We assess whether they use consistent terminology to classify species status, provide similar taxonomic classification and coverage, deliver comparable estimates of alien richness per country and identify comparable correlates of alien richness. Location Northern Europe. Methods Data on the total number of alien species as well as the number of established alien species were extracted from the online databases DAISIE and NOBANIS for 13 European countries and classified into comparable taxonomic groups. Analyses across countries examined trends in alien species richness, correlations among taxonomic groups and the explanatory power of population density, country area and per capita GDP on alien species richness. Results Alien species richness, intertaxon correlations and the significance of individual drivers of invasion were all strongly database dependent. Differences were more marked for total numbers of aliens than established aliens. Over all taxonomic groups, DAISIE had lower species richness and fewer significant intertaxon correlations but presented a greater number of significant explanatory models of alien species richness. Trends in species richness were not generally correlated between the two databases with human population density being a more important driver in DAISIE while country area had greater explanatory power in NOBANIS. Main conclusions Considerable caution should be applied when collating data from different databases because often their underlying structure and content may differ markedly. For Europe, the analysis indicates that having two contrasting databases is not an ideal basis for implementing invasive species policy and moves should be made soon to establish a central pan‐European database.     

Effects of land use and environment on alien and native macrophytes: lessons from a large‐scale survey of Australian rivers

Aim We sampled riverine macrophyte communities and environmental conditions to compare drivers of alien and native abundance and to provide a general set of environmental correlates of invasion by aquatic macrophytes. Location Streams adjacent to three land‐use types (intensive, agricultural and natural) across a large latitudinal gradient (approximately 27° S–43° S) in Australia. Sites were located near Brisbane (Queensland), Sydney (New South Wales), Canberra (Australian Capital Territory), Melbourne (Victoria) and Hobart (Tasmania). Methods Alien and native aquatic plant species cover, water quality, forest canopy and adjacent land use were measured in three catchment locations (low‐, mid‐ and upper‐catchment) in all cities. Mean richness and cover of native and alien macrophytes were compared in the five cities, three catchment locations, and three land‐use types. Correlation tests examined relationships between alien and native richness at transect, site and city scales. Canonical correspondence analysis (CCA) determined the effects of environment on cover and richness of native and alien plant groups (emergents, floating, forbs/other, graminoids and submerged). Results Variation existed in the aquatic plant community at all scales, but strong patterns emerged with respect to land use and environmental gradients. Alien abundance was more responsive to anthropogenic disturbance (e.g. greater in intensive and agricultural land‐use types, and greater where dissolved nutrients and conductivity were high) than natives, which were unaffected by land‐use type and less responsive overall to environmental gradients. Native and alien richness were uncorrelated at all scales. Main conclusions Natives and aliens of the same life form did not respond similarly to the environment, suggesting inherent differences in their ability to capitalize on anthropogenic disturbance. Our results suggest invasion‐susceptible habitats are those that receive nutrient pollutants and that occur in urban and agricultural areas low in the catchment. Our confidence in these patterns is strengthened by their consistency across a large latitudinal gradient.     

Alien plant species coexist over time with native ones in Chilean Mediterranean grasslands

Aims Alien species are commonly considered as harmful weeds capable of decreasing native biodiversity and threatening ecosystems. Despite this assumption, little is known about the long-term patterns of the native–alien relationships associated with human disturbed managed landscapes. This study aims to elucidate the community dynamics associated with a successional gradient in Chilean Mediterranean grasslands, considering both native and alien species.Methods Species richness (natives and aliens separately) and life-form (annuals and perennials) were recorded in four Chilean post-agricultural grazed grasslands each covering a broad successional gradient (from 1 to 40 years since crop abandonment). A detrended correspondence analysis (DCA), mixed model effects analyses and correlation tests were conducted to assess how this temporal gradient influenced natives and aliens through community dynamics.Important findings Our results show different life-form patterns between natives and aliens over time. Aliens were mainly represented by annuals (especially ruderals and weeds), which were established at the beginning of succession. Annual aliens also predominated at mid-successional stages, but in old grasslands native species were slightly more representative than alien ones within the community. In the late successional states, positive or no correlations at all between alien and native species richness suggested the absence of competition between both species groups, as a result of different strategies in occupation of the space. Community dynamics over time constitute a net gain in biodiversity, increasing natives and maintaining a general alien pool, allowing the coexistence of both. Biotic interactions including facilitation and/or tolerance processes might be occurring in Chilean post-agricultural grasslands, a fact that contradicts the accepted idea of the alien species as contenders.     

Contrasting historical and current land-use correlation with diverse components of current alien plant invasions in Mediterranean habitats

Habitat invasion by alien plants is strongly modulated by environmental and landscape factors. However, the effect of landscape history remains largely unknown, despite the fact that it could play an important role in many stages of invasion processes, even long after land-use changes have occurred determining invasion debts. We analysed the effects of past landscape and recent changes therein, together with habitat type and current context (i.e. climate, topography and landscape), on three components of the invasion process at habitat scale: alien species presence (i.e. at least one alien species occurring), richness (number of species found) and abundance (mean species cover). We selected 531 plots in nine habitat types in Barcelona province (7725 km²) and recorded alien (neophyte) species cover. We performed Generalized Linear Models on these invasion components using the generated data and a set of predictors of habitat, context and landscape factors obtained from plot sampling and digital cartography. The results show that invasion components are affected by diverse habitat and context factors and, in some cases, by landscape history. Alien species presence is influenced by habitat type and the current environmental context, and by the number of habitat changes in the adjacent landscape; on the other hand, species richness is only associated with the current context and species abundance is only influenced by historical cropland cover. The association between alien species presence and abundance and past and recent landscape changes suggests the existence of accumulated invasion debts at habitat scale that might be relevant to habitat management.     

Species richness,alien species and plant traits in Central Argentine mountain grasslands

Abstract. Pattern of native vegetation, distribution of alien species and variation of environmental parameters were studied in mountain grasslands in a lithologically homogeneous Córdoba mountain range in Central Argentina. CCA showed that altitude was the most important factor determining the compositional variation of the vegetation, with soil nutrient status and stoniness as additional factors. Short‐grass communities, associated with the driest habitats on plateaus, showed higher small‐scale native species richness than wet‐turf communities in valleys and tall‐grass communities on slopes. Species richness was negatively correlated with soil parameters that indicate nutrient status and water availability. Also, there was a negative correlation between soil Ca‐ and Mg‐ content and richness. High native species richness coincided with high alien species richness. When smaller units – community types – were considered, it became evident that within short‐grass vegetation, the three most species‐rich community types contained significant numbers of alien species, while the other two did not. Even within one community type, the same quadrats that contained the highest number of native species, were also characterized by the highest numbers of alien species. Evidently, the same mechanism was responsible for high richness of both native and alien species. Alien species were distinguished by a greater proportion of annuals and prostrate stoloniferous plants, by lower palatability and by smaller proportion of zoochory. DCA ordination of quadrats on the basis of plant traits as attributes resulted in a clear distinction of three main vegetation types. Short‐grass vegetation was distinguished by a predomination of late flowering species, tall‐grass vegetation by the presence of high herbaceous plants and bushes, and wet‐turf vegetation by the presence of plants with storage organs, the lack of hairy leaves, and by a high proportion of cryptophytes. Quadrats with and without alien species were distinguished as well, indicating that the occurrence of aliens may be dependent on plant traits in a particular patch of a community.     

Widespread native and alien plant species occupy different habitats

Theories to explain the success of alien species often assume that they are inherently different from native species. Although there is an increasing body of evidence showing that alien plants tend to dominate in highly human‐modified environments, the underlying reasons why widespread natives might differ in their habitat distribution have rarely been addressed. We used species distribution models to quantify the dominant environmental axes shaping the habitat of 95 widespread native and alien herbaceous species in a highly modified grassland‐dominated landscape in New Zealand. For each species, support vector machines were used to determine 1) the environmental variables that most strongly determined a species’ distribution; 2) the affinity towards a particular position along environmental axes; and 3) tolerance to environmental variation. These three measures were compared among native perennials (n = 31), alien perennials (30) and alien annuals (34). Independent of their origin, species’ distributions were defined by similar environmental variables. Nevertheless, native and alien species occupied different regions of the dominant environmental axes. Perennial natives occupied regions associated with lower human disturbance, while perennial aliens were associated with habitats that had been modified by vegetation clearance, pasture development and livestock grazing. Annual aliens differed from perennials and were associated with both semi‐natural and more intensively managed vegetation. No evidence was found that aliens had broader environmental tolerances than natives that might facilitate invasion into a wider range of environments. Thus, widespread native and alien species differ in the degree to which environmental factors shape their distribution as a result of anthropogenic perturbations to which they respond differently as well as the introduction of functional groups that are capable of exploiting novel environments.     

Contrasting response of native and alien plant species richness to environmental energy and human impact along alpine elevation gradients

Aim We tested whether the species–energy and species–human relationships vary between native and both naturalized and casual alien species richness when other environmental variables had been taken into account. Location Trento Province, a region (c. 6200 km²) on the southern border of the European Alps (Italy), subdivided into 156 contiguous (c. 37.5 km²) cells and ranging in elevation from 66 to 3769 m. Methods Data were separated into three subsets, representing richness of natives, naturalized aliens and casual aliens and separately related to temperature, human population and various environmental correlates of plant species diversity. We applied ordinary least squares and simultaneous autoregressive regressions to identify potential contrasting responses of the three plant status subsets and hierarchical partitioning to evaluate the relative importance of the predictor variables. Results Variation in alien plant species richness along the region was almost entirely explained by temperature and human population density. The relationships were positive but strongly curvilinear. Native species richness was less strongly related to either factor but was positively related to the presence of calcareous bedrock. Native species richness had a decelerating positive relationship with temperature (R²= 55%), whereas naturalized and casual aliens had a positive accelerating relationship explaining 86% and 62% of the variation in richness, respectively. Native species richness had a positive decelerating relationship with population density (R²= 42%), whilst both alien subsets had a positive accelerating relationship. Main conclusions Alien species richness was higher in areas with the most rich and diverse assemblages of native species. Areas at high altitudes are not especially prone to alien invasion due to energy constraints, low propagule pressure and disturbance, even considering a potential increased in temperature. Thus, if we consider future environmental change, we should expect a stronger response of aliens than natives in the currently warm, urbanized, low‐altitude areas than in cold, high‐altitude areas where human population density is low.     

Do alien plants on Mediterranean islands tend to invade different niches from native species?

In order to understand invasions, it is important to know how alien species exploit opportunities in unfamiliar ecosystems. For example, are aliens concentrated in niches under-exploited by native communities, or widely distributed across the ecological spectrum? To explore this question, we compared the niches occupied by 394 naturalized alien plants with a representative sample from the native flora of Mediterranean islands. When niche structure was described by a functional group categorization, the distribution of native and alien species was remarkably similar, although “succulent shrubs” and “trees with specialized animal pollination mechanisms” were under-represented in the native species pool. When niche structure was described by Grime’s CSR strategy, the positioning of aliens and natives differed more strongly. Stress-tolerance was much rarer amongst the aliens, and a competitive strategy was more prevalent at the habitat level. This pattern is similar to previous findings in temperate Europe, although in those regions it closely reflects patterns of native diversity. Stressed environments are much more dominant in the Mediterranean. We discuss a number of factors which may contribute to this difference, e.g., competitive and ruderal niches are often associated with anthropogenic habitats, and their high invasibility may be due partly to introduction patterns rather than to a greater efficiency of aliens at exploiting them. Thus far, the reasons for invasion success amongst introduced species have proved difficult to unravel. Despite some differences, our evidence suggests that alien species naturalize across a wide range of niches. Given that their ecologies therefore vary greatly, one may ask why such species should be expected to share predictable traits at all?     

Contrasting patterns in the invasions of European terrestrial and freshwater habitats by alien plants,insects and vertebrates

Aim To provide the first comparative overview on the current numbers of alien species that invade representative European terrestrial and freshwater habitats for a range of taxonomic groups. Location Europe. Methods Numbers of naturalized alien species of plants, insects, herptiles, birds and mammals occurring in 10 habitats defined according to the European Nature Information System (EUNIS) were obtained from 115 regional data sets. Only species introduced after ad 1500 were considered. Data were analysed by ANCOVA and regression trees to assess whether differences exist among taxonomic groups in terms of their habitat affinity, and whether the pattern of occurrence of alien species in European habitats interacts with macroecological factors such as insularity, latitude or area. Results The highest numbers of alien plant and insect species were found in human‐made, urban or cultivated habitats; if controlled for habitat area in the region, wetland and riparian habitats appeared to support relatively high numbers of alien plant species too. Invasions by vertebrates were more evenly distributed among habitats, with aquatic and riparian, woodland and cultivated land most invaded. Mires, bogs and fens, grassland, heathland and scrub were generally less invaded. Habitat and taxonomic group explained most variation in the proportions of alien species occurring in individual habitats related to the total number of alien species in a region, and the basic pattern determined by these factors was fine‐tuned by geographical variables, namely by the mainland–island contrast and latitude, and differed among taxonomic groups. Main conclusions There are two ecologically distinct groups of alien species (plants and insects versus vertebrates) with strikingly different habitat affinities. Invasions by these two contrasting groups are complementary in terms of habitat use, which makes an overall assessment of habitat invasions in Europe possible. Since numbers of naturalized species in habitats are correlated among taxa within these two groups, the data collected for one group of vertebrates, for example, could be used to estimate the habitat‐specific numbers of alien species for other vertebrate groups with reasonable precision, and the same holds true for insects and plants.     

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.     

Reproductive habitat selection in alien and native populations of the genus Discoglossus

The existence of suitable breeding habitats is an important factor explaining the regional presence of an anuran species. This study examined patterns of habitat selection in populations of three species of the genus Discoglossus: Discoglossus galganoi (south-western Iberian Peninsula), Discoglossus scovazzi (Morocco) and Discoglossus pictus (three different areas were included in the study: Sicily, Tunisia and north-eastern Iberian Peninsula). The populations of D. pictus on the Iberian Peninsula are allochthonous, and analysis of these patterns may provide insights into the processes that regulate the invasion phase. The hypotheses tested were: (i) congeneric species show the same patterns of habitat selection, and alien species have been established following these patterns; (ii) there are differences in species associations between assemblages structured deterministically and by chance, i.e. native versus invaded assemblages. The larval habitats of three species of this genus were characterized by measuring physical and chemical parameters of the water bodies. We examined the covariation between the presence of Discoglossus species and the species richness of sympatric anurans, and investigated a possible relationship between morphological similarity (as a proxy of functional group) and overlap in habitat use. The results showed that congeneric species are morphologically conservative and also select very similar types of aquatic habitat. The alien population and other sympatric species showed a high degree of overlap in habitat use, which was greater than that observed in the native assemblage with a similar functional richness. Species associations were not structured on the basis of morphological similarity in any of the assemblages. Among native populations, the presence of Discoglossus was either negatively correlated or not significantly correlated with species richness. Only the alien population showed a positive correlation between its presence and species richness, which suggests a loss of assemblage structure.     

Do alien and native tree species from Central Argentina differ in their water transport strategy?

In this study we assessed the water transport strategies and the abundance of alien and native tree species at a regional scale in Córdoba Mountains, Central Argentina. The aims of this study were: (i) to analyse whether alien and native tree species show divergent water transport strategies; and (ii) to explore whether species abundances of alien and natives are associated with specific trait attributes. Eight alien species and 12 native species were recorded in 50 complete vascular plant vegetation relevés. Water transport strategies were assessed through the following functional traits: minimum leaf water potential, potential water content of wood, effective leaf area, leaf area per sapwood area and wood density. Also, resource use strategies were assessed throughout the measurement of specific leaf area. We found that alien species had a higher efficiency in water transport (i.e. higher minimum leaf water potential and lower wood density values) and faster resource acquisition and use (higher specific leaf area values) than native species. We did not find evidence suggesting that the relative abundance of species was associated to water transport strategies and faster resource acquisition and use. Alien species seem to differ from natives in specific functional attributes that are absent in the resident community and might allow aliens to use more resources and at a higher rate than native species. Finally, our results show the potential of a trait‐based predictive framework for alien species, and the possible effects on ecosystem functions.     

Diversity of wetland vegetation in the Bulgarian high mountains, main gradients and context-dependence of the pH role

We fill a gap in understanding wetland vegetation diversity and relationship with environmental determinants in Bulgarian high mountains. A total of 615 phytosociological samples were taken from springs, mires, wet meadows and tall-forb habitats throughout Bulgaria, of which 234 relevés are from mire and spring vegetation above timberline. The vegetation was classified by TWINSPAN and the resulting vegetation types were reproduced by the formal definitions using the combination of Cocktail species groups based on phi-coefficient of joint co-occurrence of the species. Nine vegetation types of springs and fens have been clearly delimited above the timberline. All vegetation types include Balkan endemic species, the representation of which varies. Fens generally harbour more Balkan endemics than do springs, with the exception of species-poor high-altitude Drepanocladetum exannulati. The gradient structure of the vegetation was revealed by DCA and by CCA with forward selection of environmental factors. The major determinants of vegetation variation strongly differ above and below the timberline and likewise between springs and fens. The base-richness gradient controls the floristic variation of Bulgarian submontane fens, whereas the complete data set including both submontane and subalpine fens is governed by the altitude gradient from lowland and basin fens to subalpine fens rich in Balkan endemics. When focusing on sites above the timberline only, the first DCA axis separates fens from springs without organic matter. The major species turnover in springs follows the variation in water pH and mineral content in water, whereas fen vegetation variation is primarily controlled by succession gradient of peat accumulation. Altitude remains an important factor in all cases. Weak correlation between water pH and conductivity was found. This correlation was even statistically insignificant in fens above the timberline. Water pH is not influenced by mineral richness in Bulgarian high mountains, since it is buffered by decomposition of organic matter in fens. In springs, pH reaches maximum values due to strong aeration caused by water flow. The plant species richness decreases significantly with increasing altitude. The increase of species richness towards circumneutral pH, often found in mires, was not confirmed in Bulgarian high mountains. The correlation between species richness and pH was significant only when arctic-alpine species and allied European high-mountain species were considered separately. The richness of boreal species was independent on pH. Some of them had their optima shifted to more acidic fens as compared to regions below the timberline. Our results suggest that subalpine spring and fen vegetation should be analysed separately with respect to vegetation-environment correlations. Separate analysis of fens below and above timberline is quite appropriate.     

Comparing naturalized alien plants and recipient habitats across an east–west gradient in the Mediterranean Basin

Aim To investigate alien plant species invasion levels in different habitats and alien species traits by comparing the naturalized flora in different areas of the same biogeographical region. Location Spain, Italy, Greece and Cyprus. Methods Comparison of floristic composition, species traits and recipient habitats of naturalized alien neophytes across an east–west gradient comprising four countries in the European Mediterranean basin. Results A total of 782 naturalized neophytes were recorded; only 30 species were present in all four countries. Although floristic similarity is low, the four alien floras share the same patterns of growth form (mostly herbs), life cycle (mostly perennials) and life form (mostly therophytes, hemicryptophytes and phanerophytes). The majority of the recipient habitats were artificial. Wetlands were the natural habitats, with the highest numbers of naturalized species. Floristic similarity analyses revealed: (1) the highest floristic similarity between Italy and Spain, both of which were more similar to Greece than to Cyprus; (2) two groups of floristic similarity between habitat categories in each country (Greece–Cyprus and Italy–Spain); (3) a higher degree of homogenization in the plant assemblages in different habitats in Greece and Cyprus and a lower degree of homogenization in those in Italy and Spain; and (4) a higher degree of homogenization in artificial and natural fresh‐water habitats than in the other natural habitats. Main conclusions The floristic similarity of naturalized neophytes between the four countries is low, although the overall analysis indicates that the western group (Italy–Spain) is separated from the eastern group (Greece–Cyprus). Similar patterns emerged regarding the life‐history traits and recipient habitats. The artificial habitats and the natural wet habitats are those that are invaded most and display the greatest homogenization in all four countries. Coastal habitats display a lower degree of homogenization but a high frequency of aliens. Dry shrubs and rocky habitats display a lower degree of homogenization and a low frequency of aliens.     

Invasiveness of alien plants in Brussels is related to their phylogenetic similarity to native species

Aim Understanding the processes that drive invasion success of alien species has received considerable attention in current ecological research. From an evolutionary point of view, many studies have shown that the phylogenetic similarity between the invader species and the members of the native community may be an important aspect of invasiveness. In this study, using a coarse‐scale systematic sampling grid of 1 km², we explore whether the occupancy frequency of two groups of alien species, archaeophytes and neophytes, in the urban angiosperm flora of Brussels is influenced by their phylogenetic relatedness to native species. Location The city of Brussels (Belgium). Methods We used ordinary least‐squares regressions and quantile regressions for analysing the relationship between the occupancy frequency of alien species in the sampled grid and their phylogenetic distance to the native species pool. Results Alien species with high occupancy frequency in the sampled grid are, on average, more phylogenetically related to native species than are less frequent aliens, although this relationship is significant only for archaeophytes. In addition, as shown by the quantile regressions, the relationship between phylogenetic relatedness to the native flora and occupancy frequency is much stronger for the most frequent aliens than for rare aliens. Main conclusions Our data suggest that it is unlikely that species with very low phylogenetic relatedness to natives will become successful invaders with very high distribution in the area studied. To the contrary, under future climate warming scenarios, present‐day urban aliens of high occupancy frequency are likely to become successful invaders even outside urban areas.     

Distribution of alien plant species in relation to human disturbance on the Georgia Sea Islands

Abstract. This study investigates the effects of human disturbance and environmental factors on the distribution of alien plant species on the Georgia Sea Islands (GSI), USA. We sampled the absolute cover of native and alien plant species on two tourist islands (St. Simons Island and Jekyll Island) and on two protected National Wildlife Refuge Islands (Blackbeard Island and Wassaw Island). On each island, vegetation composition and environmental variables (soil properties and salt spray) were measured in two habitats that differed substantially in their degree of environmental stress, the more exposed primary dune and the more sheltered and inland maritime forest. Sites were further stratified within each habitat into areas that had different levels of human disturbance. Many alien species were present on all islands and the absolute cover of alien species was not significantly different among islands even though they varied substantially in their degree of accessibility and overall land use. Alien plant cover was appreciably greater in severely disturbed sites than in less disturbed sites on all islands and within both habitats. However, the difference between disturbance categories was much less pronounced in the primary dunes where human disturbance agents do not mitigate the harsh environmental conditions of this habitat (salt spray and saline soils). Alien plant abundance on the GSI is evidently more dependent upon the availability of disturbed ground than the degree of accessibility or overall island development. It appears that human disturbance increases alien cover in general, but in environments where the stress levels are not mitigated, human disturbance does little to foster alien invasions.