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.     

Phyloecology of urban alien floras

1.  Understanding the mechanisms that affect invasion success of alien species is a major issue in current ecological research. Although many studies have searched for either functional or habitat attributes that drive invasion mechanisms, few researchers have addressed the role of phylogenetic diversity of alien species.
2.  Here, using data from 21 urban floras located in Europe and eight in the USA, we show that the phylogenetic diversity of alien species is significantly lower than that of native species, both at the continental scale and at the scale of single cities.
3.  Second, we show that if archaeophytes and neophytes (non-native species introduced into Europe before and after AD 1500, respectively) are analysed separately, archaeophytes show lower phylogenetic diversity than neophytes, while the phylogenetic structure of neophytes is indistinguishable from a random sample of species from the entire species pool.
4.  Our results suggest that urban aliens are subject to environmental filters that constrain their phylogenetic diversity, although these filters act more strongly upon archaeophytes than neophytes.
5.   Synthesis. Despite the huge taxonomic diversity of plants imported into European and American cities, the strong environmental filters imposed by cities constrain the functional diversity of urban floras, which is reflected in their generally low phylogenetic diversity. Urban alien floras are mainly composed of phylogenetically related species that are well adapted to anthropogenic habitats, although these filters are stronger for species groups with longer residence times.     

Determinants of native and alien species richness in the urban flora of Rome

Until now, analytical studies on European urban floras have mostly concentrated on the central and north‐western parts of the continent. In this paper, factors determining species richness of urban flora were studied for the city of Rome, Italy, based on a comprehensive floristic survey carried out between 1985 and 1994, and updated in 2005. All species were recorded in grid cells of 1.6 km² and classified into native and alien (the latter divided into archaeophytes and neophytes). The grids were classified with respect to the prevailing habitat type, area available to vegetation, level of disturbance and geographical position within the city. Data were analysed using minimal adequate models. Total species number was determined by habitat and its interaction with position on the north‐west gradient; other variables explained much less variance. Holding other variables constant, the average species number per grid cell was highest in archaeological sites and parks, followed by woodlands and rivers, and grasslands and recent developments. Residential areas and the historical centre were poorest in species number. Towards the north of the city, species richness in corresponding habitats increases because of higher landscape heterogeneity and closer association with diaspore pools in the surroundings. Native species make up on average 84% of the total species numbers, and trends opposite to those for the total number of species were found for the proportional representation of aliens. The occurrence of alien and native species in the flora of Rome is driven by similar factors, but factors that increase representation of aliens decrease that of natives and vice versa. The representation of neophytes and native species in grid cells was easier to explain (74% of variation accounted for) than that of archaeophytes (27%); this result reflects that in terms of ecology and response to factors, archaeophytes take an intermediate position between native plants and neophytes. Proportional representation of neophytes decreased with increasing area available to vegetation, reflecting that semi‐natural vegetation is better developed where less fragmented.     

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.     

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.     

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.     

Diversity of native and alien vascular plant species of dry grasslands in central Europe

Question: Which factors determine diversity of native and alien vascular plant species in semi‐natural dry grasslands? Location: Northern limestone Alps to the southern rim of the Bohemian massif in northern Austria. Methods: In 70 randomly chosen dry grassland patches (0.008 ha ‐ 7 ha) we sampled a complete inventory of vascular plant species at each site. We analysed the correlation between species diversity of natives, archaeophytes (pre‐1500 aliens) and neophytes (post‐1500 aliens). We used GLM to study the relationship of species number (natives, neophytes, archaeophytes) to five explanatory variables (altitude, within habitat diversity, habitat diversity of adjacent areas, within land‐use diversity and land‐use in adjacent areas). Orthogonal components of these variables were derived with a PCA and used in the models. We also tested the influence of minimum residence time (MRT) and the covariables origin, mode of introduction and life form on the number of grassland sites with neophytes with analogous GLMs. Results: Native species diversity species was positively correlated with the species diversity of new, but not old invaders. GLM explains 70% of the variance in the number of native species. Patch size explained the largest part of the variation in the number of native species. PCA axes 1 and 3 were significantly related to the number of native species. Axis 1was related to on‐site habitat and land‐use diversity. The GLM of the archaeophyte diversity explains 18% of the variance. Altitude and presence of fields and grassland in the neighbourhood mainly explained archaeophyte species diversity. The GLM of neophyte diversity explains 12% of the variance. The number of neophytes was positively related to that of archaeophytes. Only PCA axis 3, which is mainly influenced by adjacent land‐use types, showed a relationship with neophytes. MRT, mode of introduction and region of origin (but not life form) were significantly related to the number of grassland sites invaded by neophytes, explaining 35% of the variance. Conclusion: Most factors governing native species diversity are not significantly related to alien species diversity. Additional determinants of the local scale diversity of alien species exist such as region of origin and historical factors (MRT, mode of introduction).     

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

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

An updated checklist of the vascular flora alien to Italy

An updated inventory of the vascular flora alien to Italy, providing details on the occurrence at regional level, is presented. The checklist includes 1597 species, subspecies, and hybrids, distributed in 725 genera and 152 families; 2 taxa are lycophytes, 11 ferns and fern allies, 33 gymnosperms, and 1551 angiosperms. 157 taxa are archaeophytes and 1440 neophytes. The alien taxa currently established in Italy are 791 (570 naturalized and 221 invasive), while 705 taxa are casual aliens, 4 are not assessed, 7 are of unknown regional distribution, 47 have not been confirmed in recent times, 3 are considered extinct or possibly extinct in the country, and 40 are doubtfully occurring in Italy. This checklist allows to establish an up-to-date number (9792) of taxa constituting the whole (native and alien) Italian flora.     

The alien flora of Greece: taxonomy,life traits and habitat preferences

The aim of the paper is the state-of-the-art assessment of the alien flora of Greece and its traits. The dataset consists of a total of 343 alien taxa, including 49 archaeophytes. The taxonomy, life traits and habitat of the 294 neophytes are analysed vs their naturalisation status. Out of the 122 (41%) naturalised neophytes, 50 are identified as exhibiting invasive behaviour. Poaceae, Asteraceae, Amaranthaceae, Solanaceae, Fabaceae, and Polygonaceae are the plant families richest in alien taxa. The majority of them are of American origin, followed by those of Asiatic and Mediterranean origin. The neophytes are predominantly herbs, most of them annuals. Yet, the perennial life cycle is equally frequent with the annual one and the proportion of phanerophytes in the alien flora is increased compared to the one of the native flora. Regarding flowering traits, most of the aliens have a long flowering period (over 1 month) and flower in late spring, summer and autumn, when few of the native plants are in bloom. Vertebrate zoochory and anemochory are the two dispersal modes mostly utilised by the alien plants (43 and 28%, respectively), while more than one dispersal mechanisms are functional for 56% of them. Artificial habitats have the highest frequencies of alien plants. The natural habitats with the highest numbers of aliens are the coastal ones and inland surface waters. Opuntia ficus-barbarica, Ailanthus altissima, Oxalis pes-caprae, Erigeron bonariensis, Amaranthus albus and Symphyotrichum squamatum are typical cases of plants characterised as invasive, having established in almost all the habitat groups identified. The diversity of the ecological characteristics of the plants suggests a potential of impacts that needs to be further assessed.     

Which factors determine plant invasions in man-made habitats in the Czech Republic?

Factors determining the invasibility of different types of anthropogenic vegetation were studied in the Czech Republic. A data set of 3420 vegetation plots recorded between 1945 and 2005, containing 913 species, was used. A set of climatic variables (mean annual temperature and precipitation, together with elevation), propagule pressure (substituted by human population density) and local habitat conditions (substituted by values of CSR life strategies and Ellenberg indicator values of native species) was obtained for each plot. All species were classified as native, archaeophytes (i.e. alien species introduced before 1500), and neophytes (i.e. aliens introduced after 1500) and their relative proportion was calculated for each plot. Regression tree models were used to determine the ecological characteristics of the most invasible man-made habitats in the Czech Republic. The plots contained on average 31.9% archaeophytes and 7.3% neophytes. Correlation between the proportions of archaeophytes and neophytes was positive and significant. Both archaeophytes and neophytes were found predominantly in strongly disturbed habitats with a high nutrient supply located at low elevations in warmer climatic areas of the Czech Republic. Archaeophytes are more influenced by local habitat conditions and preferentially colonize sunny and dry man-made habitats with higher soil reaction. Neophytes have no special preferences for local habitat conditions and their highest proportion was found mainly in disturbed habitats at low elevations. Our results show that for anthropogenic vegetation in the Czech Republic, ecological and habitat characteristics are more important factors for plant invasions than different land use in the surrounding area.     

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.     

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.     

Widely distributed native and alien plant species differ in arbuscular mycorrhizal associations and related functional trait interactions

It is debated whether alien plants in new environments benefit from being mycorrhizal and whether widely distributed natives and aliens differ in their associations with mycorrhizal fungi. Here, we compared whether species differing in their origin status, i.e. natives, archaeophytes (alien species introduced before the year 1500) and neophytes (introduced after the year 1500), and arbuscular mycorrhizal (AM) status (obligate, facultative, non‐mycorrhizal) differ in their area of occupancy in Germany (i.e. number of occupied grid cells, each ~130 km²). We used generalized linear models, incorporating main effects and up to three‐way interactions combining AM status, origin status and plant functional traits. The latter were chosen to describe the possible trade‐off in carbon allocation either towards the symbiosis or to other plant structures, such as storage organs (significant interactions involving traits were assumed to indicate the existence of such trade‐offs). AM status significantly explained the area of occupancy of natives and neophytes – with facultative mycorrhizal species occupying the largest area in both groups – but was less pronounced among archaeophytes. Archaeophytes may have reduced dependency on AM fungi, as they are generally agricultural weeds and the symbiosis potentially becomes obsolete for plants growing in habitats providing a steady provision of nutrients. Trait interactions between AM status and other functional traits were almost exclusively detected for neophytes. While facultative mycorrhizal neophytes benefit from trade‐offs with other traits related to high C cost in terms of area of occupancy, such trade‐offs were almost absent among natives. This indicates that natives and neophytes benefit differently from the symbiosis and suggests that native AM fungal partners might be less important for neophytic than for native plant species or that more time is required to establish similar relationships between neophytes and native fungal symbionts.     

Effects of different cultivation types on native and alien weed species richness and diversity in Moravia (Czech Republic)

Changes in weed species richness and beta-diversity are partly attributable to different types and intensity of disturbance and partly to broad-scale variation in environmental conditions. We compiled a data set of 434 vegetation plots of weed vegetation in root crop and cereal fields in Moravia (eastern Czech Republic) to compare the effects of environmental conditions and different disturbance regimes on species richness and beta-diversity. To detect changes in species richness, we related the variation in species richness to individual environmental conditions. To assess differences in beta-diversity between the vegetation of cereal and root crop fields, we used Whittaker's measure of beta-diversity. The relative importance of each environmental variable for the variation in species composition was evaluated using canonical correspondence analysis. All analyses were done for all vascular plant species and separately for native species, archaeophytes and neophytes. A comparison of weed vegetation of root crops and cereals showed a distinct dichotomy between these two types of weed vegetation. There was no significant difference in total species richness and native species richness; however, cereal fields were richer in archaeophytes and root crop fields were richer in neophytes. The beta-diversity of weed vegetation was higher in root crops. Environmental factors explained a significant part of the variability in richness of both natives and aliens. The richness of native species increased and beta-diversity decreased with increasing precipitation. The opposite relationship was found for archaeophytes, in both cereals and root crops. These results confirmed the importance of climatic factors and management practices for changes in weed species composition. They also showed a distinct pattern of species richness and beta-diversity of native and alien weed species.     

Czech alien flora and the historical pattern of its formation: what came first to Central Europe?

Temporal patterns of immigration to the country were analysed using 668 alien species in the flora of the Czech Republic for which the dates of the first record were available (64.8% of the total number of 1031 so-called neophytes, i.e. aliens introduced after the year 1500). After a period of initial slow increase lasting to the 1840s, the accumulation of neophytes over time could be best fitted by a linear model that explained 97% of the variance. The intensity of floristic research, which varied between periods, did not significantly affect the overall increase in the number of aliens. The effect of species traits on the year of introduction was evaluated, with continent of origin, introduction type (deliberate or accidental), life history, Grime's life strategy, onset of flowering, mode of dispersal and propagule size as explanatory variables. Species of European origin and CSR strategists arrived earlier than those with other origins and strategies. Deliberately introduced species appeared earlier than accidental arrivals, and those cultivated for utilitary reasons on average arrived earlier than ornamentals. Species capable of early flowering were remarkably more prevalent among early newcomers. A separate analysis of accidentally introduced American species also identified life history as a significant predictor of immigration time, with annuals being introduced earlier than biennials and perennials. The data contribute to an understanding of a crucial stage of the invasion process that has received little attention in the literature. The model "early alien" to Central Europe is a European species with a CSR strategy deliberately brought for cultivation as a utilitary plant. Once it escaped from cultivation, its establishment in the wild was favoured by its ability to flower early and, therefore, complete the life cycle.     

A comparison of the urban flora of different phytoclimatic regions in Italy

This study is a comparison of the spontaneous vascular flora of five Italian cities: Milan, Ancona, Rome, Cagliari and Palermo. The aims of the study are to test the hypothesis that urbanization results in uniformity of urban floras, and to evaluate the role of alien species in the flora of settlements located in different phytoclimatic regions. To obtain comparable data, ten plots of 1 ha, each representing typical urban habitats, were analysed in each city. The results indicate a low floristic similarity between the cities, while the strongest similarity appears within each city and between each city and the seminatural vegetation of the surrounding region. In the Mediterranean settlements, even the most urbanized plots reflect the characters of the surrounding landscape and are rich in native species, while aliens are relatively few. These results differ from the reported uniformity and the high proportion of aliens which generally characterize urban floras elsewhere. To explain this trend the importance of apophytes (indigenous plants expanding into man-made habitats) is highlighted; several Mediterranean species adapted to disturbance (i.e. grazing, trampling, and human activities) are pre-adapted to the urban environment. In addition, consideration is given to the minor role played by the 'urban heat island' in the Mediterranean basin, and to the structure and history of several Italian settlements, where ancient walls, ruins and archaeological sites in the periphery as well as in the historical centres act as conservative habitats and provide connection with seed-sources on the outskirts.     

Do urban areas act as foci for the spread of alien plant species? An assessment of temporal trends in the UK

Aim Given that urban landscapes often act as a point of entry for many non‐native species and urban development continues to increase as the human population rapidly expands, an understanding of the interaction between urbanization and non‐native plant species is important both in the control of potentially invasive species and in the conservation of native biodiversity. We investigated the spatial and temporal relationship between urban land cover and the distribution of non‐native species in Britain using two floristic data sets collected at two different time periods: 1987–88 and 2003–04. Location UK. Methods Using floristic data collected by the Botanical Society of the British Isles in 1987–88 (Monitoring Scheme) and 2003–04 (Local Change) in conjunction with habitat data obtained from the Land Cover Map of the UK, we conducted multiple regression analyses both within and between years on both groups of species (natives, neophytes and archaeophytes) and individual species. Results Neophytes (alien species introduced after 1500) were very strongly associated with urban land cover in both time periods and do not appear to be spreading out of urban habitats into the wider countryside. Archaeophytes (alien species introduced before 1500), however, showed a strong association with urban habitats in the earlier 1988 data set but no longer showed this association in the 2004 data set. Analysis at the individual species level showed that a large percentage of alien plant species, particularly archaeophytes, were not strongly associated with urban land cover or were negatively associated with such habitats. Main conclusions Our results suggest that there has been a reduction in the urban association of archaeophytes that is likely to have resulted from the recovery of archaeophytes associated with non‐urban (especially arable) habitats, following their decline in mid‐20th century, rather than from the movement of aliens into the wider countryside from urban habitats.     

Alien plant species and factors of invasiveness of anthropogenic vegetation in the Northwestern Balkans — a phytosociological approach

We studied the anthropogenic vegetation of the Northwest Balkans in order to determine its susceptibility to invasion by alien plant species. We compiled a dataset of 3089 vegetation plots sampled between 1939 and 2009, recording a set of variables for each sample plot in order to determine which factors have the most effect on a habitat’s vulnerability to invaders. We calculated the proportion of native species, archaeophytes and neophytes for each plot. We used regression tree models to determine the site conditions of the most invaded anthropogenic habitats. The sample plots contained an average of 12.7% alien plant species, with a low proportion of archaeophytes (4.3%) and 8.4% neophytes. Local habitat conditions proved to have the largest effect, rather than climatic variables or propagule pressure. The proportion of archaeophytes follows a different pattern than that seen in central and northern Europe, indicating that macroecological factors are more important. Neophytes show a similar distribution to other European locations.     

Changes in the functional composition of a Central European urban flora over three centuries

Documents on historical floras provide unique opportunities to analyze past changes and to show trends in biodiversity. We studied the historical and recent flora of the city of Halle in Central Germany. Our earliest records date back to the year 1687; the youngest were sampled in 2008. More than 20 other floras provide information for time in-between, covering ca. 320 years in total. We checked all historical plant occurrences for plausibility. The species turnover of 22% that took place in the study period should also yield changes in the functional composition of the flora. We identified native species and archaeophytes that went extinct since 1689 and 1856, respectively, and all neophytes that were introduced since 1689 or 1856. This ‘double’ calculation minimized the influence of so-called possible pseudo-absences. Contingency tables assisted to identify trait states which were associated with extinction or introduction. Time-series analysis identified temporal trends in trait state ratio development after testing for temporal autocorrelation. Within the study period, species of bogs, nitrogen-poor habitats or plants with helomorphic leaves got extinct more often than expected by chance. Species dispersed by humans, plants preferring nitrogen-rich or warm habitats, shrubs and trees, and species with mesomorphic leaves were, amongst others, over-represented among introduced neophytes. Land-use changes such as the transformation from agriculture to urban land use or the drainage of bogs are discussed as main drivers of these developments. Additionally, climatic changes, contamination of habitats and gardeners’ preferences for specific plants are presumed to having caused floristic changes. Our study shows the vast influence humans had and still have on biodiversity by intentionally or unintentionally selecting specific functional plant types and thus changing the composition of the flora.