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.     

Anthropogenic fires increase alien and native annual species in the Chilean coastal matorral

Aim We tested the hypothesis that anthropogenic fires favour the successful establishment of alien annual species to the detriment of natives in the Chilean coastal matorral. Location Valparaíso Region, central Chile. Methods We sampled seed rain, seedbank emergence and establishment of species in four paired burned and unburned areas and compared (using GLMM) fire resistance and propagule arrival of alien and native species. To assess the relative importance of seed dispersal and seedbank survival in explaining plant establishment after fire, we compared seed rain and seedbank structure with post‐fire vegetation using ordination analyses. Results Fire did not change the proportion of alien species in the coastal matorral. However, fire increased the number of annual species (natives and aliens) of which 87% were aliens. Fire reduced the alien seedbank and not the native seedbank, but alien species remained dominant in burned soil samples (66% of the total species richness). Seed rain was higher for alien annuals than for native annuals or perennials, thus contributing to their establishment after fire. Nevertheless, seed rain was less important than seedbank survival in explaining plant establishment in burned areas. Main conclusions Anthropogenic fires favoured alien and native annuals. Thus, fire did not increase the alien/native ratio but increased the richness of alien species. The successful establishment of alien annuals was attributable to their ability to maintain rich seedbanks in burned areas and to the greater propagule arrival compared to native species. The native seedbank also survived fire, indicating that the herbaceous community has become highly resilient after centuries of human disturbances. Our results demonstrate that fire is a relevant factor for the maintenance of alien‐dominated grasslands in the matorral and highlight the importance of considering the interactive effect of seed rain and seedbank survival to understand plant invasion patterns in fire‐prone ecosystems.     

Alien and native species in Central European urban floras: a quantitative comparison

The paper provides quantitative information on the occurrence of alien species in Central European cities and analyses factors determining the richness of alien and native floras in this habitat type. Data for 54 cities (25 Polish, 24 German, 4 Czech and 1 Austrian) were gathered, and the representation of archaeophytes (i.e. aliens introduced before 1500 ad ), neophytes (introduced after that date) and native species was expressed. In an average city there were 87.4 archaeophytes (15.2% of the city flora) and 172.4 neophytes (25.2%) giving a total of 259.7 for alien species (40.3%). The number of native species averaged 386.5. The numbers of species in each category of immigration status increased significantly with city size. For neophytes, the species-area relationship showed a higher slope (0.49) on log–log axes than for both archaeophytes (0.16) and native species (0.30). Not only the number, but also the relative contribution of neophytes to the total flora, increased with city size, indicating that neophytes are the group which are most closely associated with human activity. On the other hand, archaeophytes were better represented in smaller cities, as they were confined to rural environment. A step-wise multiple regression was used to test for environmental variables acting as significant predictors, and explained between 40 and 65% of variation in the species numbers for particular categories of immigration status, providing the best fit for neophytes. City size was the best predictor for each characteristic, except of the proportion of total aliens, where the percentage of explained variability was low (8.2%), with latitude being the only significant predictor. Temperature was another highly significant predictor for the number of archaeophytes and total aliens, reflecting the origin of aliens in warmer areas. There was an effect of region on some flora characteristics. Polish cities had significantly higher proportion of archaeophytes and of total aliens than German cities. It is concluded that the occurrence of native and alien species in urban floras follows rather different pattern.     

Patterns of alien plant diversity in the urban landscapes of global biodiversity hotspots: a case study from the Himalayas

In an age of Anthropocene, the urban landscapes are recognised as the ‘hotspots’ of human-mediated alien species introductions. As the cities provide an ideal natural experimental system to investigate the patterns of alien plant diversity in urban landscapes, the present study aimed to unravel the taxonomic, biogeographic and ecological patterns of alien flora of Srinagar—one of the largest urban centres in the Himalayan biodiversity hotspot. The alien flora of Srinagar comprises 325 species, constituting ca.35% of total flora of the city. Out of the 325 alien species documented, 157 species (43%) were recorded to be under cultivation, while 168 species (57%) were growing in the wild (i.e., outside cultivation); those growing in the wild, in turn, comprised 110 cultivation escapes and 58 accidentally introduced plant species. Biogeographically, two-third of the alien plant diversity reported from Srinagar is native to Asia-Temperate. This indicates that climatic similarity between Asia-Temperate and Kashmir Himalayas facilitate in flourishing similar floristic diversity. The study highlights a relatively higher proportion of herbaceous growth form in the aliens growing in the wild (80%) than those under cultivation (43%). Similarly, 82% of the alien species under cultivation had a perennial life span, but those growing in the wild were dominated by annuals (44%). Currently, 45 species are growing as casuals and 124 species are naturalised (including 105 naturalised non-invasive and 19 naturalised invasive). Along the continuum of casual-naturalised-invasive categories, the contribution of cultivation escapes and accidently introduced aliens contrastingly shows decreasing and increasing trends respectively. Interestingly, the results revealed that the human practice of stopping cultivation of alien escapes increased rapidly as we move along the continuum. Thus, the present study has investigated the patterns of alien plant diversity in the urban landscape of Srinagar, and the results obtained offer scientific insights toward better scientific understanding and management of plant invasions in this Himalayan city, with wider policy implications for neighbouring urbanised landscapes in the Himalayas and other mountainous regions across the world.     

Urban domestic gardens (I): Putting small‐scale plant diversity in context

Abstract. As part of a larger survey of biodiversity in gardens in Sheffield, UK, we examined the composition and diversity of the flora in two 1‐m² quadrats in each of 60 gardens, and compared this with floristic data from semi‐natural habitats in central England and derelict urban land in Birmingham, UK. Garden quadrats contained more than twice as many taxa as those from any other habitat type. Ca. 33 % of garden plants were natives and 67 % aliens, mainly from Europe and Asia. A higher proportion of garden aliens originated from Asia and New Zealand than in the UK alien flora as a whole; 18 of the 20 most frequent plants in garden quadrats were natives, mostly common weeds. Garden quadrats showed no evidence of ‘nestedness’, i.e. a tendency for scarce species to be confined to the highest diversity quadrats. Conversely, species in all semi‐natural and derelict land data sets were significantly nested. Compared to a range of semi‐natural habitats, species richness of garden quadrats was intermediate, and strikingly similar to the richness of derelict land quadrats. Although species accumulation curves for all other habitats showed signs of saturation at 120 quadrats, gardens did not. Correlations between Sørensen similarity index and physical distance were insignificant for all habitat types, i.e. there was little evidence that physical distance played any part in structuring the composition of the quadrats in any of the data sets. However, garden quadrats were much less similar to each other than quadrats from semi‐natural habitats or derelict land.     

Widespread plant species: natives versus aliens in our changing world

Estimates of the level of invasion for a region are traditionally based on relative numbers of native and alien species. However, alien species differ dramatically in the size of their invasive ranges. Here we present the first study to quantify the level of invasion for several regions of the world in terms of the most widely distributed plant species (natives vs. aliens). Aliens accounted for 51.3% of the 120 most widely distributed plant species in North America, 43.3% in New South Wales (Australia), 34.2% in Chile, 29.7% in Argentina, and 22.5% in the Republic of South Africa. However, Europe had only 1% of alien species among the most widespread species of the flora. Across regions, alien species relative to native species were either as well-distributed (10 comparisons) or more widely distributed (5 comparisons). These striking patterns highlight the profound contribution that widespread invasive alien plants make to floristic dominance patterns across different regions. Many of the most widespread species are alien plants, and, in particular, Europe and Asia appear as major contributors to the homogenization of the floras in the Americas. We recommend that spatial extent of invasion should be explicitly incorporated in assessments of invasibility, globalization, and risk assessments.     

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.     

Research into Plant Invasions in a Crossroads Region: History and Focus

The Czech Republic is a central European country whose geographical location, natural conditions, history of human settlement, and present land-use management make it relatively prone to plant invasions, hence it represents a convenient model for their study. Research in plant invasions, which date to the late 19th century, is reviewed in the present paper. A long-term floristic tradition allowed for the accumulation of a large body of floristic data on alien plants. During the 1960s–1970s, the main research focus was on their distribution. In this period, attempts were also made to predict potential invasiveness of weeds of arable land. The success rate of this prediction was about 39%. Considerable effort was put into a detailed classification of human-accompanying plants and the terminology associated with the issue. There is a high level of taxonomic research conducted in the country, and the new Flora of the Czech Republic treats the immigration status of taxa with reasonable care and detail. A complete, serious catalogue of the alien plants of the country has been published recently: there are currently 1378 alien plants (33.4% of the total flora). The core of present research in plant invasions is in ecological, biological, and biogeographical studies, focussed on (i) the history of invasion of particular species since their introduction, (ii) the role and importance of alien species in vegetation, including their participation in succession, and (iii) the major invasive species of the Czech flora and comparison of congeners. Reynoutria spp., Heracleum mantegazzianum, Oenothera spp., Pinus strobus, and Bidens frondosa are among the taxa most intensively studied in recent time. A complete list of 69 invasive plants in the Czech flora introduced after the year 1500 [following the definition of Richardson et al. (2000a) Diversity and Distributions 6: 93–107] is given. The available legal instruments relevant to the issue are reviewed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Exotic plant species invade diversity hot spots: the alien flora of northwestern Kenya

We analysed the distribution of native and alien plant species across 20 ecogeographic zones of northwestern Kenya. The source pool for the majority of aliens was Europe and America. Thus, the source pool has a biogeographic bias which explains the low proportion of aliens in the tropics: most species in the European or American source pool are not well adapted to tropical conditions. As expected, native and alien plant species showed an area effect. Correcting for this area effect. species rich zones showed a higher proportion of alien plant species in their flora. At the analysed scale, species richness of native plant communities does not increase the resistance to invasions and alien plant species invade diversity hotspots. Compared to the other ecogeographic zone, the urban area around Nairobi showed an increased richness in alien and native plant species. This is very similar to findings in Europe, although the history of urbanisation is much shorter in Kenya. The species turnover between zones (β-diversity) shows a similar pattern in native and alien plant species. Within a very short time scale the alien plant species mapped the biogeographic patterns of natives, although the geography of human activities influences the propagule pressure.     

Four arguments why so many alien species settle into estuaries, with special reference to the German river Elbe

In one of the largest European rivers, the Elbe, from its source in the Czech Republic to the German North Sea, 31 alien macrozoobenthic species have been recorded in total. Most of these species have been introduced by shipping activities. With a total number of 21 species, many of the established aliens occur—partly exclusively—in the brackish area of the Elbe estuary. In order to explain this observed settlement characteristic, four main arguments come into consideration: (1) estuaries with intensive international shipping have a higher potential infection rate than other aquatic zones; (2) brackish water species have, due to specific physiological characteristics, a better chance of being transported alive than euhaline or freshwater species and they also probably have a higher perennation and establishment potential after release; (3) brackish waters have the greatest natural ‘indigenous species minimum’, so that more alien species can potentially establish; and (4) salt-tolerant limnetic alien species introduced into inland water reached the coast at first in the estuaries. It seems that the combination of brackish water with its unsaturated ecological niches and intensive international ship traffic has the highest potential infection rate for aquatic systems with alien macrozoobenthic species. And, estuaries are subjected to a two-sided invasion pressure by alien species, via the ocean (mainly shipping) and via inland waters (mainly shipping canal construction). The identification of such patterns is an important prerequisite for the development of a forward-looking alien monitoring and management strategy.     

Past and future of predictions in plant invasions: a field test by time

Intensive interest in the alien flora of the Czech Republic stimulated one of the earliest attempts (in the early 1970s) to predict potential invaders of arable land. Another paper published recently by the same research team using the same methodology provides a relatively unique possibility to assess the success of these predictions after a quarter of a century. The predictions were successful for 39.3% of the 28 species included, while 60.7% of invasion cases must be considered as failures. Prediction was rather unsuccessful for the members of Asteraceae (14.3%), and more correct for annuals/biennials (45.5%) than for perennials (16.7%). No pattern was found with respect to the area of origin. The results indicate that past predictions based largely on intuition were less successful than modern prediction systems using the knowledge of a large number of characters and carried out using advanced computation methods. The correct identification of invaders using such systems reaches values between 61% and 91%.     

The alien flora of Kashmir Himalaya

Compilation of alien flora from phytogeographically distinct regions is of immediate relevance not only for better understanding the patterns of plant invasion but also for explicating the processes promoting invasion at local, regional or global scales. Despite being at higher risk of invasion by plants because of its European colonial past, south Asia has received very little attention in respect of characterization of its alien flora. This paucity of baseline data necessitated compilation of the first catalogue of alien flora from the Kashmir Himalaya—a phytogeographically distinct south Asian region nestled in the northwestern folds of Himalayan biodiversity hotspot. Total alien flora of the region is represented by 571 plant species, belonging to 352 genera and 104 families. It constitutes a relatively higher (29%) proportion of the total flora of the region. Families with largest number of alien representatives are Poaceae (60 species), Asteraceae (54 species), and Brassicaceae (30 species). However, families such as Amaranthaceae (83%) and Chenopodiacae (71%) show higher percentage of aliens relative to their total number of plant species in the region. Most of the alien plant species (38%) trace their origin to Europe, followed by Asia (27%) and Africa (15%). Present study also reports, for the first time, occurrence of seven plant species in this region. Each alien plant species is provided with information on the origin, habit, mode/purpose of introduction, current invasion status, altitudinal range and the primary published source.     

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?     

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.     

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.     

Alien flora of the Korean Peninsula

The Korean Peninsula has been divided into the countries of North and South Korea since 1948. This adoption of different political and socio-economic regimes has significantly changed economic activity and international exchange by those nations, thus affecting the introduction of alien plants. Our study objective was to provide a comprehensive, up-to-date inventory of the alien flora on the Korean Peninsula and to analyse how the partitioning between North and South Korea has influenced the numbers and status of those alien plants. We identified pre-1876 aliens that were brought to the Korean Peninsula before 1876, when Korea opened up to foreign countries, and also determined the post-1876 aliens that were introduced after 1876. Plants in the latter group were further classified into those that arrived before the outbreak of the Korean War in 1950 (1876–1950 aliens) and those introduced since then (post-1950 aliens). We investigated the regions of origin, family compositions, and invasion status of those alien taxa. In total, 504 alien taxa were identified, with 102 taxa being pre-1876 and 117 being 1876–1950 aliens. After the Korean War, the number of alien plants in South Korea more than doubled (276 taxa), whereas the number reported for North Korea was considerably lower (33 taxa). This might be due to enhanced economic activity and a larger human population in South Korea. Climatic differences as well as variations in the recording intensity for alien flora between North and South Korea may also have contributed to the large contrast in numbers of new alien species recorded during the last decades. Coordinated measures between the two nations are necessary if the spread of invasive alien plants onto the Korean Peninsula is to be reduced.     

Plant Invasions on an Oceanic Archipelago

Plant invasions are particularly noticeable on oceanic islands. For many ecological or evolutionary phenomena, oceanic islands offer advantages in comparison to continental regions, because they are often simpler systems. The Juan Fernández (Robinson Crusoe) Islands, located 667 km west of continental Chile, provide an especially favorable case study of plant invasions on an oceanic archipelago. They have little flora, no influence from aboriginal peoples, and good historical and recent documentation of flora, vegetation and human impacts. The total flora of the archipelago consists of 441 vascular plants, of which 209 are native (125 of them endemic) and 232 are aliens. Many alien species exist on the Juan Fernández Archipelago, but three shrubs are particularly invasive: Aristotelia chilensis, Rubus ulmifolius, and Ugni molinae. About 15% of the total area of the island has been impacted by at least one of these shrubs. In addition, the herbaceous Acaena argentea, now occurs at high abundance in 12% of the total area of the island. Comparisons of earlier and recent surveys of vegetation reveal that the area impacted by Aristotelia, Rubus, and Ugni molinae has increased tremendously. Among the most frequent aliens are Euro-Mediterranean taxa, also present on other archipelagos. A few aliens that are serious invasives on other archipelagos have been recently observed near San Juan Bautisata, the only village on the island. Effective measures involving stronger monitoring and sanitation of incoming materials (foodstuffs, building materials, etc.), cutting or poisoning of invasives, and reseeding with native species, are necessary to help preserve the native and endemic flora (and biota) of these islands.     

Distribution of alien animal species richness in the Czech Republic

Biogeographical barriers formed by natural forces over billions of years have been substantially disrupted by human activity, particularly in recent centuries. In response to these anthropogenic changes, global homogenization of biota is observed at an ever‐increasing rate, causing environmental and economic losses as well as emerging health risks. Identifying factors underlying alien species richness is essential for prevention of future introductions and subsequent spread. In this study, we examined the effects of environmental and human‐related factors on distribution of alien animal species richness in the Czech Republic (Central Europe). We compiled a set of maps showing the level of invasion of six categories of alien animal species in each of 628 grid cells (ca. 12.0 × 11.1 km) covering the Czech Republic. Relationships between alien species richness and 12 variables characterizing climatic conditions, topography, land cover, and human population size were calculated using the generalized least squares method. Species richness of all alien species, of invertebrates, and of terrestrial species showed the strongest positive relationship with mean annual temperature, while the number of black and grey (proposed prominent invaders) and aquatic species was most closely related to the presence of large rivers. Alien vertebrates showed a strong negative relationship with annual precipitation. The highest alien animal species richness was found in and near large population centers and in agricultural landscapes in warm and dry lowlands. The gateways for alien aquatic species are rather large rivers over sport fishing and aquaculture import. Compiled maps create a powerful visual communication tool, useful in development of programs to prevent future introductions.     

Effects of urban land cover on the local species pool in Britain

Regression models were used to quantify the relationship between the amount of urban land and composition of local species pools. There was no evidence that urban land cover increases the richness of plant species, based on a survey of 785 2-km squares of which 157 had > 10% urban land cover. However, the number of alien plants is significantly higher in urban areas. Complete urbanization approximately doubles the proportion of alien species, and the proportion of aliens is twice as high in southern Britain as in the north. The flora of urban tetrads consists of ubiquitous native species and introduced species characteristic of waste ground, but woodland species are poorly represented. At the tetrad scale. enhanced dispersal by man is not the main factor for maintaining the urban flora; availability of urban habitats and high levels of disturbance are more important. The planned housing expansion to greenfield sites in Britain will increase the proportion of alien species, yet the majority of native species should persist in urban areas if existing woodland is preserved.     

Rocks and Walls: Natural Versus Secondary Habitats

Walls are often considered secondary habitats for vegetation of natural rock surfaces. Compared with rocks, walls differ in many features, for example the presence of a binding material (mainly calcareous mortar), location in settlements and exposure to human impact. A data set of 1,205 phytosociological relevés recorded on horizontal wall tops, wall verticals and rock verticals in the Czech Republic was used to compare their vegetation with regard to i) species composition (frequent species, species diversity, endangered and alien species) and ii) the ecological requirements of the respective species. Gamma diversity of vascular plant species was comparable in all habitat types (242 species on wall tops, 212 species on wall verticals and 197 species on rock verticals). Wall verticals had higher beta diversity, but lower alpha diversity than rocks. Remarkable differences were found comparing the diversity of alien species. Whereas alpha and gamma diversities of aliens were higher on both wall habitats, beta diversity of aliens was the highest on rocks. The high floristic heterogeneity of walls is mainly attributable to the large pool of species from the surrounding urbanized landscape (e.g., cultivated ornamental species and synanthropic weeds) that are favoured by high nutrient inputs. In contrast, species characteristic of rocks are mainly substrate specialists. Walls and rocks share the frequent occurrence of ferns, grasses and herbs typical of forest understorey and clearings. Compared with rocks, walls are generally colonized by species requiring higher nutrient content, soil reaction, temperature and moisture. Secondary wall habitats might be suitable for some rare and endangered species, but contrary to rocks their occurrences are only accidental and temporary. The representation of aliens was considerably higher on walls (approximately 35%) than on rocks (9%).