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1. Pathways describe the processes that result in the introduction of alien species from one location to another. A framework is proposed to facilitate the comparative analysis of invasion pathways by a wide range of taxa in both terrestrial and aquatic ecosystems. Comparisons with a range of data helped identify existing gaps in current knowledge of pathways and highlight the limitations of existing legislation to manage introductions of alien species. The scheme aims for universality but uses the European Union as a case study for the regulatory perspectives. 2. Alien species may arrive and enter a new region through three broad mechanisms: importation of a commodity, arrival of a transport vector, and/or natural spread from a neighbouring region where the species is itself alien. These three mechanisms result in six principal pathways: release, escape, contaminant, stowaway, corridor and unaided. 3. Alien species transported as commodities may be introduced as a deliberate release or as an escape from captivity. Many species are not intentionally transported but arrive as a contaminant of a commodity, for example pathogens and pests. Stowaways are directly associated with human transport but arrive independently of a specific commodity, for example organisms transported in ballast water, cargo and airfreight. The corridor pathway highlights the role transport infrastructures play in the introduction of alien species. The unaided pathway describes situations where natural spread results in alien species arriving into a new region from a donor region where it is also alien. 4. Vertebrate pathways tend to be characterized as deliberate releases, invertebrates as contaminants and plants as escapes. Pathogenic micro-organisms and fungi are generally introduced as contaminants of their hosts. The corridor and unaided pathways are often ignored in pathway assessments but warrant further detailed consideration. 5. Synthesis and applications. Intentional releases and escapes should be straightforward to monitor and regulate but, in practice, developing legislation has proved difficult. New introductions continue to occur through contaminant, stowaway, corridor and unaided pathways. These pathways represent special challenges for management and legislation. The present framework should enable these trends to be monitored more clearly and hopefully lead to the development of appropriate regulations or codes of practice to stem the number of future introductions.

• Background Biological invasions are a major ecological and socio-economic problem in many parts of the world. Despite an explosion of research in recent decades, much remains to be understood about why some species become invasive whereas others do not. Recently, polyploidy (whole genome duplication) has been proposed as an important determinant of invasiveness in plants. Genome duplication has played a major role in plant evolution and can drastically alter a plant's genetic make-up, morphology, physiology and ecology within only one or a few generations. This may allow some polyploids to succeed in strongly fluctuating environments and/or effectively colonize new habitats and, thus, increase their potential to be invasive. • Scope We synthesize current knowledge on the importance of polyploidy for the invasion (i.e. spread) of introduced plants. We first aim to elucidate general mechanisms that are involved in the success of polyploid plants and translate this to that of plant invaders. Secondly, we provide an overview of ploidal levels in selected invasive alien plants and explain how ploidy might have contributed to their success. • Conclusions Polyploidy can be an important factor in species invasion success through a combination of (1) 'pre-adaptation', whereby polyploid lineages are predisposed to conditions in the new range and, therefore, have higher survival rates and fitness in the earliest establishment phase; and (2) the possibility for subsequent adaptation due to a larger genetic diversity that may assist the 'evolution of invasiveness'. Alternatively, polyploidization may play an important role by (3) restoring sexual reproduction following hybridization or, conversely, (4) asexual reproduction in the absence of suitable mates. We, therefore, encourage invasion biologists to incorporate assessments of ploidy in their studies of invasive alien species.

This paper considers key issues in plant invasion ecology, where findings published since 1990 have significantly improved our understanding of many aspects of invasions. The review focuses on vascular plants invading natural and semi-natural ecosystems, and on fundamental ecological issues relating to species invasiveness and community invasibility. Three big questions addressed by the SCOPE programme in the 1980s (which species invade; which habitats are invaded; and how can we manage invasions?) still underpin most work in invasion ecology. Some organizing and unifying themes in the field are organism-focused and relate to species invasiveness (the tens rule; the concept of residence time; taxonomic patterns and Darwin’s naturalization hypothesis; issues of phenotypic plasticity and rapid evolutionary change, including evolution of increased competitive ability hypothesis; the role of long-distance dispersal). Others are ecosystem-centred and deal with determinants of the invasibility of communities, habitats and regions (levels of invasion, invasibility and propagule pressure; the biotic resistance hypothesis and the links between diversity and invasibility; synergisms, mutualisms, and invasional meltdown). Some theories have taken an overarching approach to plant invasions by integrating the concepts of species invasiveness and community invasibility (a theory of seed plant invasiveness; fluctuating resources theory of invasibility). Concepts, hypotheses and theories reviewed here can be linked to the naturalization-invasion continuum concept, which relates invasion processes with a sequence of environmental and biotic barriers that an introduced species must negotiate to become casual, naturalized and invasive. New research tools and improved research links between invasion ecology and succession ecology, community ecology, conservation biology and weed science, respectively, have strengthened the conceptual pillars of invasion ecology.

Urbanization contributes to the loss of the world's biodiversity and the homogenization of its biota. However, comparative studies of urban biodiversity leading to robust generalities of the status and drivers of biodiversity in cities at the global scale are lacking. Here, we compiled the largest global dataset to date of two diverse taxa in cities: birds (54 cities) and plants (110 cities). We found that the majority of urban bird and plant species are native in the world's cities. Few plants and birds are cosmopolitan, the most common being Columba livia and Poa annua. The density of bird and plant species (the number of species per km2) has declined substantially: only 8% of native bird and 25% of native plant species are currently present compared with estimates of non-urban density of species. The current density of species in cities and the loss in density of species was best explained by anthropogenic features (landcover, city age) rather than by non-anthropogenic factors (geography, climate, topography). As urbanization continues to expand, efforts directed towards the conservation of intact vegetation within urban landscapes could support higher concentrations of both bird and plant species. Despite declines in the density of species, cities still retain endemic native species, thus providing opportunities for regional and global biodiversity conservation, restoration and education.

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.

Agricultural intensification is critical to meet global food demand, but intensification threatens native species and degrades ecosystems. Sustainable intensification (SI) is heralded as a new approach for enabling growth in agriculture while minimizing environmental impacts. However, the SI literature has overlooked a major environmental risk. Using data from eight countries on six continents, we show that few governments regulate conventionally bred pasture taxa to limit threats to natural areas, even though most agribusinesses promote taxa with substantial weed risk. New pasture taxa (including species, subspecies, varieties, cultivars, and plant-endophyte combinations) are bred with characteristics typical of invasive species and environmental weeds. By introducing novel genetic and endophyte variation, pasture taxa are imbued with additional capacity for invasion and environmental impact. New strategies to prevent future problems are urgently needed. We highlight opportunities for researchers, agribusiness, and consumers to reduce environmental risks associated with new pasture taxa. We also emphasize four main approaches that governments could consider as they build new policies to limit weed risks, including (i) national lists of taxa that are prohibited based on environmental risk; (ii) a weed risk assessment for all new taxa; (iii) a program to rapidly detect and control new taxa that invade natural areas; and (iv) the polluter-pays principle, so that if a taxon becomes an environmental weed, industry pays for its management. There is mounting pressure to increase livestock production. With foresight and planning, growth in agriculture can be achieved sustainably provided that the scope of SI expands to encompass environmental weed risks.

Rapid economic development in the past century has translated into severe pressures on species survival as a result of increasing land-use change, environmental pollution, and the spread of invasive alien species. However, though the impact of these pressures on biodiversity is substantial, it could be seriously underestimated if population declines of plants and animals lag behind contemporary environmental degradation. Here, we test for such a delay in impact by relating numbers of threatened species appearing on national red lists to historical and contemporary levels of socioeconomic pressures. Across 22 European countries, the proportions of vascular plants, bryophytes, mammals, reptiles, dragonflies, and grasshoppers facing medium-to-high extinction risks are more closely matched to indicators of socioeconomic pressures (i.e., human population density, per capita gross domestic product, and a measure of land use intensity) from the early or mid-, rather than the late, 20th century. We conclude that, irrespective of recent conservation actions, large-scale risks to biodiversity lag considerably behind contemporary levels of socioeconomic pressures. The negative impact of human activities on current biodiversity will not become fully realized until several decades into the future. Mitigating extinction risks might be an even greater challenge if temporal delays mean many threatened species might already be destined toward extinction.

This paper explores how a lack of taxonomic expertise, and by implication a dearth of taxonomic products such as identification tools, has hindered progress in understanding andmanaging biological invasions. It also explores how the taxonomic endeavour could benefit from studies of invasive species. We review the literature on the current situation in taxonomy with a focus on the challenges of identifying alien plant species and explorehowthis has affected the study of biological invasions. Biosecurity strategies, legislation dealing with invasive species, quarantine, weed surveillance and monitoring all depend on accurate and rapid identification of non-native taxa. However, such identification can be challenging because the taxonomic skill base in most countries is diffuse and lacks critical mass. Taxonomic resources are essential for the effective management of invasive plants and incorrect identifications can impede ecological studies. On the other hand, biological invasions have provided important tests of basic theories about species concepts. Better integration of classical alpha taxonomy and modern genetic taxonomic approaches will improve the accuracy of species identification and further refine taxonomic classification at the level of populations and genotypes in the field and laboratory. Modern taxonomy therefore needs to integrate both classical and new concepts and approaches. In particular, differing points of view between the proponents of morphological and molecular approaches should be negotiated because a narrow taxonomic perspective is harmful; the rigour of taxonomic decision- making clearly increases if insights from a variety of different complementary disciplines are combined and confronted. Taxonomy plays a critical role in the study of plant invasions and in turn benefits from the insights gained from these studies. © 2013 The Authors.

Aim: Alien species successfully colonize new ranges if they encounter favourable environmental conditions there and possess traits that match new challenges. Climate-matching approaches comparing native and exotic ranges mostly consider temperature and precipitation niches of alien species, but have largely ignored UV-B radiation. UV-B fundamentally differs between hemispheres, with much higher levels at southern than at northern latitudes. Consequently, UV-B might act at the global scale and present a so far neglected filter that species need to overcome when invading high-UV-B environments. Location: We performed two multi-species common garden experiments, conducted in the native European range (Germany) and the high-UV-B exotic range (New Zealand) to test for preadaptation to UV-B. Methods: We used 25 herbaceous species from open habitats, which we exposed in each range to three UV radiation treatments: (a) natural sunlight, (b) exclusion of UV-B while allowing natural UV-A, and (c) exclusion of UV-B and UV-A. We additionally used plant traits (TRY), global distribution data (GBIF, GloNAF) and global UV-B satellite data (glUV) to determine species-specific characteristics as fostering agents of UV-B tolerance. The joint analysis of experimental and macroecological data allowed quantification of species plasticity and identification of beneficial species traits in high-UV-B environments. Results: Our results showed an overall limiting effect of UV-B in both common gardens but the UV-stress response tended to be more pronounced in the invaded range. Across all species, we found little evidence for preadaptation by functional plant traits. In contrast, preadaptation to climatic conditions related to the species’ native UV-B niche was of greater importance for plant performance in the presence of UV-B radiation. Main conclusions: For predicting alien species’ ability to expand into high-UV-B environments, macroclimatic niche characteristics of the species’ native range might be better predictors than functional traits and should be more considered in future projection models.

Questions: (1) What are the most important abiotic environmental variables influencing succession in central European man-made habitats? (2) How do these variables interact with one another and with variation in community properties? Location: Central, western and southern parts of the Czech Republic. Habitats included old fields, urban sites, spoil heaps after coal mining, sites at water reservoirs, extracted sand pit and peatland and reclaimed sites in areas deforested by air pollution. Methods: We investigated vegetation patterns on 15 successional seres, sampled by the same methods. Time of succession over which the data were available ranged from 12 to 76 years. The cover of vascular plant species (in %) was estimated in 5 m × 5 m plots. The relationships between vegetation characteristics (species composition, total cover, cover of woody species, species number and rate of dominant species turnover) and 13 abiotic site factors, including climatic and soil variables, were tested using CCA ordination and regression models. Results: Substratum pH, the only substratum characteristic, and climate were the environmental variables significantly affecting the vegetation patterns in the course of succession. The rate of succession, measured as the turnover of dominant species, was significantly more rapid in lowland than in mountain climates. On alkaline soils, species numbers in succession increased towards warmer climates. However, acid soils prevented any increase in species numbers, regardless of the climate. Surprisingly, forms of nitrogen and contents of C, P and cations did not exhibit any significant effect on the vegetation characteristics studied. Conclusions: Our approach, to compare a number of seres, can contribute not only to our understanding of succession, but also to help restoration projects to predict vegetation change because the crucial environmental variables, as identified by this study, are easy to measure. Nomenclature: Tutin et al. (1964–1980).