Czech Academy of Sciences, Institute of Botany

Abstract. Much confusion exists in the English‐language literature on plant invasions concerning the terms ‘naturalized’ and ‘invasive’ and their associated concepts. Several authors have used these terms in proposing schemes for conceptualizing the sequence of events from introduction to invasion, but often imprecisely, erroneously or in contradictory ways. This greatly complicates the formulation of robust generalizations in invasion ecology. Based on an extensive and critical survey of the literature we defined a minimum set of key terms related to a graphic scheme which conceptualizes the naturalization/invasion process. Introduction means that the plant (or its propagule) has been transported by humans across a major geographical barrier. Naturalization starts when abiotic and biotic barriers to survival are surmounted and when various barriers to regular reproduction are overcome. Invasion further requires that introduced plants produce reproductive offspring in areas distant from sites of introduction (approximate scales: > 100 m over < 50 years for taxa spreading by seeds and other propagules; > 6 m/3 years for taxa spreading by roots, rhizomes, stolons or creeping stems). Taxa that can cope with the abiotic environment and biota in the general area may invade disturbed, seminatural communities. Invasion of successionally mature, undisturbed communities usually requires that the alien taxon overcomes a different category of barriers. We propose that the term ‘invasive’ should be used without any inference to environmental or economic impact. Terms like ‘pests’ and ‘weeds’ are suitable labels for the 50–80% of invaders that have harmful effects. About 10% of invasive plants that change the character, condition, form, or nature of ecosystems over substantial areas may be termed ‘transformers’.

The nuclear ribosomal internal transcribed spacer (ITS) region is the formal fungal barcode and in most cases the marker of choice for the exploration of fungal diversity in environmental samples. Two problems are particularly acute in the pursuit of satisfactory taxonomic assignment of newly generated ITS sequences: (i) the lack of an inclusive, reliable public reference data set and (ii) the lack of means to refer to fungal species, for which no Latin name is available in a standardized stable way. Here, we report on progress in these regards through further development of the UNITE database (http://unite.ut.ee) for molecular identification of fungi. All fungal species represented by at least two ITS sequences in the international nucleotide sequence databases are now given a unique, stable name of the accession number type (e.g. Hymenoscyphus pseudoalbidus|GU586904|SH133781.05FU), and their taxonomic and ecological annotations were corrected as far as possible through a distributed, third-party annotation effort. We introduce the term 'species hypothesis' (SH) for the taxa discovered in clustering on different similarity thresholds (97-99%). An automatically or manually designated sequence is chosen to represent each such SH. These reference sequences are released (http://unite.ut.ee/repository.php) for use by the scientific community in, for example, local sequence similarity searches and in the QIIME pipeline. The system and the data will be updated automatically as the number of public fungal ITS sequences grows. We invite everybody in the position to improve the annotation or metadata associated with their particular fungal lineages of expertise to do so through the new Web-based sequence management system in UNITE.

Biological invasions cause ecological and economic impacts across the globe. However, it is unclear whether there are strong patterns in terms of their major effects, how the vulnerability of different ecosystems varies and which ecosystem services are at greatest risk. We present a global meta-analysis of 199 articles reporting 1041 field studies that in total describe the impacts of 135 alien plant taxa on resident species, communities and ecosystems. Across studies, alien plants had a significant effect in 11 of 24 different types of impact assessed. The magnitude and direction of the impact varied both within and between different types of impact. On average, abundance and diversity of the resident species decreased in invaded sites, whereas primary production and several ecosystem processes were enhanced. While alien N-fixing species had greater impacts on N-cycling variables, they did not consistently affect other impact types. The magnitude of the impacts was not significantly different between island and mainland ecosystems. Overall, alien species impacts are heterogeneous and not unidirectional even within particular impact types. Our analysis also reveals that by the time changes in nutrient cycling are detected, major impacts on plant species and communities are likely to have already occurred.

Although research on human-mediated exchanges of species has substantially intensified during the last centuries, we know surprisingly little about temporal dynamics of alien species accumulations across regions and taxa. Using a novel database of 45,813 first records of 16,926 established alien species, we show that the annual rate of first records worldwide has increased during the last 200 years, with 37% of all first records reported most recently (1970-2014). Inter-continental and inter-taxonomic variation can be largely attributed to the diaspora of European settlers in the nineteenth century and to the acceleration in trade in the twentieth century. For all taxonomic groups, the increase in numbers of alien species does not show any sign of saturation and most taxa even show increases in the rate of first records over time. This highlights that past efforts to mitigate invasions have not been effective enough to keep up with increasing globalization.

Biological invasions are a global consequence of an increasingly connected world and the rise in human population size. The numbers of invasive alien species - the subset of alien species that spread widely in areas where they are not native, affecting the environment or human livelihoods - are increasing. Synergies with other global changes are exacerbating current invasions and facilitating new ones, thereby escalating the extent and impacts of invaders. Invasions have complex and often immense long-term direct and indirect impacts. In many cases, such impacts become apparent or problematic only when invaders are well established and have large ranges. Invasive alien species break down biogeographic realms, affect native species richness and abundance, increase the risk of native species extinction, affect the genetic composition of native populations, change native animal behaviour, alter phylogenetic diversity across communities, and modify trophic networks. Many invasive alien species also change ecosystem functioning and the delivery of ecosystem services by altering nutrient and contaminant cycling, hydrology, habitat structure, and disturbance regimes. These biodiversity and ecosystem impacts are accelerating and will increase further in the future. Scientific evidence has identified policy strategies to reduce future invasions, but these strategies are often insufficiently implemented. For some nations, notably Australia and New Zealand, biosecurity has become a national priority. There have been long-term successes, such as eradication of rats and cats on increasingly large islands and biological control of weeds across continental areas. However, in many countries, invasions receive little attention. Improved international cooperation is crucial to reduce the impacts of invasive alien species on biodiversity, ecosystem services, and human livelihoods. Countries can strengthen their biosecurity regulations to implement and enforce more effective management strategies that should also address other global changes that interact with invasions.

Plant traits-the morphological, anatomical, physiological, biochemical and phenological characteristics of plants-determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait-based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits-almost complete coverage for 'plant growth form'. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait-environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects. We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives.

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 km(2)) 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.

Ecologists and evolutionary biologists are increasingly using big-data approaches to tackle questions at large spatial, taxonomic, and temporal scales. However, despite recent efforts to gather two centuries of biodiversity inventories into comprehensive databases, many crucial research questions remain unanswered. Here, we update the concept of knowledge shortfalls and review the tradeoffs between generality and uncertainty. We present seven key shortfalls of current biodiversity data. Four previously proposed shortfalls pinpoint knowledge gaps for species taxonomy (Linnean), distribution (Wallacean), abundance (Prestonian), and evolutionary patterns (Darwinian). We also redefine the Hutchinsonian shortfall to apply to the abiotic tolerances of species and propose new shortfalls relating to limited knowledge of species traits (Raunkiæran) and biotic interactions (Eltonian). We conclude with a general framework for the combined impacts and consequences of shortfalls of large-scale biodiversity knowledge for evolutionary and ecological research and consider ways of overcoming the seven shortfalls and dealing with the uncertainty they generate.

1 An international group of scientists has built an open internet data base of life-history traits of the Northwest European flora (the LEDA-Traitbase) that can be used as a data source for fundamental research on plant biodiversity and coexistence, macro-ecological patterns and plant functional responses. 2 The species-trait matrix comprises referenced information under the control of an editorial board, for ca. 3000 species of the Northwest European flora, combining existing information and additional measurements. The data base currently contains data on 26 plant traits that describe three key features of plant dynamics: persistence, regeneration and dispersal. The LEDA-Traitbase is freely available at http://www.leda-traitbase.org. 3 We present the structure of the data base and an overview of the trait information available. 4 Synthesis. The LEDA Traitbase is useful for large-scale analyses of functional responses of communities to environmental change, effects of community trait composition on ecosystem properties and patterns of rarity and invasiveness, as well as linkages between traits as expressions of fundamental trade-offs in plants.

Abstract Aims Vegetation classification consistent with the Braun‐Blanquet approach is widely used in Europe for applied vegetation science, conservation planning and land management. During the long history of syntaxonomy, many concepts and names of vegetation units have been proposed, but there has been no single classification system integrating these units. Here we (1) present a comprehensive, hierarchical, syntaxonomic system of alliances, orders and classes of Braun‐Blanquet syntaxonomy for vascular plant, bryophyte and lichen, and algal communities of Europe; (2) briefly characterize in ecological and geographic terms accepted syntaxonomic concepts; (3) link available synonyms to these accepted concepts; and (4) provide a list of diagnostic species for all classes. Location European mainland, Greenland, Arctic archipelagos (including Iceland, Svalbard, Novaya Zemlya), Canary Islands, Madeira, Azores, Caucasus, Cyprus. Methods We evaluated approximately 10 000 bibliographic sources to create a comprehensive list of previously proposed syntaxonomic units. These units were evaluated by experts for their floristic and ecological distinctness, clarity of geographic distribution and compliance with the nomenclature code. Accepted units were compiled into three systems of classes, orders and alliances (EuroVegChecklist, EVC ) for communities dominated by vascular plants ( EVC 1), bryophytes and lichens ( EVC 2) and algae ( EVC 3). Results EVC 1 includes 109 classes, 300 orders and 1108 alliances; EVC 2 includes 27 classes, 53 orders and 137 alliances, and EVC 3 includes 13 classes, 24 orders and 53 alliances. In total 13 448 taxa were assigned as indicator species to classes of EVC 1, 2087 to classes of EVC 2 and 368 to classes of EVC 3. Accepted syntaxonomic concepts are summarized in a series of appendices, and detailed information on each is accessible through the software tool EuroVegBrowser. Conclusions This paper features the first comprehensive and critical account of European syntaxa and synthesizes more than 100 yr of classification effort by European phytosociologists. It aims to document and stabilize the concepts and nomenclature of syntaxa for practical uses, such as calibration of habitat classification used by the European Union, standardization of terminology for environmental assessment, management and conservation of nature areas, landscape planning and education. The presented classification systems provide a baseline for future development and revision of European syntaxonomy.

Abstract The number of studies dealing with plant invasions is increasing rapidly, but the accumulating body of knowledge has unfortunately also spawned increasing confusion about terminology. Invasions are a global phenomenon and comparison of geographically distant regions and their introduced biota is a crucially important methodological approach for elucidation of the determinants of invasiveness and invasibility. Comparative studies of alien floras provide substantial new insights to our understanding of general patterns of plant invasions. Such studies, using information in previously published floras and checklists, are fundamentally dependent on the quality of the assessment of particular species with respect to their taxonomic identity, time of immigration and invasion status. Three crucial decisions should be made when defining the status of a plant species in a given region: (1) whether the taxon is native or alien to that region (origin status); (2) what is its position in the invasion process, i.e., when was it introduced (residence status); and (3) what is the degree of its naturalization and possible invasion (invasion status). Standard floras differ hugely in their treatment of non­native species and those with appropriate categorization of alien species according to their status are rather rare. The present paper suggests definitions of terms associated with plant invasions and places these into the context of floras. Recommendations are outlined on how to deal with the issue of plant invasions in standard floras with the aim of contributing to a better understanding between taxonomists and ecologists and allowing more detailed comparative analyses of alien floras of various regions of the world.

Abstract With the growing body of literature assessing the impact of invasive alien plants on resident species and ecosystems, a comprehensive assessment of the relationship between invasive species traits and environmental settings of invasion on the characteristics of impacts is needed. Based on 287 publications with 1551 individual cases that addressed the impact of 167 invasive plant species belonging to 49 families, we present the first global overview of frequencies of significant and non‐significant ecological impacts and their directions on 15 outcomes related to the responses of resident populations, species, communities and ecosystems. Species and community outcomes tend to decline following invasions, especially those for plants, but the abundance and richness of the soil biota, as well as concentrations of soil nutrients and water, more often increase than decrease following invasion. Data mining tools revealed that invasive plants exert consistent significant impacts on some outcomes (survival of resident biota, activity of resident animals, resident community productivity, mineral and nutrient content in plant tissues, and fire frequency and intensity), whereas for outcomes at the community level, such as species richness, diversity and soil resources, the significance of impacts is determined by interactions between species traits and the biome invaded. The latter outcomes are most likely to be impacted by annual grasses, and by wind pollinated trees invading mediterranean or tropical biomes. One of the clearest signals in this analysis is that invasive plants are far more likely to cause significant impacts on resident plant and animal richness on islands rather than mainland. This study shows that there is no universal measure of impact and the pattern observed depends on the ecological measure examined. Although impact is strongly context dependent, some species traits, especially life form, stature and pollination syndrome, may provide a means to predict impact, regardless of the particular habitat and geographical region invaded.

Invasive species are a major element of global change and are contributing to biodiversity loss, ecosystem degradation, and impairment of ecosystem services worldwide. Research is shedding new light on the ecological and economic consequences of invasions. New approaches are emerging for describing and evaluating impacts of invasive species, and for translating these impacts into monetary terms. The harmful effects of invasions are now widely recognized, and multiscale programs are in place in many parts of the world to reduce current and future impacts. There has been an upsurge in scientific research aimed at guiding management interventions. Among the activities that are receiving the most attention and that have the most promise for reducing problems are risk assessment, pathway and vector management, early detection, rapid response, and new approaches to mitigation and restoration. Screening protocols to reduce new introductions are becoming more accurate and have been shown cost-effective.

The experiments have revealed that (NH4)2SO4 combined with KNO3 at low concentration is of advantage to the formation, growth and differentiation of pollen callus in rice, whereas the high concentration of (NH4)2SO4, whether used separately or in combination with KNO3, obviously inhibits the pollen callus formation. The optimum NH4+ concen- tration is about 7.0 mM (equal to 3.5 mM (NH4)2SO4). A basic medium containing 3.5 mM (NH4)2SO4 and 28 mM KNO3 as nitrogen sources has been established. On such medium the frequency of the pollen callus formation is higher than that on Millers me- dium, and the differentiation of shoot from pollen callus is satisfactory.

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.

Summary Much attention has been paid to negative effects of alien species on resident communities but studies that quantify community‐level effects of a number of invasive plants are scarce. We address this issue by assessing the impact of 13 species invasive in the Czech Republic on a wide range of plant communities. Vegetation in invaded and uninvaded plots with similar site conditions was sampled. All species of vascular plants were recorded, their covers were estimated and used as importance values for calculating the Shannon diversity index H′ , evenness J and Sørensen index of similarity between invaded and uninvaded vegetation. With the exception of two invasive species, species richness, diversity and evenness were reduced in invaded plots. Species exhibiting the greatest impact reduced species numbers per plot and the total number of species recorded in the communities sampled by almost 90%. A strong reduction of species number at the plot scale resulted in a marked reduction in the total species number at the landscape scale, and in less similarity between invaded and uninvaded vegetation. The decrease in species richness in invaded compared to uninvaded plots is largely driven by the identity of the invading species, whereas the major determinants of the decrease in Shannon diversity and evenness are the cover and height of invading species, and differences between height and cover of invading and dominant native species, independent of species identity. Synthesis. Management decisions based on impact need to distinguish between invasive species, as their effects on diversity and composition of resident vegetation differ largely. Tall invading species capable of forming populations with the cover markedly greater than that of native dominant species exert the most severe effects on species diversity and evenness. Since a strong impact on the community scale is associated with reduction in species diversity at higher scales, invaders with a high impact represent a serious hazard to the landscape.

Recent studies have shown that accounting for intraspecific trait variation (ITV) may better address major questions in community ecology. However, a general picture of the relative extent of ITV compared to interspecific trait variation in plant communities is still missing. Here, we conducted a meta-analysis of the relative extent of ITV within and among plant communities worldwide, using a data set encompassing 629 communities (plots) and 36 functional traits. Overall, ITV accounted for 25% of the total trait variation within communities and 32% of the total trait variation among communities on average. The relative extent of ITV tended to be greater for whole-plant (e.g. plant height) vs. organ-level traits and for leaf chemical (e.g. leaf N and P concentration) vs. leaf morphological (e.g. leaf area and thickness) traits. The relative amount of ITV decreased with increasing species richness and spatial extent, but did not vary with plant growth form or climate. These results highlight global patterns in the relative importance of ITV in plant communities, providing practical guidelines for when researchers should include ITV in trait-based community and ecosystem studies.

Research was conducted in July 2013 as part of anethnobotany research-training course in Georgia(საქართველო). After obtaining prior informed consent forinterviews and photographs, semi-structured interviewswere conducted with participants in local villages.Images were taken of the physical environment (soils, waterfeatures, and other geographical context elements), biologicalenvironment (parts of plants used, whole plants,animals, ecosystems, landscapes), and cultural environment(artifacts, villages, crop fields, and other anthropogeniclandscapes, individuals and groups interacting witheach other or the researchers, and people conducting dailytasks of life). Most of the photographs were taken opportunisticallyand only Figures 79-81 were “staged.”

Recent comprehensive data provided through the DAISIE project ( www.europe‐aliens.org ) have facilitated the development of the first pan‐European assessment of the impacts of alien plants, vertebrates, and invertebrates – in terrestrial, freshwater, and marine environments – on ecosystem services. There are 1094 species with documented ecological impacts and 1347 with economic impacts. The two taxonomic groups with the most species causing impacts are terrestrial invertebrates and terrestrial plants. The North Sea is the maritime region that suffers the most impacts. Across taxa and regions, ecological and economic impacts are highly correlated. Terrestrial invertebrates create greater economic impacts than ecological impacts, while the reverse is true for terrestrial plants. Alien species from all taxonomic groups affect “supporting”, “provisioning”, “regulating”, and “cultural” services and interfere with human well‐being. Terrestrial vertebrates are responsible for the greatest range of impacts, and these are widely distributed across Europe. Here, we present a review of the financial costs, as the first step toward calculating an estimate of the economic consequences of alien species in Europe.

Summary 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. 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. 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. 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. 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.