The paper provides the first estimate of the composition and structure of alien plants occurring in the wild in the European continent, based on the results of the DAISIE project (2004–2008), funded by the 6th Framework Programme of the European Union and aimed at “creating an inventory of invasive species that threaten European terrestrial, freshwater and marine environments”. The plant section of the DAISIE database is based on national checklists from 48 European countries/regions and Israel; for many of them the data were compiled during the project and for some countries DAISIE collected the first comprehensive checklists of alien species, based on primary data (e.g., Cyprus, Greece, F. Y. R. O. Macedonia, Slovenia, Ukraine). In total, the database contains records of 5789 alien plant species in Europe (including those native to a part of Europe but alien to another part), of which 2843 are alien to Europe (of extra-European origin). The research focus was on naturalized species; there are in total 3749 naturalized aliens in Europe, of which 1780 are alien to Europe. This represents a marked increase compared to 1568 alien species reported by a previous analysis of data in Flora Europaea (1964–1980). Casual aliens were marginally considered and are represented by 1507 species with European origins and 872 species whose native range falls outside Europe. The highest diversity of alien species is concentrated in industrialized countries with a tradition of good botanical recording or intensive recent research. The highest number of all alien species, regardless of status, is reported from Belgium (1969), the United Kingdom (1779) and Czech Republic (1378). The United Kingdom (857), Germany (450), Belgium (447) and Italy (440) are countries with the most naturalized neophytes. The number of naturalized neophytes in European countries is determined mainly by the interaction of temperature and precipitation; it increases with increasing precipitation but only in climatically warm and moderately warm regions. Of the nowadays naturalized neophytes alien to Europe, 50% arrived after 1899, 25% after 1962 and 10% after 1989. At present, approximately 6.2 new species, that are capable of naturalization, are arriving each year. Most alien species have relatively restricted European distributions; half of all naturalized species occur in four or fewer countries/regions, whereas 70% of non-naturalized species occur in only one region. Alien species are drawn from 213 families, dominated by large global plant families which have a weedy tendency and have undergone major radiations in temperate regions (Asteraceae, Poaceae, Rosaceae, Fabaceae, Brassicaceae). There are 1567 genera, which have alien members in European countries, the commonest being globally-diverse genera comprising mainly urban and agricultural weeds (e.g., Amaranthus, Chenopodium and Solanum) or cultivated for ornamental purposes (Cotoneaster, the genus richest in alien species). Only a few large genera which have successfully invaded (e.g., Oenothera, Oxalis, Panicum, Helianthus) are predominantly of non-European origin. Conyza canadensis, Helianthus tuberosus and Robinia pseudoacacia are most widely distributed alien species. Of all naturalized aliens present in Europe, 64.1% occur in industrial habitats and 58.5% on arable land and in parks and gardens. Grasslands and woodlands are also highly invaded, with 37.4 and 31.5%, respectively, of all naturalized aliens in Europe present in these habitats. Mires, bogs and fens are least invaded; only approximately 10% of aliens in Europe occur there. Intentional introductions to Europe (62.8% of the total number of naturalized aliens) prevail over unintentional (37.2%). Ornamental and horticultural introductions escaped from cultivation account for the highest number of species, 52.2% of the total. Among unintentional introductions, contaminants of seed, mineral materials and other commodities are responsible for 1091 alien species introductions to Europe (76.6% of all species introduced unintentionally) and 363 species are assumed to have arrived as stowaways (directly associated with human transport but arriving independently of commodity). Most aliens in Europe have a native range in the same continent (28.6% of all donor region records are from another part of Europe where the plant is native); in terms of species numbers the contribution of Europe as a region of origin is 53.2%. Considering aliens to Europe separately, 45.8% of species have their native distribution in North and South America, 45.9% in Asia, 20.7% in Africa and 5.3% in Australasia. Based on species composition, European alien flora can be classified into five major groups: (1) north-western, comprising Scandinavia and the UK; (2) west-central, extending from Belgium and the Netherlands to Germany and Switzerland; (3) Baltic, including only the former Soviet Baltic states; (4) east-central, comprizing the remainder of central and eastern Europe; (5) southern, covering the entire Mediterranean region. The clustering patterns cut across some European bioclimatic zones; cultural factors such as regional trade links and traditional local preferences for crop, forestry and ornamental species are also important by influencing the introduced species pool. Finally, the paper evaluates a state of the art in the field of plant invasions in Europe, points to research gaps and outlines avenues of further research towards documenting alien plant invasions in Europe. The data are of varying quality and need to be further assessed with respect to the invasion status and residence time of the species included. This concerns especially the naturalized/casual status; so far, this information is available comprehensively for only 19 countries/regions of the 49 considered. Collating an integrated database on the alien flora of Europe can form a principal contribution to developing a European-wide management strategy of alien species.
Comparative studies of closely related species may provide useful insights into the effect of species traits on invasion success since some of the biases associated with multispecies studies, such as phylogenetic effects, are considerably reduced by virtue of the experimental design. In this study seed and seedling traits of three congeneric alien species in Europe, differing in their region of origin, invasion status and history (Impatiens glandulifera, I. parviflora, I. capensis), were compared with the native I. noli-tangere in laboratory and common garden experiments. Seeds of I. glandulifera required the shortest period of stratification, germinated well both under laboratory and experimental garden conditions and the seedlings produced more biomass than those of the other species. Seeds of I. parviflora required a longer period of stratification, had the highest percentage germination but seedling emergence in the experimental garden was poorer than in I. glandulifera. Neither of these two species invasive in the Czech Republic formed soil seed banks. The native I. noli-tangere had the lowest percentage germination and formed a short-term persistent seed bank. Impatiens capensis germinated well in the laboratory, had the highest seedling emergence in the garden and its seed remained viable in the soil for three years. This indicates that in terms of germination and emergence, this species is comparable with the two invasive alien congeners and there appear to be no constraints to its invasion in the Czech Republic where it does not occur yet. Its absence may be due to a low propagule pressure; in the national flora I. capensis is listed as a potential future invader without mentioning it being cultivated in this country. Our results indicate that differences in the invasiveness of three alien species of balsams in the temperate zone of Central Europe can be attributed, at least in part, to their differing performances in the early stages of their life cycle. The short period of time required for seed stratification and the high seedling biomass of I. glandulifera might have increased its invasion potential compared to other Impatiens species occurring in the Czech Republic.
Heracleum mantegazzianum is one of the most invasive species in the Czech flora. The present study describes its flowering phenology and assess the effectiveness of protandry in preventing selfing in this self-compatible species, describes the timing of flowering in a heavily invaded area of Slavkovský les (Czech Republic) and estimates fruit set in a large sample of plants, which provides reliable data on the often exaggerated fecundity of this species. The study of flowering phenology revealed that protandry is always effective only within individual flowers, where male and female flowering phases are completely separated. In contrast, anther dehiscence in some flowers can occasionally overlap with stigma receptivity in other flowers in the same umbel, providing an opportunity for geitonogamous (i.e. between-flower) selfing. Nevertheless, the potential for selfing in H. mantegazzianum is determined mainly by an overlap in the male and female flowering phases between umbels on the same plant; at least a short overlap between some umbels was observed in 99% of the plants at the Slavkovský les. Although the degree of protandry in H. mantegazzianum favours outcrossing, the opportunity to self may be of crucial importance for an invasive plant, especially if a single plant colonizes a new location. At Slavkovský les, flowering started within one week (from 20 to 27 June 2002) at all 10 sites. The duration of flowering of an individual plantwas on average 36 days,with maximum of 60 days, and increased significantly with the number of umbels on a plant. In the second half of August, the majority of the fruits were ripe and had started to be shed. The beginning of flowering of a plant was significantly negatively correlated with the number of umbels it had – the earlier a plant started to flower the more umbels it had produced. A significant negative relationship was also found between basal diameter and beginning of flowering; plants with large basal diameters started to flower earlier. An average plant at Slavkovský les produced 20,671 fruits. Of these, 44.6% were produced by the terminal umbel, 29.3% by secondary umbels on satellites, 22.6% by secondary umbels on branches and only 3.5% by tertiary umbels. The estimated fruit number of the most fecund plant was 46,470 – compared to an average plant, the proportional contribution of tertiary umbels increased relative to the primary umbel. This study revealed a significant positive relationship between fecundity and plant basal diameter. Although the results of this study indicate that the fecundity of this species is often overestimated in the literature, the number of fruits produced by H. mantegazzianum provides this invasive species with an enormous reproductive capacity.
The Central European flora is an important source pool of plant species introduced to many regions throughout theworld. In this study,we identified a total of 759 plant species of the Central European flora that are currently recognized as alien species in Australia. We explored temporal patterns of introduction of these species to Australia in relation to method of introduction, growth form, naturalization status and taxonomy. Across all species, substantially larger numbers of species were introduced between 1840 and 1880 as well as between 1980 and the present, with a small peak of introductions within the 1930s. These patterns reflect early immigration patterns to Australia, recent improvements in fast and efficient transportation around the globe, and emigration away from difficult conditions brought about by the lead up to the Second World War respectively. We found that the majority of species had deliberate (69%) rather than accidental (31%) introductions and most species have not naturalized (66% casual species, 34% naturalized species). A total of 86 plant families comprising 31 tree species, 91 shrub species, 533 herbaceous species and 61 grass species present in Central Europe have been introduced to Australia. Differential patterns of temporal introduction of species were found as a function of both plant family and growth form and these patterns appear linked to variation in human migration numbers to Australia.
We investigated the effects of different temperature regimes and dry storage on germination of H. mantegazzianum (Apiaceae, native to Caucasus) seeds in the laboratory and linked the results with studies of seasonal seed bank depletion in a common garden experiment and under field conditions. Seeds were collected at seven sites in the Slavkovský les region, Czech Republic, cold-stratified for 2 months and germinated at seven temperature regimes. Under all temperature regimes, fresh seeds germinated to significantly higher percentages than older (1, 2, 3 years) seeds. For all storage lengths, seeds germinated best at alternating day/night temperatures of 20/5 °C. The length of the germination period had a significant effect only at low constant temperatures of 2 and 6 °C, where germination percentage increased between 2 and 6 months. Seasonal germination exhibited a distinct pattern, with rapid depletion of seed bank by the first spring after seed burial. Non-dormant seeds were present in the soil early in spring and late in autumn. The higher summer temperatures prevented dormancy breaking and another cold period of at least two months below 10 °C was needed to bring non-germinated seeds out of dormancy. The results suggest that (1) seed dormancy of H. mantegazzianum was not completely broken until the first spring, but that some seeds re-enter or retain dormancy during high summer temperatures and that (2) the threshold needed for breaking the dormancy was achieved gradually during the cold autumn and winter months. However, in a small fraction of seeds the dormancy breaking process took several years. Of seeds buried in 10 different regions of the Czech Republic, on average 8.8% survived 1 year, 2.7% 2 years and 1.2% remained viable and dormant after 3 years of burial. The ability of even small fraction of H. mantegazzianum seeds to survive for at least 3 years can result in re-invasion of this species into controlled sites.