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.
Density dependent host mortality in the interaction between the solitary endoparasitoid Aphidius colemani (Aphidiidae), and its host, the green peach aphid Myzus persicae, was examined on greenhouse peppers. The experimental approach attempted to eliminate spatial interdependence in the relationship between host density and host mortality by using different plants to measure parasitism at different spatial scales. Increasing host density at the plant scale caused a significant increase in the proportion of host mortality. However, at the shoot and leaf scale, increasing host density caused a significant decrease in host mortality. This may be one of the first experimental demonstrations of a switch from inverse to direct density dependence. The pattern is assumed to be a result of searching parasitoids using different cues at different spatial scales.
A complete list of all alien taxa ever recorded in the flora of the Czech Republic is presented as an update of the original checklist published in 2002. New data accumulated in the last decade are incorporated and the listing and status of some taxa are reassessed based on improved knowledge. Alien flora of the Czech Republic consists of 1454 taxa listed with information on their taxonomic position, life history, geographic origin (or mode of origin, distinguishing anecophyte and hybrid), invasive status (casual; naturalized but not invasive; invasive), residence time status (archaeophyte vs neophyte), mode of introduction into the country (accidental, deliberate), and date of the first record. Additional information on species performance that was not part of the previous catalogue, i.e. on the width of species’ habitat niches, their dominance in invaded communities, and impact, is provided. The Czech alien flora consists of 350 (24.1%) archaeophytes and 1104 (75.9%) neophytes. The increase in the total number of taxa compared to the previous catalogue (1378) is due to addition of 151 taxa and removal of 75 (39 archaeophytes and 36 neophytes), important part of the latter being the reclassification of 41 taxa as native, mostly based on archaeobotanical evidence. The additions represent taxa newly recorded since 2002 and reported in the national literature; taxa resulting from investigation of sources omitted while preparing the previous catalogue; redetermination of previously reported taxa; reassessment of some taxa traditionally considered native for which the evidence suggests the opposite; and inclusion of intraspecific taxa previously not recognized in the flora. There are 44 taxa on the list that are reported in the present study for the first time as aliens introduced to the Czech Republic or escaped from cultivation: Abies concolor, A. grandis, A. nordmanniana, Avena sterilis subsp. ludoviciana, A. ×vilis, Berberis julianae, B. thunbergii, Bidens ferulifolius, Buddleja alternifolia, Buglossoides incrassata subsp. splitgerberi, Buxus sempervirens, Corispermum declinatum, Cotoneaster dielsianus, C. divaricatus, Euphorbia myrsinites, Gleditsia triacanthos, Helleborus orientalis, Hieracium heldreichii, Koelreuteria paniculata, Lonicera periclymenum, Lotus ornithopodioides, Malus baccata, M. pumila, Miscanthus sacchariflorus, Morus alba, Muscari armeniacum, Paeonia lactiflora, Pennisetum alopecuroides, Pinguicula crystallina subsp. hirtiflora, P. grandiflora subsp. rosea, Podophyllum hexandrum, Pyracantha coccinea, Rhodotypos scandens, Rumex patientia × R. tianschanicus ‘Uteuša’, Salix cordata, Sarracenia purpurea, Sasa palmata ‘Nebulosa’, Scolymus maculatus, Spiraea japonica, Tagetes tenuifolia, Thuja occidentalis, Trifolium badium, Vaccinium corymbosum and Viburnum rhytidophyllum. All added and deleted taxa are commented on. Of the total number of taxa, 985 are classified as casuals, 408 as naturalized but not invasive, and 61 as invasive. The reduction in the number of invasive taxa compared to the previous catalogue is due to a more conservative approach adopted here; only taxa that currently spread are considered invasive. Casual taxa are strongly overrepresented among neophytes compared to archaeophytes (76.7% vs 39.4%), while naturalized but non-invasive taxa follow the reversed pattern (18.8% vs 57.4). However, these two groups do not significantly differ in the proportion of invasive taxa. Of introduced neophytes, 250 taxa (22.6%) are considered vanished, i.e. no longer present in the flora, while 23.3% became naturalized, and 4.5% invasive. In addition to the traditional classification based on introduction–naturalization–invasion continuum, taxa were classified into 18 population groups based on their long-term trends in metapopulation dynamics in the country, current state of their populations, and link to the propagule pressure from cultivation. Mapping these population groups onto the unified framework for biological invasions introduced by Blackburn et al. in 2011 made it possible to quantify invasion failures, and boom-and-busts, in the Czech alien flora. Depending on inclusion criteria (whether or not extinct/vanished taxa and hybrids are considered), alien taxa ever recorded in the Czech Republic contribute 29.7–33.1% to the total country’s plant diversity; taking into account only naturalized taxa, a permanent element of the country’s flora, the figure is 14.4–17.5%. Analysis of the dates of the first record, known for 771 neophytes, indicates that alien taxa in the flora have been increasing at a steady pace without any distinct deceleration trend; by extrapolating this data to all 1104 neophytes recorded it is predicted that the projected number would reach 1264 in 2050. Deliberate introduction was involved in 747 cases (51.4%), the remaining 48.6% of taxa are assumed to have arrived by unintentional pathways. Archaeophytes are more abundant in landscapes, occupy on average a wider range of habitat types than neophytes, but reach a lower cover in plant communities. The alien flora is further analysed with respect to representation of genera and families, origin and life history. and Nevejdou se dvě poslední jména autorů
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.
The development stages of a species may have an identical lower development threshold (LDT) and proportionally different durations. This phenomenon called "rate isomorphy" (RI) has been demonstrated for a number of insect species. In contrast, the growing day degrees accumulated over the period of larval development (sum of effective temperatures SET) should be plastic and vary with environment conditions. The prediction from RI is that, with changing conditions, the uniform LDT should be accompanied by differences in development time which remain proportional at different temperatures. This was tested by investigating the effect of diet on thermal requirements for development of larvae of the polyphagous species Autographa gamma (L.) (Lepidoptera: Noctuidae). The larvae were kept at 15.0, 20.3 and 26.7°C and fed on leaves of 13dicotyledoneous herb and tree species. The proportion of total development time spent on a particular diet was plotted against temperature. The existence of RI was inferred from a zero change in development time proportion with changing temperature. This rigorous test supported RI for 3 of 9 diets where development was completed in all temperatures. The LDT observed on 11 diets where the larvae completed development in at least 2 temperatures varied between 9.3 and 11.0°C while SET varied between 167 and 353 day degrees (dd). Assuming RI, LDT and SET for those 9 diets were recalculated. The recalculated LDT was 10.0°C and SET varied between 177-257 dd. The SET increased with decreasing water content and decreasing nitrogen content of food. Worsening food quality decreased food consumption, metabolic and food conversion efficiency, and the relative growth rate of the larvae. Increasing metabolic costs of development were thus positively correlated with SET. The standardized rate of growth (mg.dd-1) was typical for particular diets. Pupal mass decreased with increasing temperature and, within each temperature, with development length.
Genome size has been suggested as one of the traits associated with invasiveness of plant species. To provide a quantitative insight into the role of this trait, we estimated nuclearDNAcontent in 93 alien species naturalized in the Czech Republic, belonging to 32 families, by using flow cytometry, and compared it with the values reported for non-invading congeneric and confamilial species from the Plant DNA C-values database. Species naturalized in the Czech Republic have significantly smaller genomes than their congeners not known to be naturalized or invasive in any part of the world. This trend is supported at the family level: alien species naturalized in the Czech flora have on average a smaller genome than is the mean value for non-invading confamilials. Moreover, naturalized and non-invading species clearly differed in the frequency of five genome size categories; this difference was mainly due to very small genomes prevailing and intermediate to very large genomes underrepresented in the former group. Our results provide the first quantitative support for association of genome size with invasiveness, based on a large set of alien species across a number of plant families. However, there was no difference in the genome size of invasive species compared to naturalized but non-invasive. This suggests that small genome size provides alien plants with an advantage already at the stage of naturalization and need not be necessarily associated with the final stage of the process, i.e. invasion.
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.
This paper describes the reproductive characteristics of 93 neophytes (alien species introduced after 1500 A.D.) of the flora of the Czech Republic and compares trait values between naturalized invasive and naturalized non-invasive neophytes. Species were sampled and seed collected in the field from multiple localities in the Czech Republic. Traits related to seed production (propagule number per plant and per population), dispersal (propagule size, length/width ratio and weight; buoyancy; epizoochory; terminal velocity) and establishment (germination; seedling relative growth rate; seedling establishment) were measured for each species either in the field, in a common garden experiment or in the laboratory. Invasive species significantly differ from naturalized non-invasive species in propagule length/width ratio (by having lower ratio, i.e. more rounded propagules) and fecundity (invasive species are more fecund, both per individual plant and in terms of the population propagule production). Invasive species have proportionally fewer seedlings establishing in the autumn and better capacity for dispersal by wind than non-invasive species. The results for several traits differ depending on whether or not the effect of phylogeny is included in analytical models. Considering species relatedness expressed as a taxonomic hierarchy, invasive species have lighter propagules and higher population propagule numbers, and marginally significantly differ in producing more propagules per plant and having higher capacity for dispersal bywater.We found that most variation in invasiveness is linked to variation among species within genera. This distribution of relatedness means that predictions of whether a species will become invasive cannot be based on traits of the relatives of the given species at higher taxonomic levels. The distinction made in this paper, i.e. invasive species vs. naturalized but non-invasive species, can potentially contribute to a deeper understanding of the role of traits associated with invasiveness because the crucial transition from the naturalized to invasion stage is rarely addressed in invasion ecology.
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.