A determination of net effects from minimal adequate models.

Comparison of the number of archaeophytes, neophytes, and native species in different crop types.

To better understand the effect of species traits on plant invasion, we collected comparative data on 20 reproductive and dispersal traits of 93 herbaceous alien species in the Czech Republic, central Europe, introduced after 1500 A. D. We explain plant invasion success, expressed by two measures: invasiveness, i.e. whether the species is naturalized but non-invasive, or invasive; and dominance in plant communities expressed as the mean cover in vegetation plots. We also tested how important reproductive and dispersal traits are in models including other characteristics generally known to predict invasion outcome, such as plant height, life history and residence time. By using regression/classification trees we show that the biological traits affect invasion success at all life stages, from reproduction (seed production) to dispersal (propagule properties), and the ability to compete with resident species (height). By including species traits information not usually available in multispecies analyses, we provide evidence that traits do play important role in determining the outcome of invasion and can be used to distinguish between alien species that reach the final stage of the invasion process and dominate the local communities from those that do not. No effect of taxonomy ascertained in regression and classification trees indicates that the role of traits in invasiveness should be assessed primarily at the species level.

We tested whether two basic thermal requirements for insect development, lower developmental thresholds, i.e. temperatures at which development ceases, and sums of effective temperatures, i.e. numbers of day degrees above the lower developmental thresholds necessary to complete development, differ among insect species that proved to be successful invaders in regions outside their native range and those that did not. Focusing on species traits underlying invasiveness that are related to temperature provides insights into the mechanisms of insect invasions. The screening of thermal requirements thus could improve risk-assessment schemes by incorporating these traits in predictions of potentially invasive insect species. We compared 100 pairs of taxonomically-related species originating from the same continent, one invasive and the other not reported as invasive. Invasive species have higher lower developmental thresholds than those never recorded outside their native ranges. Invasive species also have a lower sum of effective temperatures, though not significantly. However, the differences between invasive and non-invasive species in the two physiological measures were significantly inversely correlated. This result suggests that many species are currently prevented from invading by low temperatures in some parts of the world. Those species that will overcome current climatic constraints in regions outside their native distribution due to climate change could become even more serious future invaders than present-day species, due to their potentially faster development.

This chapter provides guidance on mapping risks posed by invasive alien species to support pest risk analysis (PRA), the process required to justify phytosanitary measures. Because pest risk mapping can be challenging and resource-intensive, the situations in which risk maps are particularly useful are highlighted. The procedures described focus on mapping areas where the pest can establish and potentially cause the greatest harm. In the first stage of risk mapping, the factors that might influence the potential distribution and impacts of an invasive alien species are identified and the data are assembled and mapped. In the second stage, the maps of each factor are combined using matrix rules to generate areas of potential establishment and highest risk. These general procedures are illustrated with two examples. Risk maps for the western corn rootworm, a maize pest that has invaded Europe, are based on the combination of maps of climatic suitability, the presence of sandy soils, the distribution of grain and forage maize and the value of these commodities in Europe. Uncertainty is estimated by varying the classification of climatic suitability to obtain the worst, best and most likely scenarios. Risk maps for the common water hyacinth, an invasive plant on the Iberian Peninsula, are based on maps of climatic suitability, the distribution of suitable wetland habitats and areas of conservation importance. The chapter concludes by summarizing some of the major challenges that remain to enhance the production of risk maps for PRA and their interpretation by risk managers.

Studies of plant invasions rarely address impacts on molluscs. By comparing pairs of invaded and corresponding uninvaded plots in 96 sites in floodplain forests, we examined effects of four invasive alien plants (Impatiens glandulifera, Fallopia japonica, F. sachalinensis, and F.× bohemica) in the Czech Republic on communities of land snails. The richness and abundance of living land snail species were recorded separately for all species, rare species listed on the national Red List, and small species with shell size below 5 mm. The significant impacts ranged from 16-48% reduction in snail species numbers, and 29-90% reduction in abundance. Small species were especially prone to reduction in species richness by all four invasive plant taxa. Rare snails were also negatively impacted by all plant invaders, both in terms of species richness or abundance. Overall, the impacts on snails were invader-specific, differing among plant taxa. The strong effect of I. glandulifera could be related to the post-invasion decrease in abundance of tall nitrophilous native plant species that are a nutrient-rich food source for snails in riparian habitats. Fallopia sachalinensis had the strongest negative impact of the three knotweeds, which reflects differences in their canopy structure, microhabitat humidity and litter decomposition. The ranking of Fallopia taxa according to the strength of impacts on snail communities differs from ranking by their invasiveness, known from previous studies. This indicates that invasiveness does not simply translate to impacts of invasion and needs to be borne in mind by conservation and management authorities.

The so-called BIOCLIM variables have played a central role in the advancement of ecologicalmodellingmethods for correlative species distribution modelling. We propose to establish a register of the BIOCLIMvariables, allowing for the extension of the suite of described, readily available covariate data tosupport bioclimatic modelling. The registry will provide a simple means for researchers to add usefuldata sets in a transparent, documented manner. As a case study, we introduce and describe the set offivemost significant principal components of the first 35 BIOCLIM variables (Bio01 to Bio35) and proposethey be assigned as Bio36 to Bio40.Together, these five PCA variables capture more than 90%of thevariability in the full suite of 35 BIOCLIM variables. The PCA variables may provide a means to explore the climaticlimits of poorly known species, with a reduced risk of overfitting the models. The BIOCLIM variable registry should facilitate the expansion of the suite of variables commonly used in species distribution modelling.

Much of our current understanding of the impact of invasive species on plant communities is based on patterns occurring in the above-ground vegetation, while only few studies have examined changes in soil seed banks associated with plant invasions, despite their important role as determinants of vegetation dynamics. Here, we reviewed the literature on the impact of plant invasions on the seed bank and we provide a quantitative synthesis using a meta-analysis approach. Specifically, (1) we quantified the impact of 18 invasive alien plants on (i) species richness and (ii) density of the seed banks of invaded communities, based on 58 pair-wise invaded-uninvaded comparisons (cases); we identified (2) the invasive taxa that are responsible for the largest changes in the seed bank; and (3) the habitats where substantial changes occur. Our study showed three major findings: (1) species richness (68% of cases) and density (58% of cases) were significantly lower in native seed banks invaded by alien plants; (2) species richness and density of native and alien species were remarkably lower in seed banks invaded by large, perennial herbs compared to uninvaded sites; and (3) invaded seed banks were often associated with a larger richness and/or abundance of alien species. This study indicates a need for additional seed bank data in invasion ecology to characterize species-specific and habitat-specific impacts of plant invasions, and to determine whether changes in the seed banks of native and alien species are a symptom of environmental degradation prior to a plant invasion or whether they are its direct result. The findings of this study help improve our capacity to predict the long-term implications of plant invasions, including limitations in the recruitment of native species from the seed bank and the potential for secondary invasions by seeds of other alien species.

Many woody plants have been recently recognized as major invasive species with serious impacts on species diversity and functioning of invaded ecosystems. We evaluated the global invasion success of temperate trees and shrubs with native ranges in central Europe and explored the role of their native distribution and that of biological traits in determining whether they have become invasive in other parts of the world. Of the 94 species forming the source-area species pool, 27 % are known to be invasive in at least one region of the world. Generalized linear models on phylogenetically non-informed analyses revealed that tall woody plants flowering early in the season, and occupying many grid cells in the native range are significantly more likely to become successful invaders than species not possessing these traits. However, other traits can partly reduce the disadvantages resulting from low regional frequencies in the native range and consequent lower probability of them being introduced elsewhere. Species that do not depend for reproduction on another individual, those pollinated by wind and dispersed by animals are likely to become invasive even if they do not have extensive native distributions. However, of these traits only the regional frequency in the native range remained significant when phylogenetic relationships were taken into account. This indicates that the effect of the regional frequency is generic, valid across all woody species, and fine-tuned by advantageous biological traits inherited from common ancestors, shared by groups of phylogenetically-related species. Traits that only appeared significant in the phylogenetically non-informed analyses should be considered as specific for shrubs and trees of central Europe. Mode of reproduction was only significant in the phylogenetically-informed analysis, indicating that reproduction exclusively by seed favours invasiveness of woody species. From the management perspective, however, the predictive value of some traits is not diminished by them being phylogenetically constrained since we are not interested whether the behaviour of particular species is a result of evolutionary history but how we can treat specific cases of invasion.

There is a controversy about whether the thermal constants, lower developmental threshold, rate of development and corresponding degree days required for development, change when a species is reared under different developmental conditions. We present a more precise way of measuring these constants using the linear relationship between the rate of development and temperature. First we use the equation proposed by Ikemoto and Takai (2000) to determine the linear phase of development and then a generalised linear model having a different variance at low and high temperatures, specific for each condition, to estimate the parameters of the linear relationship. Using this method, we show that providing the difference in food quality is sufficiently great, an aphidophagous ladybird develops significantly faster and starts developing at a significantly lower temperature on a good than on a poor quality diet. Adaptive significance of the thermal constants not remaining constant is discussed in terms of a trade-off between growth and rate of development, when temperature and food quality varies.

Many alien plants are thought to be invasive because of unique traits and greater phenotypic plasticity relative to resident species. However, many studies of invasive species are unable to quantify the importance of particular traits and phenotypic plasticity in conferring invasive behavior because traits used in comparative studies are often measured in a single environment and by using plants from a single population. To obtain a deeper insight into the role of environmental factors, local differences and competition in plant invasions, we compared species of Impatiens (Balsaminaceae) of different origin and invasion status that occur in central Europe: native I. noli-tangere and three alien species (highly invasive I. glandulifera, less invasive I. parviflora and potentially invasive I. capensis). In two experiments we harvested late-stage reproductive plants to estimate performance. The first experiment quantified how populations differed in performance under varying light and moisture levels in the absence of competition. The second experiment quantified performance across these environments in the presence of intra- and inter-specific competition. The highly invasive I. glandulifera was the strongest competitor, was the tallest and produced the greatest biomass. Small size and high plasticity were characteristic for I. parviflora. This species appeared to be the second strongest competitor, especially under low soil moisture. The performance of I. capensis was within the range of the other Impatiens species studied, but sometimes limited by alien competitors. Our results suggest that invasion success within the genus Impatiens depends on the ability to grow large under a range of environmental conditions, including competition. The invasive species also exhibited greater phenotypic plasticity across environmental conditions than the native species. Finally, the decreased performance of the native I. noli-tangere in competition with other species studied indicates that this species may be possibly excluded from its sites by invading congeners.

Quantitative assessments of alien plant impacts are essential to inform management to ensure that resources are prioritized against the most problematic species and that restoration targets the worst-affected ecosystem processes. Here, we present the first detailed critique of quantitative field studies of alien plant impacts and highlight biases in the biogeography and life form of the target species, the responses assessed, and the extent to which spatial variability is addressed. Observed impacts often fail to translate to ecosystem services or evidence of environmental degradation. The absence of overarching hypotheses regarding impacts has reduced the consistency of approaches worldwide and prevented the development of predictive tools. Future studies must ensure that the links between species traits, ecosystem stocks, and ecosystem flows, as well as ecosystem services, are explicitly defined.

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.

Species gain membership of regional assemblages by passing through multiple ecological and environmental filters. To capture the potential trajectory of structural changes in regional meta-communities driven by biological invasions, one can categorize species pools into assemblages of different residence times. Older assemblages, having passed through more environmental filters, should become more functionally ordered and structured. Here we calculate the level of compartmentalization (modularity) for three different-aged assemblages (neophytes, introduced after 1500 AD; archaeophytes, introduced before 1500 AD, and natives), including 2,054 species of vascular plants in 302 reserves in central Europe. Older assemblages are more compartmentalized than younger ones, with species composition, phylogenetic structure and habitat characteristics of the modules becoming increasingly distinctive. This sheds light on two mechanisms of how alien species are functionally incorporated into regional species pools: the settling-down hypothesis of diminishing stochasticity with residence time, and the niche-mosaic hypothesis of inlaid neutral modules in regional meta-communities.

Alien plants pose significant threats to protected areas worldwide yet many studies only describe the degree to which these areas have become invaded. Research must move toward a better understanding of alien plant impacts since managers urgently require an appropriate evidence base to prioritize control/eradication targets. We analyze a global database of quantitative studies of alien plant impacts to evaluate existing knowledge of alien plant impacts within and outside protected areas. Although protected areas are a significant focus for quantitative impact studies, the biogeographic emphasis of most research effort does not coincide with the global distribution of protected areas nor the plant species or life‐forms recognized to have greatest impacts on ecosystems. While impacts were often as significant within protected areas as outside, only a minority of studies provide any subsequent management recommendations. There is therefore considerable scope to improve the evidence base on alien plant management in protected areas.

A fundamental assumption in invasion biology is that most invasive species exhibit enhanced performance in their introduced range relative to their home ranges. This idea has given rise to numerous hypotheses explaining "invasion success" by virtue of altered ecological and evolutionary pressures. There are surprisingly few data, however, testing the underlying assumption that the performance of introduced populations, including organism size, reproductive output, and abundance, is enhanced in their introduced compared to their native range. Here, we combined data from published studies to test this hypothesis for 26 plant and 27 animal species that are considered to be invasive. On average, individuals of these 53 species were indeed larger, more fecund, and more abundant in their introduced ranges. The overall mean, however, belied significant variability among species, as roughly half of the investigated species (N=27) performed similarly when compared to conspecific populations in their native range. Thus, although some invasive species are performing better in their new ranges, the pattern is not universal, and just as many are performing largely the same across ranges.

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 and managing 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 explore how this 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.