Europe's other debt crisis caused by the long legacy of future extinctions.

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.

Socioeconomic legacy yields an invasion debt.

Globalization and economic growth are widely recognized as important drivers of biological invasions. Consequently, there is an increasing need for governments to address the role of international trade in their strategies to prevent species introductions. However, many of the most problematic alien species are not recent arrivals but were introduced several decades ago. Hence, current patterns of alien-species richness may better reflect historical rather than contemporary human activities, a phenomenon which might be called "invasion debt." Here, we show that across 10 taxonomic groups (vascular plants, bryophytes, fungi, birds, mammals, reptiles, amphibians, fish, terrestrial insects, and aquatic invertebrates) in 28 European countries, current numbers of alien species established in the wild are indeed more closely related to indicators of socioeconomic activity from the year 1900 than to those from 2000, although the majority of species introductions occurred during the second half of the 20th century. The strength of the historical signal varies among taxonomic groups, with those possessing good capabilities for dispersal (birds, insects) more strongly associated with recent socioeconomic drivers. Nevertheless, our results suggest a considerable historical legacy for the majority of the taxa analyzed. The consequences of the current high levels of socioeconomic activity on the extent of biological invasions will thus probably not be completely realized until several decades into the future.

Do invasive species perform better in their new ranges?

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.

Disentangling the role of environmental and human pressures on biological invasions across Europe.

The accelerating rates of international trade, travel, and transport in the latter half of the twentieth century have led to the progressive mixing of biota from across the world and the number of species introduced to new regions continues to increase. The importance of biogeographic, climatic, economic, and demographic factors as drivers of this trend is increasingly being realized but as yet there is no consensus regarding their relative importance. Whereas little may be done to mitigate the effects of geography and climate on invasions, a wider range of options may exist to moderate the impacts of economic and demographic drivers. Here we use the most recent data available from Europe to partition between macroecological, economic, and demographic variables the variation in alien species richness of bryophytes, fungi, vascular plants, terrestrial insects, aquatic invertebrates, fish, amphibians, reptiles, birds, and mammals. Only national wealth and human population density were statistically significant predictors in the majority of models when analyzed jointly with climate, geography, and land cover. The economic and demographic variables reflect the intensity of human activities and integrate the effect of factors that directly determine the outcome of invasion such as propagule pressure, pathways of introduction, eutrophication, and the intensity of anthropogenic disturbance. The strong influence of economic and demographic variables on the levels of invasion by alien species demonstrates that future solutions to the problem of biological invasions at a national scale lie in mitigating the negative environmental consequences of human activities that generate wealth and by promoting more sustainable population growth.

Synergy between pathogen release and resource availability in plant invasion.

Why do some exotic plant species become invasive? Two common hypotheses, increased resource availability and enemy release, may more effectively explain invasion if they favor the same species, and therefore act in concert. This would be expected if plant species adapted to high levels of available resources in their native range are particularly susceptible to enemies, and therefore benefit most from a paucity of enemies in their new range. We tested this possibility by examining how resource adaptations influence pathogen richness and release among 243 European plant species naturalized in the United States. Plant species adapted to higher resource availability hosted more pathogen species in their native range. Plants from mesic environments hosted more fungi than plants from xeric environments, and plants from nitrogen-rich environments hosted more viruses than plants from nitrogen-poor environments. Furthermore, plants classified as competitors hosted more than 4 times as many fungi and viruses as did stress tolerators. Patterns of enemy release mirrored those of pathogen richness: competitors and species from mesic and nitrogen-rich environments were released from many pathogen species, while stress tolerators and species from xeric and nitrogen-poor environments were released from relatively few pathogen species. These results suggest that enemy release contributes most to invasion by fast-growing species adapted to resource-rich environments. Consequently, enemy release and increases in resource availability may act synergistically to favor exotic over native species.

Ecological impacts of invasive alien plants: a meta-analysis of their effects on species, communities and ecosystems.

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.

Protected-area boundaries as filters of plant invasions.

Human land uses surrounding protected areas provide propagules for colonization of these areas by non-native species, and corridors between protected-area networks and drainage systems of rivers provide pathways for long-distance dispersal of non-native species. Nevertheless, the influence of protected-area boundaries on colonization of protected areas by invasive non-native species is unknown. We drew on a spatially explicit data set of more than 27,000 non-native plant presence records for South Africa's Kruger National Park to examine the role of boundaries in preventing colonization of protected areas by non-native species. The number of records of non-native invasive plants declined rapidly beyond 1500 m inside the park; thus, we believe that the park boundary limited the spread of non-native plants. The number of non-native invasive plants inside the park was a function of the amount of water runoff, density of major roads, and the presence of natural vegetation outside the park. Of the types of human-induced disturbance, only the density of major roads outside the protected area significantly increased the number of non-native plant records. Our findings suggest that the probability of incursion of invasive plants into protected areas can be quantified reliably.

Developmental database for phenology models: related insect and mite species have similar thermal requirements.

Two values of thermal requirements, the lower developmental threshold (LDT), that is, the temperature at which development ceases, and the sum of effective temperatures, that is, day degrees above the LDT control the development of ectotherms and are used in phenology models to predict time at which the development of individual stages of a species will be completed. To assist in the rapid development of phenology models, we merged a previously published database of thermal requirements for insects, gathered by online search in CAB Abstracts, with independently collected data for insects and mites from original studies. The merged database comprises developmental times at various constant temperatures on 1,054 insect and mite species, many of them in several populations, mostly pests and their natural enemies, from all over the world. We show that closely related species share similar thermal requirements and therefore, for a species with unknown thermal requirements, the value of LDT and sum of effective temperatures of its most related species from the database can be used.

Controls on pathogen species richness in plants' introduced and native ranges: roles of residence time, range size and host traits.

Introduced species escape many pathogens and other enemies, raising three questions. How quickly do introduced hosts accumulate pathogen species? What factors control pathogen species richness? Are these factors the same in the hosts' native and introduced ranges? We analysed fungal and viral pathogen species richness on 124 plant species in both their native European range and introduced North American range. Hosts introduced 400 years ago supported six times more pathogens than those introduced 40 years ago. In hosts' native range, pathogen richness was greater on hosts occurring in more habitat types, with a history of agricultural use and adapted to greater resource supplies. In hosts' introduced range, pathogen richness was correlated with host geographic range size, agricultural use and time since introduction, but not any measured biological traits. Introduced species have accumulated pathogens at rates that are slow relative to most ecological processes, and contingent on geographic and historic circumstance.

Planting intensity, residence time, and species traits determine invasion success of alien woody species.

We studied the relative importance of residence time, propagule pressure, and species traits in three stages of invasion of alien woody plants cultivated for about 150 years in the Czech Republic, Central Europe. The probability of escape from cultivation, naturalization, and invasion was assessed using classification trees. We compared 109 escaped-not-escaped congeneric pairs, 44 naturalized-not-naturalized, and 17 invasive-not-invasive congeneric pairs. We used the following predictors of the above probabilities: date of introduction to the target region as a measure of residence time; intensity of planting in the target area as a proxy for propagule pressure; the area of origin; and 21 species-specific biological and ecological traits. The misclassification rates of the naturalization and invasion model were low, at 19.3% and 11.8%, respectively, indicating that the variables used included the major determinants of these processes. The probability of escape increased with residence time in the Czech Republic, whereas the probability of naturalization increased with the residence time in Europe. This indicates that some species were already adapted to local conditions when introduced to the Czech Republic. Apart from residence time, the probability of escape depends on planting intensity (propagule pressure), and that of naturalization on the area of origin and fruit size; it is lower for species from Asia and those with small fruits. The probability of invasion is determined by a long residence time and the ability to tolerate low temperatures. These results indicate that a simple suite of factors determines, with a high probability, the invasion success of alien woody plants, and that the relative role of biological traits and other factors is stage dependent. High levels of propagule pressure as a result of planting lead to woody species eventually escaping from cultivation, regardless of biological traits. However, the biological traits play a role in later stages of invasion.

Separating habitat invasibility by alien plants from the actual level of invasion.

Habitats vary considerably in the level of invasion (number or proportion of alien plant species they contain), which depends on local habitat properties, propagule pressure, and climate. To determine the invasibility (susceptibility to invasions) of different habitats, it is necessary to factor out the effects of any confounding variables such as propagule pressure and climate on the level of invasion. We used 20 468 vegetation plots from 32 habitats in the Czech Republic to compare the invasibility of different habitats. Using regression trees, the proportion of alien plants, including archaeophytes (prehistoric to medieval invaders) and neophytes (recent invaders), was related to variables representing habitat properties, propagule pressure, and climate. The propagule pressure was expressed as the proportion of surrounding urban and industrial or agricultural land, human population density, distance from a river, and history of human colonization in the region. Urban and industrial land use had a positive effect on the proportion of both archaeophytes and neophytes. Agricultural land use, higher population density, and longer history of human impact positively affected the proportion of archaeophytes. Disturbed human-made habitats with herbaceous vegetation were most invaded by both groups of aliens. Neophytes were also relatively common in disturbed woody vegetation, such as broad-leaved plantations, forest clearings, and riverine scrub. These habitats also had the highest proportion of aliens after removing the effect of propagule pressure and climate, indicating that they are not only the most invaded, but also most invasible. These habitats experience recurrent disturbances and are rich, at least temporarily, in available nutrients, which supports the hypothesis that fluctuating resources are the major cause of habitat invasibility. The least invaded habitats were mires and alpine-subalpine grasslands and scrub. After removing the effect of propagule pressure and climate, some habitats actually invaded at an intermediate level had very low proportions of aliens. This indicates that these habitats (e.g., dry, wet, and saline grasslands, base-rich fens, and broad-leaved deciduous woodlands) are resistant to invasion.

Getting the right traits: reproductive and dispersal characteristics predict the invasiveness of herbaceous plant species.

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.

Invasive Insects Differ from Non-Invasive in Their Thermal Requirements.

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.

Developmental rate isomorphy in insects and mites.

When the proportion of total developmental time spent in a particular developmental stage does not change with temperature, an organism shows "rate isomorphy." This is the case only if the lower developmental threshold is the same for all developmental stages. In this study, the incidence of rate isomorphy in seven species of mites and 342 species from 11 insect orders (some represented by several populations) was determined. Whether a species shows rate isomorphy or not was determined over a range of temperatures where the relationship between the rate of development and temperature is linear. Proportion of total developmental time spent in a particular stage was plotted against temperature and the existence of rate isomorphy inferred from a zero change in proportion. Rate isomorphy was detected in 243 (57%) of 426 populations. In the rest of the cases, rate isomorphy was violated by deviations in the proportion of time spent in a stage by an average of 0.2% (range 4.5E-06% to 2.8%) at the mean of the range of temperatures of all the data sets (11 degrees C). The violations occurred most frequently at the extremes of the linear phase, which is attributed to methodical biases, mortality at low temperatures, or too coarse an estimate of developmental time at high temperatures. Similarly, a meta-analysis also revealed an overall prevalence of rate isomorphy. Consequently, in insect and mite species, all the developmental stages appear to have the same population-specific lower developmental threshold. The existence of rate isomorphy could be of great practical importance, for example, in the timing of life-history events and in determining preadult thermal requirements. There are also indications that it may act as a phylogenetic constraint.

A general rule for the dependence of developmental rate on temperature in ectothermic animals.

In animals that do not regulate their body temperature by the production of heat, the proportion of the total developmental time spent in a particular developmental stage does not change with temperature. In the quasi-linear region of the relationship between developmental rate and temperature, all of the developmental stages appear to have the same species-specific lower developmental threshold. This trait, which is called developmental isomorphy, constrains developmental adaptations of ectotherms to their environments and facilitates the precise timing of life-history events.

Plant species richness of nature reserves: the interplay of area, climate and habitat in a central European landscape.

Aim To detect regional patterns of plant species richness in temperate nature reserves and determine the unbiased effects of environmental variables by mutual correlation with operating factors. Location The Czech Republic. Methods Plant species richness in 302 nature reserves was studied by using 14 explanatory variables reflecting the reserve area, altitude, climate, habitat diversity and prevailing vegetation type. Backward elimination of explanatory variables was used to analyse the data, taking into account their interactive nature, until the model contained only significant terms. Results A minimal adequate model with reserve area, mean altitude, prevailing vegetation type and habitat diversity (expressed as the number of major habitat types in the reserve) accounted for 53.9% of the variance in species number. After removing the area effect, habitat diversity explained 15.6% of variance, while prevailing vegetation type explained 29.6%. After removing the effect of both area and vegetation type, the resulting model explained 10.3% of the variance, indicating that species richness further increased with habitat diversity, and most obviously towards warm districts. After removing the effects of area, habitat diversity and climatic district, the model still explained 9.4% of the variance, and showed that species richness (i) significantly decreased with increasing mean altitude and annual precipitation, and with decreasing January temperature in the region of the mountain flora, and (ii) increased with altitudinal range in regions of temperate and thermophilous flora. Main conclusions We described, in quantitative terms, the effects of the main factors that might be considered to be determining plant species richness in temperate nature reserves, and evaluated their relative importance. The direct habitat effect on species richness was roughly equal to the direct area effect, but the total direct and indirect effects of area slightly exceeded that of habitat. It was shown that the overall effect of composite variables such as altitude or climatic district can be separated into particular climatic variables, which influence the richness of flora in a context-specific manner. The statistical explanation of richness variation at the level of families yielded similar results to that for species, indicating that the system of nature conservation provides similar degrees of protection at different taxonomic levels.

Invasiveness does not predict impact: response of native land snail communities to plant invasions in riparian habitats.

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.

Effect of intra- and interspecific competition on the performance of native and invasive species of Impatiens under varying levels of shade and moisture.

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.

Bias and error in understanding plant invasion impacts.

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.