Vojtěch Jarošík

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.

A proposed unified framework for biological invasions

There has been a dramatic growth in research on biological invasions over the past 20 years, but a mature understanding of the field has been hampered because invasion biologists concerned with different taxa and different environments have largely adopted different model frameworks for the invasion process, resulting in a confusing range of concepts, terms and definitions. In this review, we propose a unified framework for biological invasions that reconciles and integrates the key features of the most commonly used invasion frameworks into a single conceptual model that can be applied to all human-mediated invasions. The unified framework combines previous stage-based and barrier models, and provides a terminology and categorisation for populations at different points in the invasion process.

A global assessment of invasive plant impacts on resident species, communities and ecosystems: the interaction of impact measures, invading species' traits and environment

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.

Geographical and taxonomic biases in invasion ecology

Invasive alien species come from most taxonomic groups, and invasion biology is searching for robust cross-taxon generalizations and principles. An analysis of 2,670 papers dealing with 892 invasive species showed that all major groups of invaders are well studied, but that most information on the mechanisms of invasion has emerged from work on a limited number of the most harmful invaders. A strong geographical bias, with Africa and Asia understudied, inhibits a balanced understanding of invasion, because we might be lacking knowledge of specific invasion mechanisms from poorly studied, regionally specific habitats. International cooperation is required to achieve a more geographically balanced picture of biological invasions. Invasive species with the greatest impact are best studied, but more studies of species that are naturalized but not (yet) invasive are needed to improve understanding of the mechanisms acting during the naturalization phase of invasions and leading to successful invasion.

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.

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.

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.

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.

ALIEN PLANTS IN TEMPERATE WEED COMMUNITIES: PREHISTORIC AND RECENT INVADERS OCCUPY DIFFERENT HABITATS

Variables determining the number of native and alien plants on arable land in Central Europe are identified. Species richness of 698 samples of weed floras recorded in the Czech Republic in plots of a standard size of 100 m 2 in 1955-2000 was studied in relation to altitudinally based floristic region, soil type, type of cultivated crop, climatic variables, altitude, year of the record, crop cover and height, and human population density in the region. Vascular plant species were classified into native and alien, the latter divided in archaeophytes, introduced before AD 1500, and neophytes, introduced after this date. The use of minimal adequate models in the analysis of covariance allowed determination of the net effects of mutually correlated environmental variables. Models for particular species groups explained 33-48% of variation in species numbers and 27-51% in propor- tions; however, explanatory variables affected native species, archaeophytes, and neophytes differently. The number and proportion of neophytes increased in 1955-2000, whereas the number of native species and archaeophytes declined (in archaeophytes more slowly in the warm than in the moderate to cool altitudinal floristic region). In warm and dry regions and on dry soils, where most archaeophytes find optimum conditions, fewer native species are able to persist in weed communities than in colder and wetter regions. Archaeophytes respond like neophytes to some variables (climate, seasonal development of crop) and alternatively like native species to other variables (increasing agricultural intensification through time, human population density). Archaeophytes are common in old crops intro- duced with the beginning of agriculture (cereals), but are poorly represented in relatively recently introduced crops (rape, maize), where neophytes are most numerous. These patterns reflect the history of plant invasions in Central Europe. Neolithic agriculture, introduced from the Near East in the sixth millenium BC, brought archaeophytes with crops and, by creating intense and continuous propagule pressure and imposing new agricultural man- agement, facilitated their invasion. By contrast, the crops introduced during the past five centuries and their specific agrotechnical management have supported spreading of other weed species, mainly invaders from overseas.

Trends in species diversity and composition of urban vegetation over three decades

Abstract Question: What was the change in diversity of urban synantropic vegetation in a medium-sized Central European city during the period of increasing urbanization (1960s–1990s)? Location: The city of Plzeň, an industrial centre of the western part of the Czech Republic. Methods: Sampling of various types of synanthropic vegetation, conducted in the 1960s, was repeated by using the same methods in the 1990s. This yielded 959 relevés, of which 623 were made in the 1960s and 336 in the 1990s. The relevés were assigned to the following phytosociological classes: Chenopodietea, Artemisietea vulgaris, Galio-Urticetea, Agropyretea repentis and Plantaginetea majoris. Total number of vascular plant species, evenness index J, number of alien species (classified into archaeophytes and neophytes), and mean Ellenberg indicator values for light, temperature, continentality, moisture, soil reaction, and nutrients were obtained for each relevé. Results: From 1960s to 1990s, there was a significant decrease of species richness and diversity in synanthropic vegetation. The proportion of archaeophytes decreased in most vegetation types, indicating the contribution of this group of species, often confined to specific rural-like habitats, to the observed impoverishment of ruderal vegetation. The proportion of neophytes did not change between the two periods. Comparison between 1960s and 1990s indicated a decrease in light, temperature, moisture, soil reaction and nutrient indicator values in some vegetation types. In both periods, Artemisieta, Galio-Urticetea and Chenopodietea formed a distinct group harbouring more species than Agropyretea and Plantaginetea. Neophytes, i.e. recently introduced species, were most represented in the early successional annual vegetation of Chenopodietea, rather than in perennial vegetation of the other classes. Conclusions: Synanthropic vegetation of Plzeň exhibited a general trend of decrease in species diversity. Nomenclature: Ehrendorfer (1973) for species; Moravec et al. (1995) for phytosociological units.