Predicting and explaining plant invasions through analysis of source area floras: some critical considerations

Comparing species that become invasive with others from the same regional species pool that do not invade raises several issues about the accuracy of analyses attempting to define the determinants of invasiveness. The delimitation of the source area and deciding which species group(s) to include are especially relevant in analyses focusing on species originating in Europe. Historical patterns of immigration of alien species into Europe must be considered since European floras comprise a complex mix of native species, historical introductions (archaeophytes) and relative newcomers (neophytes). We make three main points: (1) Archaeophytes (species introduced to Europe before the discovery of America) differ from natives in a number of traits and in historical association with people; it is misleading to lump archaeophytes with native taxa. (2) Taxa from climatically and geographically different regions, representing distinct floristic geoelements, need to be treated separately, and not as a homogenous pool of potential invaders. Restricting the source species pool to native taxa with comparable phytogeographical characteristics reduces the variation associated with chance of dispersal by humans from the source area. (3) For prediction, a clear distinction should be made between accuracy (the proportion of those found to be alien that were also predicted to be there) and reliability (or predictive value, the proportion of those predicted to become aliens that do so). Information accumulated over centuries by botanists in Central Europe provides an excellent opportunity to deal with these issues and avoid spurious results. To illustrate these issues, we revisit a recently published study of Central‐European plant species as invaders in two Argentinean provinces ( Prinzing et al., 2002 ) to explore and demonstrate the implications of the above points. We hope that future studies will build on these points to achieve more reliable predictions.     

Habitat Overlap and Facilitation in Tamarisk and Box Elder Stands: Implications for Tamarisk Control Using Native Plants

Invasive plants are typically managed using top‐down control techniques that focus on the removal of the target organism. Bottom‐up control limits the resources available to the undesired species by manipulating disturbance, competition, and successional processes, and thus may prevent reinvasion. Tamarisk species (Tamarix sp.) have invaded riparian areas throughout western North America, resulting in expansive control efforts. A companion study has shown that a native competitor, Box elder (Acer negundo), is capable of outcompeting and killing established Tamarisk through light interception in canyons of Dinosaur National Monument (DNM), Colorado. The goal of this study was to determine the feasibility of using Box elder as a bottom‐up control agent by (1) determining the distributional overlap of the two species in DNM; (2) determining if Tamarisk facilitates Box elder establishment; and (3) analyzing Box elder seedling survival across a range of physical gradients. The distribution of Tamarisk and Box elder overlapped considerably throughout the study area. Box elder seedlings were planted under Tamarisk canopies or areas with the canopy removed. Survival was significantly higher under Tamarisk canopies, indicating that Tamarisk facilitates Box elder seedling survival. Box elder seedling survival was tested across soil texture, litter depth, groundwater depth, and shade intensities indicative of conditions found in the canyons of DNM, and survival was high for all treatments. The manipulation of competitive and successional processes through the promotion of Box elder and other native tree establishment is suggested as a means of bottom‐up Tamarisk control to complement traditional control techniques.     

Growth,phenology, and biomass allocation of alien Solidago species in central Europe

Plant invasion is a major threat to the integrity of an ecosystem. Exceptionally successful invaders in Europe are the American species of Solidago genus. In this study we examined growth, reproduction, and phenology of Solidago species, of American origin, growing in central Europe (S. altissima, S. canadensis, S. gigantea, S. graminifolia). These taxa were compared with two native species: Solidago virgaurea and Tanacetum vulgare. We observed high differentiation in height, number of shoots, and biomass production between individuals within taxa. Generally, the invasive species produced substantially (two to five times) more biomass than the native ones, being statistically significant in the comparison of alien Solidago graminifolia and S. gigantea versus native S. virgaurea and T. vulgare. The ratio of biomass of reproductive parts to overall biomass varied considerably among years, but generally the lowest one was for Solidago altissima, and the highest for S. graminifolia. It shows a lack of a clear pattern of differentiation between alien and native species in terms of biomass investment in reproduction. We observed a general tendency of allocation of a major part of biomass in rhizomes by phalanx species (S. graminifolia and S. gigantea), while species with guerrilla strategy (S. altissima and S. canadensis) invested more biomass in stems and leaves. However, because of the high variability there was no clear, stable pattern of statistically significant differences between these two groups. The results suggest that S. graminifolia reveals a strong potential of invasion, in spite of its, so far limited, distribution in Europe.     

Native and alien plant species richness in relation to spatial heterogeneity on a regional scale in Germany

Aim The aim of our study was to reveal relationships between richness patterns of native vs. alien plant species and spatial heterogeneity across varying landscape patterns at a regional scale. Location The study was carried out in the administrative district of Dessau (Germany), covering around 4000 km². Methods Data on plant distribution of the German vascular flora available in grid cells covering 5′ longitude and 3′ latitude (c. 32 km²) were divided into three status groups: native plants, archaeophytes (pre 1500 AD aliens) and neophytes (post 1500 AD aliens). Land use and abiotic data layers were intersected with 125 grid cells comprising the selected area. Using novel landscape ecological methods, we calculated 38 indices of landscape composition and configuration for each grid cell. Principal components analysis (PCA) with a set of 29 selected, low correlated landscape indices was followed by multiple linear regression analysis. Results PCA reduced 29 indices to eight principal components (PCs) that explained 80% cumulative variance. Multiple linear regression analysis was highly significant and explained 41% to 60% variance in plant species distribution (adjusted R²) with three significant PCs (tested for spatial autocorrelation) expressing moderate to high disturbance levels and high spatial heterogeneity. Comparing the significance of the PCs for the species groups, native plant species richness is most strongly associated with riverine ecosystems, followed by urban ecosystems, and then small‐scale rural ecosystems. Archaeophyte and neophyte richness are most strongly associated with urban ecosystems, followed by small‐scale rural ecosystems and riverine ecosystems for archaeophytes, and riverine ecosystems and small‐scale rural ecosystems for neophytes. Main conclusions Our overall results suggest that species richness of native and alien plants increases with moderate levels of natural and/or anthropogenic disturbances, coupled with high levels of habitat and structural heterogeneity in urban, riverine, and small‐scale rural ecosystems. Despite differences in the order of relevance of PCs for the three plant groups, we conclude that at the regional scale species richness patterns of native plants as well as alien plants are promoted by similar factors.     

Inverted invasions: Native plants can frequently colonise urban and highly disturbed habitats

There is an enormous body of literature on plant invasions, including many investigations of the types of introduced species that are most likely to invade natural ecosystems. In this study we turn invasion biology upside down, and ask what sort of native species colonise novel anthropogenic habitats such as roadside lawns, infrequently tended road shoulders, railway embankments and fire trails. We quantified species richness and cover in roadside lawns and infrequently tended road shoulders in five regions of New South Wales, Australia. The native vegetation in these regions included sclerophyll forest, fertile and infertile Eucalypt‐dominated woodlands, rainforest, and semi‐arid woodland. We performed a complementary survey of sites spanning five disturbance levels within the region containing sclerophyll forest vegetation. Although many non‐native species were present in disturbed, novel habitats, a total of 136 native species were also found. Most of these native species were in sites with low levels of disturbance (fire trails and railway embankments), but 35 native species were found to colonise roadside lawns, our most highly‐disturbed vegetation type. There was a significant negative relationship between the disturbance level in novel habitats and the number and cover of native species. Native species that colonised novel habitats were disproportionately likely be generalist species whose natural habitat includes both high and low light and high and low disturbance conditions. The native species colonising novel habitats also tended to have traits associated with a fast life‐history, including short stature and small seeds. A surprisingly high number of native plant species are colonising novel, anthropogenic habitats. Our findings highlight the potential importance of urban ecosystems for conservation and restoration biology.     

Contrasting patterns in the invasions of European terrestrial and freshwater habitats by alien plants,insects and vertebrates

Aim To provide the first comparative overview on the current numbers of alien species that invade representative European terrestrial and freshwater habitats for a range of taxonomic groups. Location Europe. Methods Numbers of naturalized alien species of plants, insects, herptiles, birds and mammals occurring in 10 habitats defined according to the European Nature Information System (EUNIS) were obtained from 115 regional data sets. Only species introduced after ad 1500 were considered. Data were analysed by ANCOVA and regression trees to assess whether differences exist among taxonomic groups in terms of their habitat affinity, and whether the pattern of occurrence of alien species in European habitats interacts with macroecological factors such as insularity, latitude or area. Results The highest numbers of alien plant and insect species were found in human‐made, urban or cultivated habitats; if controlled for habitat area in the region, wetland and riparian habitats appeared to support relatively high numbers of alien plant species too. Invasions by vertebrates were more evenly distributed among habitats, with aquatic and riparian, woodland and cultivated land most invaded. Mires, bogs and fens, grassland, heathland and scrub were generally less invaded. Habitat and taxonomic group explained most variation in the proportions of alien species occurring in individual habitats related to the total number of alien species in a region, and the basic pattern determined by these factors was fine‐tuned by geographical variables, namely by the mainland–island contrast and latitude, and differed among taxonomic groups. Main conclusions There are two ecologically distinct groups of alien species (plants and insects versus vertebrates) with strikingly different habitat affinities. Invasions by these two contrasting groups are complementary in terms of habitat use, which makes an overall assessment of habitat invasions in Europe possible. Since numbers of naturalized species in habitats are correlated among taxa within these two groups, the data collected for one group of vertebrates, for example, could be used to estimate the habitat‐specific numbers of alien species for other vertebrate groups with reasonable precision, and the same holds true for insects and plants.     

The role of non‐native plants and vertebrates in defining patterns of compositional dissimilarity within and across continents

Aim Human activities have led to the spread and establishment of increasing numbers of non‐native species. Here we assess whether non‐native plant and vertebrate species have affected species compositions within and across Europe and North America. We also assess the effects of intra‐continental species exchange using the example of vertebrates. Location European countries and North America (states in the contiguous United States and provinces of Canada). Methods We measured compositional dissimilarity of native and non‐native assemblages of vascular plants and vertebrates and related these patterns to climatic dissimilarity and geographical distance. We considered three categories of non‐native species (introduced after ad 1500), namely: those (1) originating outside of both continents, (2) native to one continent and non‐native to the other, and (3) native in a particular region of a continent but non‐native in another region. Results The presence of non‐native plants and vertebrates led to more homogeneous species compositions between continents and to less homogeneous species composition within Europe compared with the native assemblages. In North America, the presence of non‐native plants led to more homogeneous species compositions and the presence of non‐native vertebrates had no effect. Species compositions being more homogeneous than the native composition were found for the three categories of non‐native vertebrate species for both continents. Between continents, climate was a better predictor of compositional dissimilarity for non‐native plants, whereas for vertebrates the explanatory power of climate and geographical distance were comparable. By contrast, within continents, climate was a better predictor of compositional dissimilarity of both plants and vertebrates. Conclusions We found clear evidence for biotic homogenization as a consequence of species displacement. However, in relation to overall species richness this effect was rather small, indicating that floras and faunas are still quite distinct. Therefore, claiming that we already face homogeneous biotas might be premature, although clear indications are visible which should raise a note of caution, especially in the light of increasing globalization.     

Patterns of Plant Invasions: A Case Example in Native Species Hotspots and Rare Habitats

Land managers require landscape-scale information on where exotic plant species have successfully established, to better guide research, control, and restoration efforts. We evaluated the vulnerability of various habitats to invasion by exotic plant species in a 100,000 ha area in the southeast corner of Grand Staircase-Escalante National Monument, Utah. For the 97 0.1-ha plots in 11 vegetation types, exotic species richness (log₁₀) was strongly negatively correlated to the cover of cryptobiotic soil crusts (r = −0.47, P < 0.001), and positively correlated to native species richness (r = 0.22, P < 0.03), native species cover (r = 0.23, P < 0.05), and total nitrogen in the soil (r = 0.40, P < 0.001). Exotic species cover was strongly positively correlated to exotic species richness (r = 0.68, P < 0.001). Only 6 of 97 plots did not contain at least one exotic species. Exotic species richness was particularly high in locally rare, mesic vegetation types and nitrogen rich soils. Dry, upland plots (n = 51) had less than half of the exotic species richness and cover compared to plots (n = 45) in washes and lowland depressions that collect water intermittently. Plots dominated by trees had significantly greater native and exotic species richness compared to plots dominated by shrubs. For the 97 plots combined, 33% of the variance in exotic species richness could be explained by a positive relationship with total plant cover, and negative relationships with the cover of cryptobiotic crusts and bare ground. There are several reasons for concern: (1) Exotic plant species are invading hot spots of native plant diversity and rare/unique habitats. (2) The foliar cover of exotic species was greatest in habitats that had been invaded by several exotic species.(3) Continued disturbance of fragile cryptobiotic crusts by livestock, people, and vehicles may facilitate the further invasion of exotic plant species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Plant biodiversity patterns along a climatic gradient and across protected areas in West Africa

Knowledge of spatial patterns of biological diversity is fundamental for ecological and biogeographical analyses and for priority setting in nature conservation, particularly in West Africa where the existing high biodiversity is increasingly threatened by human activities. The maximum entropy approach was used to model the geographic distribution of 3,393 vascular plant species at a spatial resolution of 0.0833°. Species richness decreases along temperature and precipitation gradients with high species numbers in the south and lower numbers towards the north of the transect. All centres of plant species diversity are confined to humid areas in concordance with the high positive correlation between species richness and rainfall which appears to be the most important delimiter for the distribution ranges of many species in the area. The effectiveness of the existing protected areas at regional and national levels is investigated based on the proportion of species covered. Considering the whole study area, 95% of all species are covered by protected areas according to their distribution ranges. However, the proportion of species covered is considerably lower for some countries such as Benin and Togo. Our results could provide guidance for essential land use management interventions to decision‐makers and conservationists in the region.     

中国农业生态系统外来种入侵及其管理现状

农业生态系统极易遭受外来生物入侵。作者根据文献资料和多年工作观察统计出入侵我国农业生态系统的外来生物共计92科175属239种, 其中植物155种, 动物55种, 微生物29种, 植物多为有意引入后逸生, 而动物和微生物则主要是无意引入。外来入侵种发生数量呈现从南到北、从东到西逐渐减少的趋势。这些入侵种中, 来源于美洲的最多(占45.04%), 其次是欧洲(22.90%); 菜地(包括温室大棚)和果园入侵种最多, 分别达64.85%和66.53%, 而半年期的秋熟旱地和夏熟旱地分别占34.31%和23.85%。其中17种外来杂草、10种害虫、7种病原菌为恶性有害生物, 应作为防除的重点目标。目前农业生态系统外来入侵物种的控制以化学防治为主, 但由于长期施用化学农药, 在侵入我国农田的入侵种中, 已有51种在世界不同地区演化出抗药性生物型, 因而需重视生物防治、农业和生态防治以及检疫等的综合应用。今后外来种对农业生态系统的入侵格局、机制和趋势, 入侵途径以及生物入侵和抗药性生物型对农业生态系统中有害生物群落演替的影响、转基因作物导致的生物入侵等问题值得关注。     

Geographical patterns and determinants of species richness in Mexico across selected families of vascular plants: implications for conservation

Mexico is considered a megadiverse country containing more than 10% of the world's biodiversity. The distribution of this species richness and endemism is different among the different Mexican states. We examined the species richness patterns of 13 families of vascular plants (including ferns, gymnosperms and angiosperms) in Mexico using political divisions (states) as units of analysis. We analysed the species richness values (absolute richness, endemic richness and restrictive richness) of these plant families using stepwise multiple regression analysis, assessing their relationship with a set of 10 environmental variables (expressed as heterogeneity coefficients). A combined cluster analysis with multidimensional scaling analysis (MDS) and an analysis of similarities were also undertaken to define the spatial–geographical patterns. Additionally, we proposed a methodological strategy to determine which states of Mexico have priorities for conservation. Our results suggested that the three species richness values used were significantly predicted by environmental factors, especially by climatic heterogeneity. Notwithstanding that a linear pattern was recognized, the Mexican states were gathered in four groups, which were confirmed by the MDS and the cluster analysis: (1) the Yucatan Peninsula, (2) arid Mexico, (3) the Mexican Transition Zone and (4) the megadiverse states. We proposed that 12 Mexican states include all the environmental conditions and are candidates for developing conservation programmes: (1) Quintana Roo, Tabasco and Yucatán, (2) Baja California, Chihuahua and Sinaloa, (3) Guerrero, Jalisco and Nuevo León and (4) Chiapas, Oaxaca and Veracruz.     

Paddlefish (Polyodon spathula) in Europe: An aquaculture species and a potential invader

The paddlefish (Polyodon spathula) was first introduced to Europe in 1974, mainly due to its potential for rearing in natural polyculture ponds and large temperate reservoirs. The information on the history of paddlefish aquaculture efforts in Europe is scarce, as well as data on current paddlefish aquaculture status and trends. In addition, there is a lack of data on its presence and potential establishment in the wild, while its invasive potential and associated risks and impacts are largely unknown. In order to evaluate its current status in Europe, we conducted a survey among scientists, aquaculture producers and other stakeholders, and reviewed literature and data on the Internet. Based on the results obtained, we discuss the potential and the challenges in European paddlefish aquaculture development, and analyze paddlefish invasive potential and risks associated with its naturalization. Paddlefish aquaculture is well established only regionally in Europe, but offers relatively high potential for further development in pond farms. Nevertheless, future development will require careful planning, especially regarding market development and improved marketing strategies. While paddlefish likely represents a low‐risk invader, improved control and reporting on trade and intentional and unintentional releases will be required. Given the lack of knowledge on potential impacts following its introduction, due caution seems highly advisable.     

Phylogenetic beta diversity of native and alien species in European urban floras

Aim Human activities have weakened biogeographical barriers to dispersal, increasing the rate of introduction of alien plants. However, their impact on beta diversity and floristic homogenization is poorly understood. Our goal is to compare the phylogenetic beta diversity of native species with that of two groups of alien species, archaeophytes and neophytes (introduced before and after ad 1500, respectively), across European urban floras to explore how biological invasions affect phylogenetic turnover at a continental scale. Location Twenty European cities located in six countries between 49 and 53° N latitude in continental Europe and the British Isles. Methods To compare the phylogenetic beta diversity of native and alien species we use the average phylogenetic dissimilarity of individual floras from their group centroid in multivariate space. Differences in phylogenetic beta diversity among different species groups are then assessed using a randomization test for homogeneity of multivariate dispersions. Results Across European urban floras, and when contrasted with natives, archaeophytes are usually associated with lower levels of phylogenetic beta diversity while neophytes tend to increase phylogenetic differentiation. Main conclusions While archaeophytes tend to promote limited homogenization in phylogenetic beta diversity, because of their diverse geographical origin together with short residence times in the invaded regions, neophytes are not promoting biotic homogenization of urban floras across Europe. Therefore, in spite of the increasing rate of alien invasion, an intense phylogenetic homogenization of urban cities is not to be expected soon.     

Impacts of invasive plants on resident animals across ecosystems,taxa, and feeding types: a global assessment

As drivers of global change, biological invasions have fundamental ecological consequences. However, it remains unclear how invasive plant effects on resident animals vary across ecosystems, animal classes, and functional groups. We performed a comprehensive meta‐analysis covering 198 field and laboratory studies reporting a total of 3624 observations of invasive plant effects on animals. Invasive plants had reducing (56%) or neutral (44%) effects on animal abundance, diversity, fitness, and ecosystem function across different ecosystems, animal classes, and feeding types while we could not find any increasing effect. Most importantly, we found that invasive plants reduced overall animal abundance, diversity and fitness. However, this significant overall effect was contingent on ecosystems, taxa, and feeding types of animals. Decreasing effects of invasive plants were most evident in riparian ecosystems, possibly because frequent disturbance facilitates more intense plant invasions compared to other ecosystem types. In accordance with their immediate reliance on plants for food, invasive plant effects were strongest on herbivores. Regarding taxonomic groups, birds and insects were most strongly affected. In insects, this may be explained by their high frequency of herbivory, while birds demonstrate that invasive plant effects can also cascade up to secondary consumers. Since data on impacts of invasive plants are rather limited for many animal groups in most ecosystems, we argue for overcoming gaps in knowledge and for a more differentiated discussion on effects of invasive plant on native fauna.     

The distribution of range sizes of native and alien plants in four European countries and the effects of residence time

Aim Do the statistical distributions of range sizes of native and alien species differ? If so, is this because of residence time effects? And can such effects indicate an average time to a maximum? Location Ireland, Britain, Germany and the Czech Republic. Methods The data are presence or absence of higher plants in mapping units of 100 km² (Ireland and Britain) or c. 130 km² (Germany and the Czech Republic) in areas varying from 79 to 357 thousand km². Logit transforms of range sizes so defined were tested for normality, and examined by ANOVA, and by loess, ordinary least square (OLS) and reduced major axis regressions. Results Current range sizes, in logits, are near normally distributed. Those of native plants are larger than those of naturalized neophytes (plants introduced since 1500 ad ) and much larger than those of casual neophytes. Archaeophytes (introduced earlier) have range sizes slightly larger than natives, except in Ireland. Residence time, the time since an invasive species arrived in the wild at a certain place, affects range sizes. The relationships of the range of naturalized neophytes to residence time are effectively straight in all four places, showing no significant curvature or asymptote back to 1500, though there are few records between 1500 and 1800. The relationships have an r² of only about 10%. Both OLS regressions and reduced major axes can be used to estimate the time it takes for the range of a naturalized neophyte to reach a maximum. Main conclusions Established neophytes have smaller range size distributions than natives probably because many have not yet reached their maximum. We estimate it takes at least 150 years, possibly twice that, on average, for the maximum to be reached in areas of the order of 10⁵ km². Policy needs to allow for the variation in rates of spread and particularly the long time needed to fill ranges. Most naturalized neophytes are still expanding their ranges in Europe.     

Dark diversity reveals importance of biotic resources and competition for plant diversity across habitats

Species richness is the most commonly used metric to quantify biodiversity. However, examining dark diversity, the group of missing species which can potentially inhabit a site, can provide a more thorough understanding of the processes influencing observed biodiversity and help evaluate the restoration potential of local habitats. So far, dark diversity has mainly been studied for specific habitats or large‐scale landscapes, while less attention has been given to variation across broad environmental gradients or as a result of local conditions and biotic interactions. In this study, we investigate the importance of local environmental conditions in determining dark diversity and observed richness in plant communities across broad environmental gradients. Using the ecospace concept, we investigate how these biodiversity measures relate to abiotic gradients (defined as position), availability of biotic resources (defined as expansion), spatiotemporal extent of habitats (defined as continuity), and species interactions through competition. Position variables were important for both observed diversity and dark diversity, some with quadratic relationships, for example, plant richness showing a unimodal response to soil fertility corresponding to the intermediate productivity hypothesis. Interspecific competition represented by community mean Grime C had a negative effect on plant species richness. Besides position‐related variables, organic carbon was the most important variable for dark diversity, indicating that in late‐succession habitats such as forests and shrubs, dark diversity is generally low. The importance of highly competitive species indicates that intermediate disturbance, such as grazing, may facilitate higher species richness and lower dark diversity.