Internal transport of alien and native plants by geese and ducks: an experimental study

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The global invasion success of Central European plants is related to distribution characteristics in their native range and species traits

Aim Determining which traits predispose a species to become invasive is a fundamental question of invasion ecology, but traits affect invasiveness in concert with other factors that need to be controlled for. Here, we explore the relative effects of biological traits of plant species and their distributional characteristics in the native range on invasion success at two stages of invasion. Location Czech Republic (for native species); and the world (for alien species). Methods The source pool of 1218 species of seed plants native to Central Europe was derived from the flora of the Czech Republic, and their occurrence in 706 alien floras all over the world was recorded, distinguishing whether they were listed as an ‘alien’ or a ‘weed’ in the latest version of Randall’s ‘Global compendium of weeds’ database. The latter type of occurrence was considered to indicate species ability to invade and cause economic impact, i.e. a more advanced stage of invasion. Using the statistical technique of regression trees, we tested whether 19 biological traits and five distributional characteristics of the species in their native range can be used to predict species success in two stages of invasion. Results The probability of a species becoming alien outside its native distribution range is determined by the size of its native range, and its tolerance of a wide range of climates acquired in the region of origin. Biological traits play only an indirect role at this stage of invasion via determining the size of the native range. However, the ability of species to become a weed is determined not only by the above characteristics of native distribution, but also directly by biological traits (life form and strategy, early flowering, tall stature, generative reproduction, number of ploidy levels and opportunistic dispersal by a number of vectors). Species phylogenetic relatedness plays only a minor role; it is more important at the lowest taxonomic levels and at the later stage of invasion. Main conclusion The global success of Central European species as ‘weeds’ is determined by their distributional characteristics in the native ranges and by biological traits, but the relative importance of these determinants depends on the stage of invasion. Species which have large native ranges and are common within these ranges should be paid increased attention upon introductions, and the above biological traits should be taken into account in screening systems applied to evaluate deliberate introductions of alien plants to new regions.     

Invasion by a non‐native ecosystem engineer alters distribution of a native predator

Aim

Shifts in diet composition, abundance or distribution of native predators can occur as a result of exotic prey introductions. We examined effects of non‐native earthworms and anthropogenic landscape disturbance on habitat selection by the American robin (Turdus migratorius), a generalist predator, at landscape and local levels. We also investigated whether robins could act as vectors of spread for earthworm cocoons (egg cases).

Location

Boreal forest of Alberta, Canada.

Methods

We conducted robin and earthworm surveys at campgrounds, well pads, roads, pipelines, seismic lines and forest interiors across northern Alberta. At a subset of paired locations that had similar habitats and anthropogenic disturbance levels, we sampled both robins and earthworms.

Results

Both groups were most likely to occur at campgrounds, well pads and roads. Furthermore, robins were more likely to occur at locations where earthworms were present in our paired local‐level surveys. This correlation between robin and earthworm distributions could be due to robins acting as a vector for earthworm spread, rather than robins’ use of earthworms as prey. However, in tests using captive robins, earthworm cocoons did not survive digestion.

Main conclusions

Robin and earthworm distributions were correlated, likely due to robins’ use of earthworms as prey. These results suggest exotic prey can strongly influence native predators at both landscape and local levels, with shifts in native predator distributions occurring as a result of spatial variability in exotic prey distributions. Although the impacts of ecosystem engineering by earthworms have been previously demonstrated, our study provides evidence that effects of earthworms can also cascade upwards via trophic interactions.     

Phylogenetic patterns differ for native and exotic plant communities across a richness gradient in Northern California

Aim Increasingly, ecologists are using evolutionary relationships to infer the mechanisms of community assembly. However, modern communities are being invaded by non‐indigenous species. Since natives have been associated with one another through evolutionary time, the forces promoting character and niche divergence should be high. On the other hand, exotics have evolved elsewhere, meaning that conserved traits may be more important in their new ranges. Thus, co‐occurrence over sufficient time‐scales for reciprocal evolution may alter how phylogenetic relationships influence assembly. Here, we examined the phylogenetic structure of native and exotic plant communities across a large‐scale gradient in species richness and asked whether local assemblages are composed of more or less closely related natives and exotics and whether phylogenetic turnover among plots and among sites across this gradient is driven by turnover in close or distant relatives differentially for natives and exotics. Location Central and northern California, USA. Methods We used data from 30 to 50 replicate plots at four sites and constructed a maximum likelihood molecular phylogeny using the genes: matK, rbcl, ITS1 and 5.8s. We compared community‐level measures of native and exotic phylogenetic diversity and among‐plot phylobetadiversity. Results There were few exotic clades, but they tended to be widespread. Exotic species were phylogenetically clustered within communities and showed low phylogenetic turnover among communities. In contrast, the more species‐rich native communities showed higher phylogenetic dispersion and turnover among sites. Main conclusions The assembly of native and exotic subcommunities appears to reflect the evolutionary histories of these species and suggests that shared traits drive exotic patterns while evolutionary differentiation drives native assembly. Current invasions appear to be causing phylogenetic homogenization at regional scales.     

Disentangling the effects of land use, shrub cover and climate on the invasion speed of native and introduced pines in grasslands

Aims To determine how changes in land use, climate and shrub cover affect the invasion dynamics of native (Pinus sylvestris L.) and introduced (Pinus nigra Arn. subsp. nigra) pines in grasslands. To analyse how these factors interact and affect seedling recruitment, a bottleneck in the lifecycle of many trees. Such information is required to manage the dynamics of these species. Location Grands Causses, calcareous plateaus (Southern France). Methods We used both published and unpublished demographic and dispersal data to assess population growth and invasion speed of invading pines. A demographic and spatially explicit model, which included density dependence and stochasticity in dispersal, demography and environment, was run for different scenarios of sheep grazing pressure (nil, extensive or intensive), shrub cover (0, 10 or 20%) and drought frequency (past‐to‐present or future). For each scenario, population growth rate, invasion speed and elasticity of invasion speed to each demographic and dispersal parameter were computed. Results Grazing was the main factor for limiting invasion speed. Shrub cover reduced tree spread under nil or extensive grazing pressure, but increased it under intensive grazing pressure. Although dry years led to nil seedling establishment rates, an increase in their frequency had surprisingly few effects on pine invasion speed. This last result remained unchanged when very dry years, inducing seedling, but also sapling mortality were introduced. In most environmental conditions, population growth rate and invasion speed were higher for the introduced than for the native pine. Elasticity analysis highlighted the importance of demographic parameters on invasion speed, notably adult and sapling survival. Main conclusion Tree invasion speed may rely at least as much on human activities, like sheep grazing, tree cutting and non‐native trees introduction, as on changes in climate factors. Therefore, human activities need to be explicitly taken into account in the prediction and management of tree dynamics.     

Phenology in a warming world: differences between native and non‐native plant species

Phenology is a harbinger of climate change, with many species advancing flowering in response to rising temperatures. However, there is tremendous variation among species in phenological response to warming, and any phenological differences between native and non‐native species may influence invasion outcomes under global warming. We simulated global warming in the field and found that non‐native species flowered earlier and were more phenologically plastic to temperature than natives, which did not accelerate flowering in response to warming. Non‐native species' flowering also became more synchronous with other community members under warming. Earlier flowering was associated with greater geographic spread of non‐native species, implicating phenology as a potential trait associated with the successful establishment of non‐native species across large geographic regions. Such phenological differences in both timing and plasticity between native and non‐natives are hypothesised to promote invasion success and population persistence, potentially benefiting non‐native over native species under climate change.     

Invasive alien predator causes rapid declines of native European ladybirds

Aim Invasive alien species (IAS) are recognized as major drivers of biodiversity loss, but few causal relationships between IAS and species declines have been documented. In this study, we compare the distribution (Belgium and Britain) and abundance (Belgium, Britain and Switzerland) of formerly common and widespread native ladybirds before and after the arrival of Harmonia axyridis, a globally rapidly expanding IAS. Location Europe Methods We used generalized linear mixed‐effects models (GLMMs) to assess the distribution trends of eight conspicuous and historically widespread and common species of ladybird within Belgium and Britain before and after the arrival of H. axyridis. The distribution data were collated largely through public participatory surveys but verified by a recognized expert. We also used GLMMs to model trends in the abundance of ladybirds using data collated through systematic surveys of deciduous trees in Belgium, Britain and Switzerland. Results Five (Belgium) and seven (Britain) of eight species studied show substantial declines attributable to the arrival of H. axyridis. Indeed, the two‐spot ladybird, Adalia bipunctata, declined by 30% (Belgium) and 44% (Britain) over 5 years after the arrival of H. axyridis. Trends in ladybird abundance revealed similar patterns of declines across three countries. Main conclusion Together, these analyses show H. axyridis to be displacing native ladybirds with high niche overlap, probably through predation and competition. This finding provides strong evidence of a causal link between the arrival of an IAS and decline in native biodiversity. Rapid biotic homogenization at the continental scale could impact on the resilience of ecosystems and severely diminish the services they deliver.     

Contrasting patterns of genome‐wide polymorphism in the native and invasive range of the marine mollusc Crepidula fornicata

Selection processes are believed to be an important evolutionary driver behind the successful establishment of nonindigenous species, for instance through adaptation for invasiveness (e.g. dispersal mechanisms and reproductive allocation). However, evidence supporting this assumption is still scarce. Genome scans have often identified loci with atypical patterns of genetic differentiation (i.e. outliers) indicative of selection processes. Using microsatellite‐ and AFLP‐based genome scans, we looked for evidence of selection following the introduction of the mollusc Crepidula fornicata. Native to the northwestern Atlantic, this gastropod has become an emblematic invader since its introduction during the 19th and 20th centuries in the northeastern Atlantic and northeastern Pacific. We examined 683 individuals from seven native and 15 introduced populations spanning the latitudinal introduction and native ranges of the species. Our results confirmed the previously documented high genetic diversity in native and introduced populations with little genetic structure between the two ranges, a pattern typical of marine invaders. Analysing 344 loci, no outliers were detected between the introduced and native populations or in the introduced range. The genomic sampling may have been insufficient to reveal selection especially if it acts on traits determined by a few genes. Eight outliers were, however, identified within the native range, underlining a genetic singularity congruent with a well‐known biogeographical break along the Florida. Our results call into question the relevance of AFLP genome scans in detecting adaptation on the timescale of biological invasions: genome scans often reveal long‐term adaptation involving numerous genes throughout the genome but seem less effective in detecting recent adaptation from pre‐existing variation on polygenic traits. This study advocates other methods to detect selection effects during biological invasions—for example on phenotypic traits, although genome scans may remain useful for elucidating introduction histories.     

Growth‐form and spatiality driving the functional difference of native and alien aquatic plants in Europe

Trait‐based approaches are widely used in community ecology and invasion biology to unravel underlying mechanisms of vegetation dynamics. Although fundamental trade‐offs between specific traits and invasibility are well described among terrestrial plants, little is known about their role and function in aquatic plant species. In this study, we examine the functional differences of aquatic alien and native plants stating that alien and native species differ in selected leaf traits. Our investigation is based on 60 taxa (21 alien and 39 native) collected from 22 freshwater units of Hungarian and Italian lowlands and highlands. Linear mixed models were used to investigate the effects of nativeness on four fundamental traits (leaf area, leaf dry matter content, specific leaf area, and leaf nitrogen content), while the influence of growth‐form, altitude, and site were employed simultaneously. We found significantly higher values of leaf areas and significantly lower values of specific leaf areas for alien species if growth‐form was included in the model as an additional predictor.We showed that the trait‐based approach of autochthony can apply to aquatic environments similar to terrestrial ones, and leaf traits have relevance in explaining aquatic plant ecology whether traits are combined with growth‐forms as a fixed factor. Our results confirm the importance of traits related to competitive ability in the process of aquatic plant invasions. Alien aquatic plants can be characterized as species producing soft leaves faster. We argue that the functional traits of alien aquatic plants are strongly growth‐form dependent. Using the trait‐based approach, we found reliable characteristics of aquatic plants related to species invasions, which might be used, for example, in conservation management.     

Living in two worlds: Evolutionary mechanisms act differently in the native and introduced ranges of an invasive plant

Identifying the factors that influence spatial genetic structure among populations can provide insights into the evolution of invasive plants. In this study, we used the common reed (Phragmites australis), a grass native in Europe and invading North America, to examine the relative importance of geographic, environmental (represented by climate here), and human effects on population genetic structure and its changes during invasion. We collected samples of P. australis from both the invaded North American and native European ranges and used molecular markers to investigate the population genetic structure within and between ranges. We used path analysis to identify the contributions of each of the three factors—geographic, environmental, and human‐related—to the formation of spatial genetic patterns. Genetic differentiation was observed between the introduced and native populations, and their genetic structure in the native and introduced ranges was different. There were strong effects of geography and environment on the genetic structure of populations in the native range, but the human‐related factors manifested through colonization of anthropogenic habitats in the introduced range counteracted the effects of environment. The between‐range genetic differences among populations were mainly explained by the heterogeneous environment between the ranges, with the coefficient 2.6 times higher for the environment than that explained by the geographic distance. Human activities were the primary contributor to the genetic structure of the introduced populations. The significant environmental divergence between ranges and the strong contribution of human activities to the genetic structure in the introduced range suggest that invasive populations of P. australis have evolved to adapt to a different climate and to human‐made habitats in North America.     

Assessing the cryptic invasion of a domestic conspecific: American mink in their native range

Control of invasions is facilitated by their early detection, but this may be difficult when invasions are cryptic due to similarity between invaders and native species. Domesticated conspecifics offer an interesting example of cryptic invasions because they have the ability to hybridize with their native counterparts, and can thus facilitate the introgression of maladaptive genes. We assessed the cryptic invasion of escaped domestic American mink (Neovison vison) within their native range. Feral mink are a known alien invader in many parts of the world, but invasion of their native range is not well understood. We genetically profiled 233 captive domestic mink from different farms in Ontario, Canada and 299 free‐ranging mink from Ontario, and used assignments tests to ascertain genetic ancestries of free‐ranging animals. We found that 18% of free‐ranging mink were either escaped domestic animals or hybrids, and a tree regression showed that these domestic genotypes were most likely to occur south of a latitude of 43.13°N, within the distribution of mink farms in Ontario. Thus, domestic mink appear not to have established populations in Ontario in locations without fur farms. We suspect that maladaptation of domestic mink and outbreeding depression of hybrid and introgressed mink have limited their spread. Mink farm density and proximity to mink farms were not important predictors of domestic genotypes but rather, certain mink farms appeared to be important sources of escaped domestic animals. Our results show that not all mink farms are equal with respect to biosecurity, and thus that the spread of domestic genotypes can be mitigated by improved biosecurity.     

Responses of a native plant species from invaded and uninvaded areas to allelopathic effects of an invader

Invaders exert new selection pressures on the resident species, for example, through competition for resources or by using novel weapons. It has been shown that novel weapons aid invasion but it is unclear whether native species co‐occurring with invaders have adapted to tolerate these novel weapons. Those resident species which are able to adapt to new selective agents can co‐occur with an invader while others face a risk of local extinction. We ran a factorial common garden experiment to study whether a native plant species, Anthriscus sylvestris, has been able to evolve a greater tolerance to the allelochemicals exerted by the invader, Lupinus polyphyllus. Lupinus polyphyllus produces allelochemicals which potentially act as a novel, strong selective agent on A. sylvestris. We grew A. sylvestris seedlings collected from uninvaded (naïve) and invaded (experienced) sites growing alone and in competition with L. polyphyllus in pots filled with soil with and without activated carbon. Because activated carbon absorbs allelochemicals, its addition should improve especially naïve A. sylvestris performance in the presence of the invader. To distinguish the allelochemicals absorption and fertilizing effects of activated carbon, we grew plants also in a mixture of soil and fertilizer. A common garden experiment indicated that the performances of naïve and experienced A. sylvestris seedlings did not differ when grown with L. polyphyllus. The addition of activated carbon, which reduces interference by allelochemicals, did not induce differences in their performances although it had a positive effect on the aboveground biomass of A. sylvestris. Together, these results suggest that naïve and experienced A. sylvestris plants tolerated equally the invader L. polyphyllus and thus the tolerance has not occurred over the course of invasion.     

Slashing Phragmites (Phragmites australis) prior to planting does not promote native vegetation establishment

Phragmites or Common Reed (Phragmites australis) is a tall rhizomatous cosmopolitan grass. While native to Australia, it can be invasive in wetlands, forming dense monocultures and reducing their ecological integrity. We assessed the potential for the cutting of Phragmites reeds prior to planting to promote the establishment of indigenous shrubs that might ultimately outcompete Phragmites. We established ten 5 m × 5 m quadrats in an area dominated by Phragmites, brush‐cut the reeds to ground level in five of them and left five uncut as controls. Within each quadrat, we planted 20 plants (~40 cm tall) of each of five indigenous shrub species, unguarded (4 plants/m²). We surveyed the plants one month after planting and annually for the following four years for survival, height and browsing damage. Browsing damage to plants was common (>50%) and unaffected by cutting. After four years, overall plant survival rates were ~25% and mean plant heights for the five shrub species ranged between 120 and 174 cm. Cutting of Phragmites had no positive effect on plant survival or height. In fact, two Melaleuca species grew taller in the uncut quadrats. Cutting of Phragmites reed beds prior to planting is unlikely to promote the establishment of woody plantings. However, planting within established Phragmites with or without prior brush‐cutting is worthy of further trialling as a potential tool for reinstating native diversity at Phragmites‐dominated sites.     

Trophic consequences of non‐native species for commercial fish in a backwater bay of the Three Gorges Reservoir,Southwest China

Trophic niche overlap in native and alien fish species can lead to competitive interactions whereby non‐native fishes outcompete indigenous individuals and eventually affect the viability of natural populations. The species Erythroculter mongolicus and Erythroculter ilishaeformis (belonging to the Culterinae), which are two commercially important fish species in the backwater bay of the Pengxi River in the Three Gorges Reservoir (TGR), were threatened by competition from the non‐native Coilia ectenes (lake anchovy). The latter is an alien species introduced into the lower reaches of the Yangtze River in China and now widespread in the TGR. The trophic consequences of non‐native lake anchovy invasion for E. mongolicus and E. ilishaeformis were assessed using stable isotope analysis (δ¹³C and δ¹⁵N) and associated metrics including the isotopic niche, measured as the standard ellipse area. The trophic niche of native E. mongolicus had little overlap (<15%) with the alien fish species and was significantly reduced in size after invasion by lake anchovy. This suggests that E. mongolicus shifted to a more specialized diet after invasion by lake anchovy. In contrast, the trophic niche overlap of native fish E. ilishaeformis with the alien fish species was larger (>50%) and the niche was obviously increased, implying that fish in this species exploited a wider dietary base to maintain their energetic requirements. Thus, marked changes for the native E. mongolicus and E. ilishaeformis were detected as the trophic consequences of invasion of non‐native lake anchovy.