Consistent performance of invasive plant species within and among islands of the Mediterranean basin

Since the success of an invasive species depends not only upon its intrinsic traits but also on particular characteristics of the recipient habitat, assessing the performance of an invader across habitats provides a more realistic analysis of risk. Such an analysis will not only provide insights into the traits related to invasiveness, but also the habitat characteristics that underpin vulnerability to invasion that, taken together, will facilitate the selection of management strategies to mitigate the invader’s effect. In the present study, we considered the Mediterranean basin islands as an excellent study region to test how the same invasive species perform in different habitats within a single island, and to scale up differences among islands with similar climate. We tested how the performance of three widespread plant invaders with clonal growth but contrasting life-history traits, a deciduous tree Ailanthus altissima, a succulent subshrub Carpobrotus spp., and an annual geophyte Oxalis pes-caprae, varied depending upon the species identity, habitat, and invaded island. The environmental parameters considered were habitat type, elevation, species diversity in the invaded plot, and several soil traits (% C, % N, C/N, pH, and relative humidity). The study documents that the performance of these three important and widespread plant invaders is dependent mainly on species identity, and less upon the invaded island’s general features. Likewise, differences in performance among habitats were only significant in the case of Ailanthus, whereas Carpobrotus and Oxalis appear to perform equally well in different environments. Ailanthus thus appears to have a broader spectrum of invasiveness, being able to invade a larger number of habitat types. On the contrary, Carpobrotus spp. have not yet invaded habitats different from those where the species have been originally introduced and where they are still commonly spread by humans. Oxalis distribution is mainly related to agricultural activities and disturbed sites, and the total area infested by this geophyte may be more reflection of the extent of suitable habitats than of invasiveness or ecological impact. Our results confirm the potential for these species to significantly alter the functioning of ecosystems in the Mediterranean islands and highlight the risk to other islands not yet invaded.     

Local and regional assessments of the impacts of plant invaders on vegetation structure and soil properties of Mediterranean islands

Aims Although biological invasions occur throughout the world, and some invaders are widespread in many habitats, few studies on the ecological impact of invaders have examined multiple sites. We tested how the impact of three widespread plant invaders changed depending on the identity of the species and the invaded island. We also tested whether relative species loss was lower in species‐rich communities than in species‐poor ones. Location We conducted floristic surveys and soil analyses in eight Mediterranean Basin islands: Crete and Lesbos (Greece), Sardinia (Italy), Corsica, Bagaud and Porquerolles (France), and Mallorca and Menorca (Spain). Methods We compared native species richness and diversity, proportion of life forms, soil percentage nitrogen, percentage organic carbon, C/N, and soil pH in nearby paired plots of 2 × 2 m: one control and one invaded by either the deciduous tree Ailanthus altissima, the succulent subshrubs Carpobrotus spp. or the annual geophyte Oxalis pes‐caprae, across eight Mediterranean Basin islands. Results On average, the presence of invaders reduced species diversity, Carpobrotus spp. exhibiting the largest impact and Oxalis the least. However, the relative impact was island‐dependent, and was positively but weakly associated with the species richness of the recipient community. Therophytes were the life form that experienced the largest decrease across islands. The effects of invasion on soil properties were very variable. Total N changed (increased) only in plots invaded by Ailanthus, significantly decreasing the C/N ratio. The presence of this tree increased soil pH, whereas the opposite was found in plots invaded by the other two species. Organic C increased in plots invaded by Ailanthus and Carpobrotus species. Main conclusions By conducting an analysis at multiple sites, we found that the three plant invaders had an impact on plant community structure not entirely concordant with changes in soil properties. The impacts depended on the identity of the species and of the invaded island, suggesting that impact of invaders is context‐specific. The impact in terms of species loss was not lower in species‐rich than in species‐poor communities.     

Widespread resistance of Mediterranean island ecosystems to the establishment of three alien species

Although some invasive plants are cosmopolitan, not all ecosystems are invaded to the same degree. Yet there is little experimental work on how ecosystem resistance to invasion at the establishment phase differs among ecosystems. We conducted two field sowing experiments in two consecutive years to examine establishment of the deciduous tree Ailanthus altissima, the succulent subshrub Carpobrotus spp. and the annual geophyte Oxalis pes‐caprae in coastal dunes, shrublands and oldfields in more than 200 sites across six Mediterranean Basin islands differing in climatic conditions and local species richness. Establishment success (i.e. percentage of plots with at least one seedling) and rates (i.e. seedling to sown seed ratio) were low, especially for Ailanthus even when accounting for differences in seed viability. Oxalis was capable of producing a new cohort of seedlings the year following planting. By contrast, all Ailanthus seedlings and half the Carpobrotus seedlings died following the first summer. Differences in establishment success and rates among ecosystems were species‐, island‐ and year‐dependent. Differences in precipitation and mean temperature were associated with differences in establishment rates across sites. Establishment rates tended to be positively correlated with cumulative precipitation and negatively with mean T^a. Unexpectedly, native species richness was not a good predictor of seedling establishment, except for higher Oxalis establishment success in species rich habitats. By conducting field sowing tests at multiple sites across a region we found that except for Oxalis, Mediterranean island ecosystems are quite resistant to invader establishment. These results suggest that differences in the degree of invasion between ecosystems and islands might be more dependent upon the influence of invasion event factors (e.g. propagule pressure) or factors acting at a later life‐history stages rather than differences in the resistance imposed by ecosystems to invader recruitment. Moreover, our results support the notion that in Mediterranean ecosystems invasions are highly idiosyncratic events and strongly dependent on water availability conditions.     

Are islands more susceptible to plant invasion than continents? A test using Oxalis pes‐caprae L. in the western Mediterranean

Aim We tested the relative vulnerability of islands to Oxalis pes‐caprae L. invasion compared to mainland regions. Oxalis pes‐caprae is a South African annual geophyte that reproduces via bulbils, and has spread in many Mediterranean and temperate regions of the world where introduced. Our study is one of the first detailed regional analyses of the occurrence and local abundance of a non‐native plant. Methods We conducted an extensive survey (2000 sampling points) to examine local and coarse‐scale patterns in both the occurrence and abundance of O. pes‐caprae on islands and in neighbouring mainland regions of Spain. Location We analysed occurrence (number of samples where present) and abundance (percentage cover) on two Balearic Islands (Menorca and Mallorca) and in two mainland administrative provinces of Spain (Murcia and València). Results Oxalis pes‐caprae occurrence was consistently higher on islands. Occurrence varied among habitats, being the highest in tree groves and the lowest in forests and shrublands. It was never found in these two habitats on the mainland. Mean O. pes‐caprae abundance was greatest in tree groves on the mainland, and in field margins and old fields on the islands. However, in general there were not significant differences in local abundances between island and mainland locations. Main conclusions These findings suggest that local processes (such as the biotic resistance of plant communities) are less important than coarse‐scale phenomena (such as environmental driving forces) in explaining differences in the invasion patterns observed between islands and adjacent mainland regions. We suggest that O. pes‐caprae has occupied a larger proportion of available habitats on islands due to: (1) its strong dependence on domestic animal and human‐mediated dispersal which are probably greater on the islands than in mainland areas, and (2) the smaller area encompassed by islands that, over a comparable period of time, enables a greater proportion of available habitats to be colonized (and hence higher occurrence) than equivalent larger mainland areas.     

Contrasting effects of invasive plants in plant–pollinator networks

The structural organization of mutualism networks, typified by interspecific positive interactions, is important to maintain community diversity. However, there is little information available about the effect of introduced species on the structure of such networks. We compared uninvaded and invaded ecological communities, to examine how two species of invasive plants with large and showy flowers (Carpobrotus affine acinaciformis and Opuntia stricta) affect the structure of Mediterranean plant–pollinator networks. To attribute differences in pollination to the direct presence of the invasive species, areas were surveyed that contained similar native plant species cover, diversity and floral composition, with or without the invaders. Both invasive plant species received significantly more pollinator visits than any native species and invaders interacted strongly with pollinators. Overall, the pollinator community richness was similar in invaded and uninvaded plots, and only a few generalist pollinators visited invasive species exclusively. Invasive plants acted as pollination super generalists. The two species studied were visited by 43% and 31% of the total insect taxa in the community, respectively, suggesting they play a central role in the plant–pollinator networks. Carpobrotus and Opuntia had contrasting effects on pollinator visitation rates to native plants: Carpobrotus facilitated the visit of pollinators to native species, whereas Opuntia competed for pollinators with native species, increasing the nestedness of the plant–pollinator network. These results indicate that the introduction of a new species to a community can have important consequences for the structure of the plant–pollinator network.     

Potential for higher invasiveness of the alien Oxalis pes-caprae on islands than on the mainland

The higher vulnerability of islands to invasions compared to mainland areas has been partially attributed to a simplification of island communities, with lower levels of natural enemies and competitors on islands conferring vacant niches for invaders to establish and proliferate. However, differences in invader life-history traits between populations have received less attention. We conducted a broad geographical analysis (i.e. 1050 km wide transect) of plant traits comparing insular and mainland populations to test the hypothesis that alien plants from insular populations have the potential for higher invasiveness than their alien mainland counterparts. For this purpose plants of the annual geophyte Oxalis pes-caprae were grown from bulbs collected in the Balearic islands and the Spanish mainland under common greenhouse conditions. There were no significant differences in bulb emergence and plant survival between descendants from insular and mainland populations. However, Oxalis descendants from insular populations produced 20% more bulbs without reducing allocation to bulb size, above-ground biomass or flowering than descendants from mainland populations. Based on the lack of sexual reproduction in Oxalis and the dependence of invasion on bulb production, our study suggests that the higher occurrence of Oxalis in the Balearic islands than in the Spanish mainland can partially be explained by genetically based higher propagation potential of insular populations compared to mainland populations.     

Pollen flow between flowers of the same morph in invasive populations of Oxalis pes-caprae L. in the western Mediterranean region

Oxalis pes-caprae is a tristylous South African geophyte that is invasive in regions with Mediterranean climate. Given the introduction of the short-styled morph only, vegetative reproduction was, until recently, the reproductive mode known for this invaded area. The detection of new floral morphs, fruit and seed production in natural populations of the invaded range, together with the reported weakening in the heteromorphic incompatibility system, raised the question on whether the short-styled flowers are able to exchange pollen among themselves, and if so, which mechanisms account for it. Thus, flower morphometric analysis and field experiments were conducted to assess pollinator visitation rates and pollen flow in three invasive populations. Flower morphological changes suggesting the evolution from tristyly towards semi-homostyly were observed. Moreover, O. pes-caprae flowers were visited by different functional groups of pollinators, mostly hymenopteran. Dye flow experiments revealed that the short-styled flowers were able to successfully exchange pollen among themselves in natural conditions. All of this, together with the weakening of the incompatibility system, constitutes further data that might help to explain the recently reported occurrence of sexual reproduction in this invaded range, which by itself bears important implications for the invasion process of O. pes-caprae.     

The impact of <Emphasis Type="Italic">Carpobrotus</Emphasis> cfr. <Emphasis Type="Italic">acinaciformis</Emphasis> (L.) L. Bolus on soil nutrients,microbial communities structure and native plant communities in Mediterranean ecosystems

Background and aims

Carpobrotus spp. are amongst the most impactful and widespread plant invaders of Mediterranean habitats. Despite the negative ecological impacts on soil and vegetation that have been documented, information is still limited about the effect by Carpobrotus on soil microbial communities. We aimed to assess the changes in the floristic, soil and microbial parameters following the invasion by Carpobrotus cfr. acinaciformis within an insular Mediterranean ecosystem.

Methods

Within three study areas a paired-site approach, comparing an invaded vs. a non-invaded plot, was established. Within each plot biodiversity indexes, C and N soil content, pH and microbial biomass and structure (bacterial and fungal) were assessed.

Results

Invaded plots showed a decrease of α-species richness and diversity. The least represented plant species in invaded plots were those related to grassland habitats. In all invaded soils, a significant increase of carbon and nitrogen content and a significant decrease of pH were registered. Carpobrotus significantly increased bacterial and fungal biomass and altered soil microbial structure, particularly favoring fungal growth.

Conclusions

Carpobrotus may deeply impact edaphic properties and microbial communities and, in turn, these strong modifications probably increase its invasive potential and its ability to overcome native species, by preventing their natural regeneration.

     

Spreading of the invasive Carpobrotus aff. acinaciformis in Mediterranean ecosystems: The advantage of performing in different light environments

Question: Do specific environmental conditions affect the performance and growth dynamics of one of the most invasive taxa (Carpobrotus aff. acinaciformis) on Mediterranean islands? Location: Four populations located on Mallorca, Spain. Methods: We monitored growth rates of main and lateral shoots of this stoloniferous plant for over two years (2002–2003), comparing two habitats (rocky coast vs. coastal dune) and two different light conditions (sun vs. shade). In one population of each habitat type, we estimated electron transport rate and the level of plant stress (maximal photochemical efficiency Fv/ Fm) by means of chlorophyll fluorescence. Results: Main shoots of Carpobrotus grew at similar rates at all sites, regardless habitat type. However, growth rate of lateral shoots was greater in shaded plants than in those exposed to sunlight. Its high phenotypic plasticity, expressed in different allocation patterns in sun and shade individuals, and its clonal growth which promotes the continuous search of available resources, contributed to a good growth and photochemical efficiency of Carpobrotus in the relatively moderate shade of the understories of Mediterranean shrublands and woodlands. Each main shoot of a Carpobrotus clone (which can have several dozens main shoots) grows ca. 40 cm per year, which explains its vigorous habitat colonization capacity. Conclusion: The highly plastic morphological response to different light regimes of this taxon contributes to a rapid colonization of heterogeneous coastal Mediterranean environments spreading well beyond the open sand dune systems where it has been often reported.     

Control of invasive species for the conservation of biodiversity in Mediterranean islands. The LIFE PonDerat project in the Pontine Archipelago,Italy

We present the EU LIFE PonDerat project, which is aimed at restoring the natural ecosystems of the Pontine Archipelago, a group of islands located off the western coast of Italy. The spread of invasive species is a major environmental threat on these islands, which are rich in rare habitats and endemic species and are important sites for the conservation of Mediterranean biodiversity. The project focuses on the conservation of species and habitats that are protected by EU laws but are currently threatened by introduced plants and animals. The main targets of the control measures are black rats, feral goats, mouflons and invasive plants of the genus Carpobrotus. Conservation measures focus on the shearwaters Puffinus yelkouan and Calonectris diomedea, which are endemic to the Mediterranean and are listed in Annex I of the European Bird Directive. Conservation measures also focus on island habitats of great biogeographical value, which are listed in Annex I of the European Habitats Directive and are seriously threatened by introduced herbivores and by invasive plants. The main outcome expected from this project is that the restoration of ecosystem functions will enhance the breeding success and population size of shearwaters and lead to the recovery of target habitats.     

Commercially Important Trees as Invasive Aliens – Towards Spatially Explicit Risk Assessment at a National Scale

Alien species that are desirable and commercially important in parts of the landscape, but damaging invaders in other parts, present a special challenge for managers, planners, and policy-makers. Objective methods are needed for identifying areas where control measures should be focussed. We analysed the distribution of forestry plantations and invasive (self-sown) stands of Acacia mearnsii and Pinus spp. in South Africa; these two taxa account for 60% of the area under commercial plantations and 54% of the area invaded by alien trees and shrubs. The distribution of commercial forestry plantations and invasive stands of these taxa were mapped and the data was digitised and stored as Geographic Information System (GIS) (Arc/Info) layers. A series of environmental parameters were derived from GIS layers of climate, topography, geology, land use, and natural vegetation. The current distribution of the two taxa was subdivided into three groups according to the degree of invasion, planting history and the precision of the data collection. We used regression-tree analysis to relate, for each taxon, the distribution of invasive stands with environmental variables, and to derive habitat suitability maps for future invasion. The current distribution of invasive stands in South Africa was largely influenced by climatic factors. At a national scale, the distribution of large commercial plantations was a poor predictor of areas invaded by both taxa. Using environmental factors identified by the regression trees, we found that 6.6% and 9.8% of natural habitats currently not invaded and untransformed by urbanisation or agriculture are suitable for invasion by Pinus spp. and A. mearnsii, respectively. We then derived guidelines for policy on alien plant management based on vegetation type, degree of transformation, extent of invasion, and the risk of future alien spread. These factors were used to identify demarcated areas where these alien species can be grown with little risk of invasions, and areas where special measures are needed to manage spread from plantations.     

Plant invasions on small Mediterranean islands: An overview

Biological invasions have become one of the main drivers of habitat degradation and a leading cause of biodiversity loss in island ecosystems worldwide. The spread of invasive species poses a particular environmental threat on the islands of the Mediterranean Basin, which are hot spots of biodiversity and contain rare habitats and endemic species, especially on small islands, which are highly vulnerable to biodiversity loss. Following a recent survey, in this paper we aim to provide an overview of the present-day non-native vascular flora of small Mediterranean islands based on a sample of 37 islands located in the middle of the Mediterranean Sea, off the coast of Italy. By comparing the current data with those gathered during a previous survey conducted in the same study area, we also aim to highlight the main changes that have occurred in non-native plant species diversity, establishment and distribution in recent years and to present a first general overview of the most prominent plant taxa in the island’s introduced flora, focusing on those most responsible for these changes and those that pose the greatest environmental threats. We recorded 203 non-native plant species, 147 of which have established on at least one of the islands investigated. Overall, we detected a sharp increase in the number of species, in their levels of establishment and in the extent of their distribution within the study area in recent years. This may be explained by the intensification of research on plant invasions, as well as to new introduction, escape, establishment and invasion events on the islands in recent decades. The most remarkable plants detected include acacias and succulents, two groups that appear to be emerging very rapidly and to be posing new threats to the conservation of the islands’ natural environment, especially the genus Carpobrotus, whose spread into natural habitats containing rare and endemic taxa is seriously threatening biodiversity on both a local and global scale. On the whole, our results show that the plant invasion phenomenon in the study area has in recent years intensified considerably. As this process seems likely to continue, we should expect more establishment events in the future and the further spread of species that are already present. This is of particular conservation concern on the islands investigated in this survey, which are rich in endemisms, but have been facing deep socio-economic and environmental transformations in these last decades as a consequence of the abandonment of traditional management practices and the development of tourism. Our study thus confirms that plant invasions on Mediterranean islands are a serious environmental problem that threatens biodiversity conservation not only in the Mediterranean biogeographic region, but also on the global scale, and highlights the need to further increase efforts aimed at preventing, controlling or mitigating the effects of plant invasions in island ecosystems.     

Impacts of alien plants and man‐made disturbance on soil‐growing bryophyte and lichen diversity in coastal areas of Sardinia (Italy)

Abstract

Seventy phytosociological relevés were performed in 1 m × 1 m plots at 14 study sites spread along sandy shores in northern and southern Sardinia (Italy). The plots were selected in different habitat types (open dunes, native Juniperus woodlands, maquis, and plantations with Acacia, Eucalyptus and Pinus) according to a stratified sampling method in order to investigate impacts deriving from different levels of Carpobrotus spp. cover, dry litter from exotic trees, and other disturbance types. The quantile regression and logistic regression analyses revealed that the reduction in the amount of bryophyte and lichen cover on sand dunes of the study area is caused either by a high cover of Carpobrotus spp. mats or by a high cover of dry exotic litter in dense, unmanaged or poorly managed forest plantations. Additional detrimental effects are often driven by other kinds of man‐made disturbances. Forest management in the coastal areas of Sardinia should be gradually modified to take into account the conservation of bryophytes and lichens. Some of the biological indicators used are quite widespread in the Mediterranean coastal habitats or are exclusively associated with sand dunes; therefore, they can also be conveniently used as indicators of biological impacts in other countries or islands of the same biogeographical region.     

Impact of invasive plants in Mediterranean habitats: disentangling the effects of characteristics of invaders and recipient communities

While the effects of an invasive alien plant that has become dominant in a community may seem obvious, there are few studies that attempt to understand how impacts vary according to the characteristics of invaders and recipient communities. For this purpose, the vegetation of invaded and non-invaded plots was sampled for eight different invasive species in a variety of habitats within the French continental Mediterranean region. Most of the observed impact variation was species-specific, with greater effects on community-level metrics found for Carpobrotus spp. and Reynoutria × bohemica and lower effects for Amorpha fruticosa, Ambrosia artemisiifolia and Phyla filiformis. Some trends were consistent with competition-driven processes, with higher impact found in the presence of rhizomatous and creeping perennial invasive species compared to annuals, or in habitats with sparse vegetation. The importance of community characteristics such as the cover of the invasive plant or the differences in cover between the invader and the native dominant species confirmed previous results obtained in Central Europe. Therefore, such variables, easy to measure and with a generic value, could be profitably integrated into risk assessment methods to improve the prediction of the most threatened habitats. Beyond the overall decline in species diversity, the presence of some invasive species was associated with significant changes in species composition, with a filtering toward more shade-tolerant and nitrophilous ruderal species. Managers should consider replacement of resident species by species with different ecological preferences together with simple community-level metrics, to decide whether management is justified.     

Anthropogenic subsidies mitigate environmental variability for insular rodents

The exogenous input of nutrients and energy into island systems fuels a large array of consumers and drives bottom-up trophic cascades in island communities. The input of anthropogenic resources has increased on islands and particularly supplemented non-native consumers with extra resources. We test the hypothesis that the anthropogenic establishments of super-abundant gulls and invasive iceplants Carpobrotus spp. have both altered the dynamics of an introduced black rat Rattus rattus population. On Bagaud Island, two habitats have been substantially modified by the anthropogenic subsidies of gulls and iceplants, in contrast to the native Mediterranean scrubland with no anthropogenic inputs. Rats were trapped in all three habitats over two contrasting years of rainfall patterns to investigate: (1) the effect of anthropogenic subsidies on rat density, age-ratio and growth rates, and (2) the role of rainfall variability in modulating the effects of subsidies between years. We found that the growth rates of rats dwelling in the non-subsidized habitat varied with environmental fluctuation, whereas rats dwelling in the gull colony maintained high growth rates during both dry and rainy years. The presence of anthropogenic subsidies apparently mitigated environmental stress. Age ratio and rat density varied significantly and predictably among years, seasons, and habitats. While rat densities always peaked higher in the gull colony, especially after rat breeding in spring, higher captures of immature rats were recorded during the second year in all habitats, associated with higher rainfall. The potential for non-native rats to benefit from anthropogenic resources has important implications for the management of similar species on islands.     

Low individual-level dietary plasticity in an island-invasive generalist forager

The ability of invasive mammals to adjust their diet in response to new or variable resources is often proposed to explain their invasion success on islands with differing environmental conditions, especially islands with strong spatiotemporal changes in the nature and abundance of their resources. In this study, we investigated how habitat heterogeneity and seasonal fluctuation in resource quality affect dietary breadth and plasticity in an island-invasive rodent, the black rat Rattus rattus, on a small Mediterranean island. We tested for dietary plasticity of rats at both the individual and population levels by using traditional dietary and stable isotope analyses at successively increasing time scales, coupled with a long-term study of individual rats in three habitats of close proximity. Dietary and movement analyses both indicated that R. rattus is able to exploit a wide range of resources and habitats. However, dietary plasticity and habitat breadth were far narrower at the individual level. Results revealed that rats exclusively used resources found in their local habitat, and very few individuals moved among adjacent habitats in pursuit of higher-quality resources, despite those resources being abundant in their immediate environment. This counterintuitive finding suggests that intraspecific interactions must restrict rat mobility. Our results suggest that even on small islands, accessibility of patchy and high-quality resources to individuals from the entire population is not systematic. This result has important implications when quantifying invasive rodent impacts on patchily distributed species, especially when studies use indirect methods such as dietary analyses as a substitute for direct observations of predatory behavior.     

Can herbarium records be used to map alien species invasion and native species expansion over the past 100 years?

Aim To determine if the temporal and spatial pattern of alien plant invasion and native plant expansion can be observed using 100 years of herbarium data from Oklahoma, USA, and to eliminate herbarium collection biases in such analyses. Location Oklahoma, USA. Methods Using herbarium records from the Oklahoma Vascular Plants Database from 1903 to 2004, we reconstructed the spatial and temporal collection history of two alien invasive taxa (Lonicera japonica and Tamarix spp.) and three native expansive species (Ambrosia psilostachya, Amphiachyris dracunculoides and Juniperus virginiana). To compare the overall collecting trend, groups of native non‐expansive taxa were selected as counterparts. We recorded the year of the first collection in each township in Oklahoma for all taxa. The cumulative number of occupied townships was log‐transformed, plotted against time and modelled with linear regression. The slope of the linear regression represented collection trend over time for the non‐expansive counterpart group. However, for the invasive and expansive species, the regression slope represented the collection effort plus the invasion or expansion rate. We calculated the proportion of invasive and expansive species to non‐expansive species by dividing the cumulative number of townships for each invasive or expansive species by the cumulative number of townships occupied by the counterpart group (proportion curve). Results Maps of the collection records of invasive and expansive taxa illustrated no discernible spatial invasion or expansion pattern. The slopes of the linear regression for alien invasive taxa were significantly steeper than those of their associated native non‐expansive counterparts, indicating an increase in abundance. Juniperus virginiana, L. japonica and Tamarix spp. exhibited one or more periods during which they were collected at a disproportionately higher rate than their native non‐expansive counterparts. Main conclusions Patterns of species invasion and expansion in Oklahoma were detected using techniques developed for regions with longer collecting plant histories. The proportion curve analysis eliminated some biases inherent in herbarium data by reducing the effect of collecting effort. Both the regression model and proportion curve analyses illustrate the temporal invasion patterns of alien invasive species. The native species did not show a clear expansion pattern. The information found in recently established herbaria may not be sensitive enough to detect the increase in abundance of native species.     

Predation contributes to invasion resistance of benthic communities against the non-indigenous tunicate <Emphasis Type="Italic">Ciona intestinalis</Emphasis>

Marine anthropogenic structures offer novel niches for introduced species but their role in the subsequent invasion to natural habitats remains unknown. Upon arrival in new environments, invaders must overcome biotic resistance from native competitors and predators if they are to establish successfully in natural habitats. We tested the hypotheses that (1) artificial structures (e.g., suspended aquaculture installations) present a niche opportunity for invasive species by providing a refuge from native benthic predators, and (2) native predators in natural benthic habitats suppress successful colonization by invaders. A recruitment experiment showed that the ascidians Pyura chilensis (native) and Ciona intestinalis (invasive) could recruit to both suspended artificial structures and natural benthic habitats. Ciona, however, was only able to establish adult populations on artificial structures. In natural benthic habitats Ciona only recruited and grew in predator-exclusion cages, because without this protection predation prevented its establishment. In predation experiments, native invertebrate and fish predators removed all invasive ascidians (recruits and adults) in benthic habitats, which contrasted with the high adult survival of the native ascidian P. chilensis. The refuge from a number of benthic predators facilitates the establishment of large populations of invasive species on suspended structures. We present a conceptual model of the invasion processes that includes the anthropogenic structures as a transitional stepping-stone that facilitates invasion by enhancing and prolonging propagule supply to surrounding natural communities. Those established invaders might then overcome biotic resistance during time periods when populations of consumers or competitors are weakened by natural or anthropogenic disturbances. Our results suggest that the conservation of natural habitats with a high diversity of native predators can be an effective means to prevent the spread of invasive species growing on suspended structures.     

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.     

Deposition of exotic bird-dispersed seeds into three habitats of a fragmented landscape in the northeastern United States

Dispersal by frugivorous birds facilitates invasion by many exotic plants. We measured the seed rain of ornithochorous native and exotic plants at three habitats of a fragmented landscape of the northeastern United States for 1 year. We studied maple-beech forests, old fields, and abandoned conifer plantations. Across all sites we collected 2,196 ornithochorous seeds, including seeds from six exotic species and 10 native species. The majority (90%) of collected seeds were from exotic species. Seed dispersal was broadly similar among habitats, though seed rain of exotic species was higher in old fields than forested habitats. Seed rain was not strongly influenced by artificial perches for most species. However, seeds of exotic species were more commonly found in traps under an artificial perch in old fields. Seed rains for the exotic Elaeagnus umbellata, Rhamnus cathartica, and Rosa multiflora were positively associated with local density of mature plants. Seed rain of R. cathartica was positively associated with abundance of seedlings but not saplings, suggesting that post-dispersal mortality was important. Seed dispersal of the exotic Lonicera spp. was high in all habitats, accounting for 66% of all seeds collected. With the exception of Lonicera spp., seed rain of common exotic invaders was affected by the abundance of seed sources, and these species might be effectively controlled by elimination of local fruiting plants. Fruits of Lonicera morrowii, which has extensively invaded our area, are apparently a common component in the diet of frugivores.