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Positive interactions such as mutualism and facilitation between invasive species can magnify their impact on invaded ecosystems. Yet, few studies have explicitly evaluated symbioses between invasive species that span the terrestrial–aquatic interface. We conducted two experiments to investigate the fitness outcomes (growth and germination) of frugivory and ichthyochory involving invasive channel catfish (Ictalurus punctatus) that consume fruits and pass seeds of invasive Russian olive (Elaeagnus angustifolia). Our results indicate that catfish frugivory of olive fruit provides an energy source that allows the catfish to maintain body mass and seeds passed by catfish were four times more likely to germinate than untreated seeds, suggesting catfish are viable olive seed dispersers. As such, we concluded that the interaction of frugivory of invasive olive fruits by invasive catfish and ichthyochory resulting from passed olive seeds constitute a facultative mutualism spanning the aquatic–terrestrial interface. The impact of the olive–catfish mutualism on the abundance, persistence and spread of both invasive species requires further study. Researchers and managers should consider the potential of positive interactions across ecotones to facilitate invasions [ABSTRACT FROM AUTHOR]

Non‐native ants can cause ecosystem‐wide ecological change, and these changes are generally assumed to be negative. Despite this, the evidence base has never been holistically synthesised to quantify whether and to what degree non‐native ants impact native species diversity.In this study, we performed a meta‐analysis of the effects of ant invasion on animal communities. We extracted data from 46 published articles investigating abundance (156 effect sizes) and richness (53 effect sizes) responses of animal taxa to ant invasion in locations relatively unimpacted by other stressors (e.g. human disturbance, other non‐native species) to help isolate the effects of invasion.Overall, local animal diversity declined severely, with species abundance and richness lower by 42.79% and 53.56%, respectively, in areas with non‐native ants compared with intact uninvaded sites. We then combined responses of individual animal taxa extracted from an article into a single response to represent the 'community' abundance (40 effect sizes) or richness (28 effect sizes) response to non‐native ants represented in each article. Local communities decreased substantially in total abundance (52.67%) and species richness (53.47%) in invaded sites.These results highlight non‐native ants as the drivers, rather than passengers, of large net‐negative reductions to animal community diversity in relatively undisturbed systems around the world, approximately halving local species abundance and richness in invaded areas. Improved international prevention processes, early detection systems harnessing emerging technologies, and well‐designed control measures deployable by conservation practitioners are urgently needed if these effects are to be mitigated, prevented or reversed. [ABSTRACT FROM AUTHOR]

Soil biota have strong effects on plants, but we have a poor understanding of how plant chemistry might modify these effects. We examined the effect of soil biota associated with an exotic invasive tree, Prosopis juliflora, versus that associated with native species, from seven sites across India on conspecifics and two other plant species. We then measured changes in species‐specific soil biota effects (identified as plant–soil feedbacks, PSFs) when leaf leachate from P. juliflora or from native plant species was added to soil containing respective live and sterile soil inoculum.We quantified the amino acid L‐tryptophan from leaf leachate of P. juliflora, Leucaena leucocephala (another invader), and two native species. We also tested effects of P. juliflora or native species soil inoculum amendment of tryptophan on P. juliflora, P. cineraria and L. leucocephala across seven sites. We then quantified the microbially metabolized derivatives of tryptophan, phytohormone indole‐3‐acetic acid (IAA) and intermediates after adding tryptophan into P. juliflora and native soils.Soil biota associated with P. juliflora generated positive effects on conspecifics and L. leucocephala, but negative effects on the native congener P. cineraria. When P. juliflora leaf leachate was added to soil with live P. juliflora inoculum, PSFs became more positive for P. juliflora and other species, compared to leaf leachate amended with sterile soil inoculum. Native leaf leachate interacted weakly with soil biota to impact biomass of conspecifics and heterospecifics.There was roughly 10× more tryptophan in the leaf leachate of P. juliflora than in the leaf leachate of other species. Tryptophan generally increased positive PSFs associated with P. juliflora relative to soil biota associated with other plant species. When tryptophan was added to live P. juliflora soil, IAA and its intermediates were produced at five of seven sites, and at four of these sites soil biota from P. juliflora had positive PSFs.Synthesis. These results provide the first experimental evidence that a chemical leached from the leaves of an invader regulates PSFs. Our results indicate that canopy effects and PSFs, which are usually studied independently, can interact in ways that strongly affect conspecifics and neighbouring species. [ABSTRACT FROM AUTHOR]

Contemporary ecosystems commonly support multiple non‐native species, which can alter community structure and directly or indirectly interact with one another. Novel interactions between invasive species can result in the facilitation or suppression of one or both invaders, making it important to understand the underlying mechanisms that affect invasion outcomes.Goldfish Carassius auratus and Chinese mystery snails Cipangopaludina chinensis have been introduced widely through the aquarium trade and co‐occur in their non‐native ranges. We used an outdoor mesocosm experiment to assess separate and combined effects of goldfish and mystery snails on urban pond community structure and to investigate whether goldfish and mystery snails affected one another positively, neutrally or negatively. We predicted that the distinct trophic levels of each species and the anti‐predator traits of mystery snails would lead to positive interactions between invaders and synergistic effects on the aquatic community.Goldfish directly increased turbidity and decreased filamentous algae, zooplankton, juvenile native snails and survival of amphibians via consumptive effects. Goldfish indirectly increased periphyton, phytoplankton and body size of surviving native green frogs Lithobates clamitans through trophic cascades and/or release from competition. Mystery snails had more subtle effects including reducing growth rates of native grey tree frogs Hyla versicolor, likely through competitive interactions.With goldfish present, mystery snails showed increased reproductive success (higher biomass of juveniles), likely due to indirect competitive release from other herbivores. In contrast to their effects on native snails, goldfish did not consume juvenile mystery snails, likely due to their unique traits (e.g. large size and shell thickness). Mystery snails slightly reduced goldfish growth but did not affect their survival.These results suggest unidirectional facilitation between invaders via indirect changes in the food web. Overall, our study found strong direct and indirect effects of invasive aquarium species, particularly goldfish, and provides experimental support for the invasional meltdown hypothesis within urban waterbodies, which are especially prone to invasions. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Simple Summary: Monk Parakeets are invasive birds globally. Through the construction of communal nests, they shape urban bird life. Monk Parakeets were introduced in Chile in 1972 and are now widely distributed in urban environments. Through the description of ecological parameters, we aim to assess the state of the Monk Parakeet in Santiago of Chile after 50 years of invasion. We characterized 1458 Monk Parakeets' nests on 546 trees, 91% of which were introduced species. Tasmanian blue gum and black locust were the trees preferred by parakeets for nesting. The average nest height was at 14.2 m with an occupancy rate of 89.7% and associated to nest height. During two reproductive seasons, chambers had an average of 4.5 eggs and 4.2 nestlings. Results and conclusions obtained will help in understanding the ecology of this avian invasion. Other urban metropolises such as Washington DC, Mexico City, Rome, Berlin, Paris, London, Tokyo, Nairobi, Casablanca and City of Singapore also suffer Monk Parakeet invasion. To aid in the understanding of Monk Parakeet invasive biology, we aim to contribute to better informed decisions in invasive synanthropic species management. Monk Parakeets are considered one of the most invasive bird species given its unique capacity among psittacines to build their own communal nests. Originally introduced as pets in houses from where they escaped or were released, they are currently considered invasive in more than 20 countries worldwide. This is the case in Chile, where Monk Parakeets were introduced during the 1970s. Between 2016 and 2019 we searched Monk Parakeets' nests structures in the Santiago metropolis region. We identified 1458 Monk Parakeets' communal nests on 546 trees belonging to 34 tree species. Ninety-one percent of the occupied trees were also introduced. Paraná pine and cedar of Lebanon were the tree species with highest abundance of nests, averaging more than four nests/tree/species, with 23 and 18 maximum number of nests, respectively. Tasmanian blue gum and black locust were selected by parakeets more often than expected, based on availability. From all trees, 24.6% denoted health problems and 47.3% were pruned. The average nest height was 14.2 m and nests were observed mainly in secondary branches (59.3%). The occupancy rate was 89.7% and was associated to nest height and type of branch. During two reproductive seasons we quantified eggs and nestlings in chambers averaging 4.5 and 4.2, respectively. We provide a rough population size estimate and the characteristics of Monk Parakeets nest and tree selectivity, aiming to characterize several decades of a neglected urban invasion to warrant strategies for improved management measures. [ABSTRACT FROM AUTHOR]

Positive interactions have been hypothesised to influence plant community dynamics and species invasions. However, their prevalence and importance relative to negative interactions remain unclear to understand community change and invasibility. We examined pairwise biotic interactions using over 50 years of successional data to assess the prevalence of positive interactions and their effects on each focal species (either native or exotic). We found that positive interactions were widespread and the relative frequency of positive and negative interactions varied with establishment stage and between native and exotic species. Specifically, positive interactions were more frequent during early establishment and less frequent at later stages. Positive interactions involving native species were more frequent and stronger than those between exotic species, reducing the importance of invasional meltdown on succession. Our study highlights the role of positive native interactions in shielding communities from biological invasion and enhancing the potential for long‐term resilience. [ABSTRACT FROM AUTHOR]

Non-native predators are known to cause ecological impacts through heightened consumption of resources and decimation of native species. One such species is Micropterus salmoides. Often introduced for sport fishing worldwide; it has been listed by International Union for Conservation of Nature and Natural Resources as one of the 100 of the world's worst invasive species. Whilst impacts conferred towards native prey are well known, its relationship with other non-native species has received much less attention. Therefore, the aim of this study was to experimentally investigate the feeding preference of M. salmoides towards native prey (Geophagus iporangensis) compared to non-native prey (Oreochromis niloticus and Coptodon rendalli) using comparative consumption and the Manly-Chesson selectivity index. We found a higher consumption by M. salmoides towards the non-native prey and a tendency for this non-native preference to increase when prey availability increased. Our results are in line with the hypothesis that the invasion of the non-native predator can be facilitated by the high abundance and reproductive rate of the non-native prey. This is relevant considering the interaction of multiple invaders in an ecosystem, in particular environments modified by humans. Interspecific relationships are complex and their understanding is necessary for environmental management decision-making. [ABSTRACT FROM AUTHOR]

Ongoing globalisation and climate change are causing plant species to invade new habitats and thereby alter biodiversity and ecosystem functioning. Since numbers of plant invasions continue to increase globally, it is crucial to investigate the effects of multiple co-occurring alien species on native communities. Furthermore, priority effects due to the earlier emergence of certain species affecting fitness of later arriving species can shape community structure and affect native species performance. We investigate in a common garden pot experiment the interactions among five alien-native species pairs. First we focus on the effect of growing with either one or two alien neighbour species on a native plant, second we alter the arrival time of the alien or native neighbour by 3 weeks. Generally, native species performance decreased when surrounded by two alien species compared to only one, although the magnitude of this effect varied depending on species, with one species even performing better with alien neighbours than in monoculture. Species performance greatly decreased when arriving second in the pot, for both native and alien species. In contrast, alien species tended to benefit more from arriving early. Given that we studied annual ruderal species, their potentially lower competitive ability might explain why we detected negative effects of late arrival. We highlight the need to further elucidate underlying mechanisms of small-scale invasion dynamics to achieve generalisations concerning the response of multiple alien and native plants given their species-specific differences in response to neighbour species and arrival time. [ABSTRACT FROM AUTHOR]

Background and aims: Naturalized alien and native plants can impact each other directly when they grow next to each other, but also indirectly through their soil legacies. These alien-native interactions can also be modified by the presence of a third alien or native species. However, it is unknown how the performance of co-growing alien and native species is affected by their soil legacies and by the presence of an additional species.In our two-phase plant-soil-feedback experiment, soils were first conditioned by eight herbaceous species, four of which are naturalized and four of which are native to Germany. We then grew all 16 pairwise alien-native species combinations on soil conditioned by the respective alien or native species, on a mixture of soils conditioned by both species or on control soil. Each pair of test plants was grown on these soils without or with an additional alien or native species.Soil conditioning, and particularly conspecific soil conditioning, reduced growth of the alien and native test plants. The addition of another species also reduced growth of the test plants. However, the negative conspecific soil-legacy effect on alien test plants was reduced when the additional species was also alien.The negative conspecific plant-soil feedback for alien and native plants in our study could promote their coexistence. However, as partial alleviation of negative conspecific effects on alien plants occurred with an additional alien species, the chances of coexistence of alien and native species might decrease when there are multiple alien species present.Methods: Naturalized alien and native plants can impact each other directly when they grow next to each other, but also indirectly through their soil legacies. These alien-native interactions can also be modified by the presence of a third alien or native species. However, it is unknown how the performance of co-growing alien and native species is affected by their soil legacies and by the presence of an additional species.In our two-phase plant-soil-feedback experiment, soils were first conditioned by eight herbaceous species, four of which are naturalized and four of which are native to Germany. We then grew all 16 pairwise alien-native species combinations on soil conditioned by the respective alien or native species, on a mixture of soils conditioned by both species or on control soil. Each pair of test plants was grown on these soils without or with an additional alien or native species.Soil conditioning, and particularly conspecific soil conditioning, reduced growth of the alien and native test plants. The addition of another species also reduced growth of the test plants. However, the negative conspecific soil-legacy effect on alien test plants was reduced when the additional species was also alien.The negative conspecific plant-soil feedback for alien and native plants in our study could promote their coexistence. However, as partial alleviation of negative conspecific effects on alien plants occurred with an additional alien species, the chances of coexistence of alien and native species might decrease when there are multiple alien species present.Results: Naturalized alien and native plants can impact each other directly when they grow next to each other, but also indirectly through their soil legacies. These alien-native interactions can also be modified by the presence of a third alien or native species. However, it is unknown how the performance of co-growing alien and native species is affected by their soil legacies and by the presence of an additional species.In our two-phase plant-soil-feedback experiment, soils were first conditioned by eight herbaceous species, four of which are naturalized and four of which are native to Germany. We then grew all 16 pairwise alien-native species combinations on soil conditioned by the respective alien or native species, on a mixture of soils conditioned by both species or on control soil. Each pair of test plants was grown on these soils without or with an additional alien or native species.Soil conditioning, and particularly conspecific soil conditioning, reduced growth of the alien and native test plants. The addition of another species also reduced growth of the test plants. However, the negative conspecific soil-legacy effect on alien test plants was reduced when the additional species was also alien.The negative conspecific plant-soil feedback for alien and native plants in our study could promote their coexistence. However, as partial alleviation of negative conspecific effects on alien plants occurred with an additional alien species, the chances of coexistence of alien and native species might decrease when there are multiple alien species present.Conclusion: Naturalized alien and native plants can impact each other directly when they grow next to each other, but also indirectly through their soil legacies. These alien-native interactions can also be modified by the presence of a third alien or native species. However, it is unknown how the performance of co-growing alien and native species is affected by their soil legacies and by the presence of an additional species.In our two-phase plant-soil-feedback experiment, soils were first conditioned by eight herbaceous species, four of which are naturalized and four of which are native to Germany. We then grew all 16 pairwise alien-native species combinations on soil conditioned by the respective alien or native species, on a mixture of soils conditioned by both species or on control soil. Each pair of test plants was grown on these soils without or with an additional alien or native species.Soil conditioning, and particularly conspecific soil conditioning, reduced growth of the alien and native test plants. The addition of another species also reduced growth of the test plants. However, the negative conspecific soil-legacy effect on alien test plants was reduced when the additional species was also alien.The negative conspecific plant-soil feedback for alien and native plants in our study could promote their coexistence. However, as partial alleviation of negative conspecific effects on alien plants occurred with an additional alien species, the chances of coexistence of alien and native species might decrease when there are multiple alien species present. [ABSTRACT FROM AUTHOR]

Increasing rates of invasions in ecosystems worldwide necessitate experiments to determine the role of biotic interactions in the success and impact of multiple alien species. Here, we examined competitive and facilitative interactions among various combinations of three widespread and often co‐occurring invaders: the zebra mussel Dreissena polymorpha, and the macrophytes Elodea canadensis and Elodea nuttallii.Using a mesocosm‐based, factorial experimental design, we assessed the effect of interspecific competition on macrophyte growth rates in the absence and presence at varying biomass of D. polymorpha.Growth rates (wet g/day) of E. canadensis and E. nuttallii were similar when grown in isolation. When grown together, in the absence of D. polymorpha, E. canadensis growth was not significantly reduced in the presence of E. nuttallii and vice versa. In the presence of D. polymorpha (26.0 ± 1 mm), monocultural growth of E. canadensis was largely unaffected, while E. nuttallii growth was strongly enhanced. Low (2.64 g) and medium (3.96 g) mussel biomass led to negative interspecific effects between E. canadensis and E. nuttallii; at high (5.28 g) mussel biomass, the effect of interspecific competition was negated.Overall, D. polymorpha alleviated competitive interactions between the two invasive macrophytes when all three species co‐occurred, and substantially enhanced growth of E. nuttallii with increasing mussel biomass, thereby suggesting a possible influence on the relative dominance of these macrophytes in the field.Our study demonstrates how facilitations can cause shifts in dominance among closely related invaders. The consequences of such facilitations for the structure and function of communities remain to be explored generally. [ABSTRACT FROM AUTHOR]

1. Interactions between invertebrate herbivores with different feeding modes are common on long‐lived woody plants. In cases where one herbivore facilitates the success of another, the consequences for their shared host plant may be severe. Eastern hemlock (Tsuga canadensis), a canopy‐dominant conifer native to the eastern U.S., is currently threatened with extirpation by the invasive stylet‐feeding hemlock woolly adelgid (Adelges tsugae). The effect of adelgid on invasive hemlock‐feeding folivores remains unknown. 2. This study evaluated the impact of feeding by hemlock woolly adelgid on gypsy moth (Lymantria dispar) larval preference for, and performance on, eastern hemlock. To assess preference, 245 field‐grown hemlocks were surveyed for gypsy moth herbivory damage and laboratory paired‐choice bioassays were conducted. To assess performance, gypsy moth larvae were reared to pupation on adelgid‐infested or uninfested hemlock foliage, and pupal weight, proportional weight gain, and larval period were analysed. 3. Adelgid‐infested hemlocks experienced more gypsy moth herbivory than did uninfested control trees, and laboratory tests confirmed that gypsy moth larvae preferentially feed on adelgid‐infested hemlock foliage. Gypsy moth larvae reared to pupation on adelgid‐infested foliage gained more weight than larvae reared on uninfested control foliage. 4. These results suggest that the synergistic effect of adelgid and gypsy moth poses an additional threat to eastern hemlock that may increase extirpation risk and ecological impact throughout most of its range. [ABSTRACT FROM AUTHOR]

Thestrengthofbioticinteractions withinanecological communityaffectsthe susceptibility of the community to invasion by introduced taxa. In microbial communities, cross-feeding is a widespread type of biotic interaction that has the potential to affect community assembly and stability. Yet, there is little understanding of how the presence of cross-feeding within a community affects invasion risk. Here, I develop a metabolite-explicit model where native microbial taxa interact through both cross-feeding and competition for metabolites. I use this model to study how the strength of biotic interactions, especially cross-feeding, influence whether an introduced taxon can join the community. I found that stronger cross-feeding and competition led to much lower invasion risk, as both types of biotic interactions lead to greater metabolite scarcity for the invader. I also evaluated the impact of a successful invader on community composition and structure. The effect of invaders on the native community was greatest at intermediate levels of cross-feeding; at this ‘critical’ level of cross-feeding, successful invaders generally cause decreased diversity, decreased productivity, greater metabolite availability, and decreased quantities of metabolites exchanged among taxa. Furthermore, these changes resulting from a successful primary invader made communities further susceptible to future invaders. The increase in invasion risk was greatest when the network of metabolite exchange between taxa was minimally redundant. Thus, this model demonstrates a case of invasional meltdown that is mediated by initial invaders disrupting the metabolite exchange networks of the native community. [ABSTRACT FROM AUTHOR]

The availability of compatible mutualistic soil microbes could influence the invasion success of non‐native plant species. Specifically, there may be spatial variation in the distribution of compatible microbes, and species‐specific variation in plant host ability to associate with available microbes. Although either or both factors could promote or limit invasion, the scale over which most studies are conducted makes it difficult to examine these two possibilities simultaneously. However, this is critical to identifying a role of soil microbes in invasion.A series of recent research projects focused on interactions between Australian Acacia and nitrogen‐fixing bacteria (rhizobia) at multiple spatial scales, from the local to the inter‐continental, has allowed us to evaluate this question. Collectively, this research reveals that nodulation, performance and rhizobial community composition are all broadly similar across spatial scales and differentially invasive species.Synthesis and applications. We argue that current research provides convincing evidence that interactions with rhizobia do not determine invasion success in Acacia, but instead highlights key knowledge gaps that remain unfilled. Importantly, the ease with which non‐native Acacia species form mutualistic associations with rhizobia, regardless of invasive status, highlights the critical need to understand the impacts of all non‐native Acacia on native soil communities. [ABSTRACT FROM AUTHOR]

Widespread habitat-forming invaders inhabiting marinas, such as the spaghetti bryozoan Amathia verticillata, allow exploring facilitation processes across spatiotemporal contexts. Here we investigate the role of this bryozoan as habitat for native and exotic macrofaunal assemblages across different ecoregions of Western Mediterranean and East Atlantic coasts, and a monthly variation over a year. While only 7 (all peracarid crustaceans) of the 54 associated species were NIS, they dominated macrofaunal assemblages in terms of abundance, raising the potential for invasional meltdown. NIS richness and community structure differed among marinas but not among ecoregions, highlighting the importance of marina singularities in modulating facilitation at spatial scale. Despite facilitation did not depend on bryozoan abundance fluctuations, it was affected by its deciduous pattern, peaking in summer and disappearing in late winter. Monitoring A. verticillata in marinas, especially in summer periods, may improve the detection and management of multiple associated NIS. • Amathia verticillata host a diverse epifaunal community. • The bryozoan was prone to invasional meltdown. • Seven NIS found, all peracaridean crustaceans. • NIS richness and community structure differed among marinas but not among ecoregions. • Facilitation did not depend on bryozoan abundance. [ABSTRACT FROM AUTHOR]

Background: Alliaria petiolata (garlic mustard) is a widespread allelopathic invasive species in North America, but despite extensive research we do not fully understand how it impacts plant communities. Aims: We tested the hypothesis that A. petiolata facilitates the introduction of additional alien species as predicted by the invasional meltdown hypothesis. Methods: We conducted co-occurrence surveys to compare the numbers of alien and native species in communities surrounding A. petiolata plants with those surrounding an allelopathic native species. We also conducted an experiment to compare recruitment of native and alien species in pots dominated by either A. petiolata or a native species. Results: A. petiolata plots in the co-occurrence surveys had relatively high proportions of bare ground, suggesting habitat modification, plus lower species richness and disproportionately fewer alien species. There was a greater tendency for native species than alien species to establish in A. petiolata pots. Conclusions: A. petiolata does not promote additional alien species, and in fact may be a more effective competitor against alien than native species in North America. This could help explain its invasive success in disturbed habitats, and illustrates the need to consider separately alien and native species when assessing the outcomes of competition. [ABSTRACT FROM AUTHOR]

Aim: Mutualisms are often disrupted for plants introduced to new ranges, yet many of these plants have managed to obtain effective mutualistic associations in their new ranges. There are two potential pathways for non‐native plants to reassemble mutualisms: cointroduction (i.e. familiar associations with cointroduced mutualists) or ecological fitting (i.e. forming or adapting novel associations with resident native mutualists). We assessed the importance of each pathway for mutualist reassembly in four Australian Acacia species (A. baileyana, A. dealbata, A. decurrens and A. melanoxylon) and their associated nitrogen‐fixing rhizobial symbionts in two non‐native locations. Location: Native ranges of acacias in south‐eastern Australia and two non‐native ranges in New Zealand and South Africa. Methods: Rhizobia associated with each acacia species in each country were isolated and identified based on DNA sequencing of the housekeeping recA gene and the symbiotic nodA gene. Separate phylogenies were reconstructed for each gene region to infer biogeographic histories of acacia‐associated rhizobia. Selected rhizobial strains for each acacia species by country combination were used as inocula in a glasshouse experiment and early growth kinetics and nitrogen fixation efficiency of acacia seedlings were compared between inoculum treatments to determine symbiotic effectiveness. Results: All isolated rhizobial strains belonged to the genus Bradyrhizobium. Phylogenetic analyses revealed almost no country‐ or species‐specific clusters of these strains for either gene region and indicated that most acacia‐associated bradyrhizobia in New Zealand and South Africa were cointroduced from Australia. These results were supported by little variation in the growth performances of acacia seedlings, irrespective of inoculum treatment. Main conclusions: This study revealed that cointroduction of Australian acacias and their rhizobia may be more prevalent than previously thought. Additionally, a single rhizobium cointroduction event may be sufficient to facilitate the establishment of effective mutualisms in numerous Acacia species, potentially leading to an invasion meltdown. [ABSTRACT FROM AUTHOR]

Positive interactions enhance biodiversity and ecosystem function, but can also exacerbate biological invasions. Facilitation of exotic invaders by exotic foundation species (invasional meltdown) has been studied extensively, but facilitation of exotic invaders by native foundation species has attracted less attention. Specifically, very few studies have examined the extent that native foundation species facilitate native and exotic competitors. Understanding the processes that mediate interactions between native and exotic species can help explain, predict, and improve management of biological invasions. Here, we examined the effects of native foundation shrubs on the relative abundance of the annual plant community - including native and exotic taxa - from 2015-2018 in a desert ecosystem at Carrizo Plain National Monument, California, USA (elevation: 723 m). Shrub effects varied by year and by the identity of annual species, but shrubs consistently enhanced the abundance of the annual plant community and facilitated both native (n=17 species) and exotic (n=4 species) taxa. However, at the provenance level, exotic annuals were facilitated 2.75 times stronger in abundance than native annuals, and exotic annuals were always more abundant than natives both near and away from shrubs. Our study reaffirms facilitation as an important process in the organisation of plant communities and confirms that both native and exotic species can form positive associations with native foundation species. However, facilitation by native foundation species can exacerbate biological invasions by increasing the local abundance of exotic invaders. Thus, the force of facilitation can have a dark side relevant to ecosystem function and management. [ABSTRACT FROM AUTHOR]

The aim of the study is to present a complete and updated fish inventory of inland waters of the Emilia-Romagna region, Northern Italy, and to highlight the presence of fully exotic fish communities. Overall, based on 208 sampling locations, the observed fish fauna consisted of 45 species, 22 native and 23 exotics. A significant element of the inventory is the identification of xenodiversity hotspots (spatially clustered sites, one lowland and one upland region), where a complete substitution of native species by exotic species was observed (in total seven sites in the lowland and two sites in the upland with no native species presence). These xenodiversity hotspots were found to host specific combinations of exotic species, which may be able to constitute balanced exotic communities. The hotspots of the lowland region are located in the northeast lowland part of the territory, hosting exotic species combinations mainly composed by wels catfish (Silurus glanis Linnaeus, 1758, a large predator), common carp (Cyprinus carpio Linnaeus, 1758, a large benthivore), crucian carp (Carassius spp., a small-bodied generalist) and other less dominant exotic species. The hotspots in the upland region were located in the southwest part of the territory and were dominated by only one exotic species (rainbow trout, Oncorhynchus mykiss (Walbaum, 1792)). A difference between these xenodiversity hotspots is that in the lowland the introductions were mostly unintentional and are not continued, while in the upland the introduction of rainbow trout is intentional and currently carried out by local fishermen. [ABSTRACT FROM AUTHOR]

The invasion paradox refers to the existence of biotic resistance and biotic acceptance hypotheses. According to the biotic resistance hypothesis, the higher the richness present in a community, the more resistant it is against invaders. In contrast, the biotic acceptance hypothesis states that if the environment is suitable for a high richness of native species, it will be similarly suitable for establishment of non-native species. Previous studies on terrestrial plant communities considered the scale-dependent nature of native and non-native relationships: the relationship tends to be negative at small scales (biotic resistance) and positive at broader scales (biotic acceptance). We tested the hypothesized role of spatial scale on the relationship between non-native and native species richness using a spatially nested design and a long-term dataset (146 communities sampled during 13 years) of fish species richness from the upper Paraná River floodplain, Brazil. Contrary to expectations, non-native fish species richness was positively correlated with native species richness at all spatial scales. Mobility of vertebrates, mode of invasion, and environmental disturbance may affect the role of spatial scale in potentially mediating relative importance of biotic acceptance versus resistance. In this context, the present study provides a unique contribution towards resolving the invasion paradox. [ABSTRACT FROM AUTHOR]

Abstract: Aim: Darwin's naturalization hypothesis states that dissimilarity to native species may benefit alien species establishment due to empty niches and reduced competition. We here add a new dimension to large‐scale tests of community invasibility, investigating the role that previously established alien species play in facilitating or hindering new invasions in plant communities. Location: Permanent grasslands across France (including mainland and Corsica), as a receding ecosystem of great conservation importance. Methods: Focusing on 121 alien plant species occurring in 7,215 vegetation plots, we quantified biotic similarity between new invaders and resident alien species (i.e., alien species with longer residence times) based on phylogenetic and trait distances. Additionally, we calculated distances to native species for each alien species and plot. Using multispecies distribution models, we analysed the influence of these biotic similarity measures and additional covariates on establishment success (presence/absence) of new invaders. Results: We found that biotic similarity to resident alien species consistently increased establishment success of more recently introduced species. Phylogenetic relatedness to previous invaders had an equally strong positive effect as relatedness to native species. Conversely, trait similarity to natives hindered alien establishment as predicted by Darwin's naturalization hypothesis. These results highlight that various mechanisms may act simultaneously to determine alien establishment success. Main conclusions: Our results suggest that, with greater similarity among alien species, invasion success increases. Such a pattern may arise either due to actual facilitation among invaders or as a result of weaker competitive interactions among invaders than between native and alien species, leading to an indirect facilitative effect. Alternatively, recent environmental changes (e.g., eutrophication, climate change) may have added new environmental filters. Determining how initial invasions might pave the road for subsequent invasions is crucial for effective multispecies management decisions and contributes a new aspect to our understanding of community assembly. [ABSTRACT FROM AUTHOR]

In the absence of empirical evidence, invasive species are often assumed to have negative impacts because of their conspicuously high abundance. The giant African land snail Achatina ( Lissachatina) fulica is one such invader where its impact in natural ecosystems remains completely untested. On Christmas Island (Indian Ocean), A. fulica has become established across large tracts of rainforest following the impacts of invasive yellow crazy ant ( Anoplolepis gracilipes) in mutualism with non-native scale insects. Yellow crazy ants facilitate the secondary invasion of A. fulica by extirpating native red land crabs ( Gecarcoidea natalis) that are normally effective predators of A. fulica. We used a multifaceted approach to investigate some potential impacts of abundant A. fulica in invaded rainforest. Over the course of a wet season, diel activity transects showed that A. fulica consumed detrital material almost exclusively. However, stable isotope analysis did not confidently identify A. fulica as a predominantly detritivorous species. We found no statistically significant treatment effects of A. fulica exclusion on standing leaf litter and seedling recruitment processes during a 6-month manipulative field study. However, litter cover and biomass did remain slightly higher where A. fulica were excluded, albeit with overlapping confidence intervals with control plots. Our study constitutes the first empirical test for impact of A. fulica in a natural ecosystem and suggests that for Christmas Island rainforest, this species is not a damaging invader. Other studies will need to assess the impacts of A. fulica in other natural areas before these findings could be considered broadly applicable. [ABSTRACT FROM AUTHOR]

Positive interactions between exotic species may increase ecosystem-level impacts and potentially facilitate the entry and spread of other exotic species. Invader-facilitated invasion success-'secondary invasion'-is a key conceptual aspect of the well-known invasional meltdown hypothesis, but remains poorly defined and empirically underexplored. Drawing from heuristic models and published empirical studies, we explore this form of 'secondary invasion' and discuss the phenomenon within the recognized conceptual framework of the determinants of invasion success. The term 'secondary invasion' has been used haphazardly in the literature to refer to multiple invasion phenomena, most of which have other more accepted titles. Our usage of the term secondary invasion is akin to 'invader-facilitated invasion,' which we define as the phenomenon in which the invasion success of one exotic species is contingent on the presence, influence, and impacts of one or more other exotic species. We present case studies of secondary invasion whereby primary invaders facilitate the entry or establishment of exotic species into communities where they were previously excluded from becoming invasive. Our synthesis, discussion, and conceptual framework of this type of secondary invasion provides a useful reference to better explain how invasive species can alter key properties of recipient ecosystems that can ultimately determine the invasion success of other species. This study increases our appreciation for complex interactions following invasion and highlights the impacts of invasive species themselves as possible determinants of invasion success. We anticipate that highlighting 'secondary invasion' in this way will enable studies reporting similar phenomena to be identified and linked through consistent terminology. [ABSTRACT FROM AUTHOR]

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 genusCarpobrotus. Conservation measures focus on the shearwatersPuffinus yelkouanandCalonectris 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. [ABSTRACT FROM PUBLISHER]

Aim Typically, non-indigenous species have been studied in relation to either the invaded (recipient) community or the donor community. However, we still lack a broad understanding of the mechanisms underlying the establishment of non-indigenous species that combines both perspectives. Since the opening of the Suez Canal, hundreds of species have invaded the Mediterranean from the Red Sea, forming a unique system in which the entire species pool (donor, non-indigenous and recipient) is known. Focusing on species ecological traits, we assess the drivers underlying the establishment of non-indigenous species from the onset of the invasion to the present. Location The Mediterranean and Red Seas. Methods We compiled traits for shallow-water hard-bottom-associated fishes in the Mediterranean (recipient community) and the Red Sea (donor community). We compared the multivariate community trait structure of non-indigenous species with the donor and recipient assemblages, using a novel method to objectively assign trait-weights, thereby increasing the robustness of the results. Patterns were contrasted with two types of null models which provide distinct insights into the ecological processes. Results Non-indigenous fishes are very diverse ecologically, substantially increasing the total community trait diversity of the Mediterranean. Trait similarity between non-indigenous and indigenous Mediterranean species was lower than expected, indicating that non-indigenous fishes tend to occupy relatively vacant niches within the Mediterranean. However, we further found that over time non-indigenous species display increased trait similarity to indigenous Mediterranean species. This suggests that trait constraints associated with successful establishment are weakening. Main conclusions Non-indigenous species establish in relatively vacant ecological niches, opening the possibility for predicting establishment using ecological traits. However, the weakening of trait constraints through time suggests that forecasting future establishment may be difficult. Faced with an accelerating pace of invasion, it appears that the Mediterranean is going to be transformed into an extension of the Red Sea in terms of trait and species composition. [ABSTRACT FROM AUTHOR]

The entry of secondary invaders into, or their expansion within, native communities is contingent on the changes wrought by other (primary) invaders. When primary invaders have altered more than one property of the recipient community, standard descriptive and modeling approaches only provide a best guess of the mechanism permitting the secondary invasion. In rainforest on Christmas Island, we conducted a manipulative field experiment to determine the mechanism of invasion success for a community of land snails dominated by non-native species. The invasion of rainforest by the yellow crazy ant ( Anoplolepis gracilipes) has facilitated these land snails, either by creating enemy-free space and/or increased habitat and resources (in the form of leaf litter) through the removal of the native omnivorous-detritivorous red land crab ( Gecarcoidea natalis). We manipulated predator densities (high and low) and leaf litter (high and low) in replicated blocks of four treatment combinations at two sites. Over the course of one wet season (five months), we found that plots with high leaf litter biomass contained significantly more snails than those with low biomass, regardless of whether those plots had high or low predation pressure, at both the site where land crabs have always been abundant, and at the site where they have been absent for many years prior to the experiment. Each site was dominated by small snail species (<2 mm length), and through handling size and predation experiments we demonstrated that red crabs tend not to handle and eat snails of that size. These results suggest that secondary invasion by this community of non-native land snails is facilitated most strongly by habitat and resource augmentation, an indirect consequence of red land crab removal, and that the creation of enemy-free space is not important. By using a full-factorial experimental approach, we have confidently determined-rather than inferred-the mechanism by which primary invaders indirectly facilitate a community of secondary invaders. [ABSTRACT FROM AUTHOR]

Biological invasions are a key threat to freshwater biodiversity, and identifying determinants of invasion success is a global conservation priority. The establishment of introduced species is predicted to be hindered by pre-existing, functionally similar invasive species. Over a five-year period we, however, find that in the River Lee (UK), recently introduced non-native virile crayfish (Orconectes virilis) increased in range and abundance, despite the presence of established alien signal crayfish (Pacifastacus leniusculus). In regions of sympatry, virile crayfish had a detrimental effect on signal crayfish abundance but not vice versa. Competition experiments revealed that virile crayfish were more aggressive than signal crayfish and outcompeted them for shelter. Together, these results provide early evidence for the potential over-invasion of signal crayfish by competitively dominant virile crayfish. Based on our results and the limited distribution of virile crayfish in Europe, we recommend that efforts to contain them within the Lee catchment be implemented immediately. [ABSTRACT FROM AUTHOR]

When an invasive predator encounters native and invasive prey, two scenarios are possible: the predator may benefit from the presence of naïve native prey or choose prey from its region of origin, reflecting their common evolutionary history., To determine interactions between an invasive predator and native and invasive prey, we used the Ponto-Caspian racer goby Babka gymnotrachelus as predator and gammarids as prey: native Gammarus fossarum and Ponto-Caspian Dikerogammarus villosus and Pontogammarus robustoides. We hypothesised that prey origin would affect fish preferences and growth rate and conducted a series of laboratory experiments on fish predation and growth and estimated profitability of prey of different origin., The goby preferred native prey to the Ponto-Caspian gammarids, irrespective of prey motility, the presence of shelters or waterborne chemical cues. Moreover, fish grew better when fed native prey., Thus, we suggest that fish selectivity was based on the assessment of prey quality during direct contact with gammarids. A diet consisting of Ponto-Caspian gammarids did not facilitate an invader originating from the same region, which benefited more from the presence of a local prey species., Ponto-Caspian gammarids and gobies are successful invaders in inland waters, usually main rivers. The gobies, in contrast to the invasive gammarids, enter smaller tributaries that serve as refugia for native gammarids. We show that the gobies may benefit from the presence of native prey species in such locations. [ABSTRACT FROM AUTHOR]

With accelerating rates of invasion being documented in many ecosystems, communities of interacting invasive species are becoming increasingly common. Opposing theories predict that invaders can either hinder or promote one another's success. Additionally, evidence suggests that co-occurring invaders can interact to amplify or mitigate one another's impacts on ecosystems. However, there has not been a quantitative review on interactions among multiple invasive animals. Here I use a meta-analysis approach to show that, across a global scale, the mean interaction among invaders was to reduce one another's performance. This pattern was consistent when considering interactions between marine animals but interactions were neutral overall in terrestrial and freshwater ecosystems. Crucially, individual studies showed that neutral interactions were the most common interaction type. Further, I demonstrate that the combined ecological impacts of multiple invaders were frequently the sum of their independent effects (additive) but the mean effect was non-additive and less than predicted (antagonistic). In both meta-analyses, the disparity between the most frequent and mean interaction type indicates that case studies of multiple invasions commonly have different outcomes to global trends. These results will help predict how co-occurring invasive animals interact and assist in developing management strategies for problematic invaders in our changing world. [ABSTRACT FROM AUTHOR]

Ecosystems with multiple exotic species may be affected by facilitative invader interactions, which could lead to additional invasions (invasional meltdown hypothesis). Experiments show that one-way facilitation favours exotic species and observational studies suggest that reciprocal facilitation among exotic species may lead to an invasional meltdown. We conducted a mesocosm experiment to determine whether reciprocal facilitation occurs in wetland communities., We established communities with native wetland plants and aquatic snails. Communities were assigned to treatments: control (only natives), exotic snail ( Pomacea maculata) invasion, exotic plant ( Alternanthera philoxeroides) invasion, sequential invasion (snails then plants or plants then snails) or simultaneous invasion (snails and plants)., Pomacea maculata preferentially consumed native plants, so A. philoxeroides comprised a larger percentage of plant mass and native plant mass was lowest in sequential (snail then plant) invasion treatments., Even though P. maculata may indirectly facilitate A. philoxeroides, A. philoxeroides did not reciprocally facilitate P. maculata. Rather, ecosystems invaded by multiple exotic species may be affected by one-way facilitation or reflect exotic species' common responses to abiotic factors or common paths of introduction. [ABSTRACT FROM AUTHOR]

Humans have effectively transported thousands of species around the globe and, with accelerated trade; the rate of introductions has increased over time. Aquatic ecosystems seem at particular risk from invasive species because of threats to biodiversity and human needs for water resources. Here, we review some known aspects of aquatic invasive species (AIS) and explore several new questions. We describe impacts of AIS, factors limiting their dispersal, and the role that humans play in transporting AIS. We also review the characteristics of species that should be the greatest threat for future invasions, including those that pave the way for invasions by other species ('invasional meltdown'). Susceptible aquatic communities, such as reservoirs, may serve as stepping stones for invasions of new landscapes. Some microbes disperse long distance, infect new hosts and grow in the external aquatic medium, a process that has consequences for human health. We also discuss the interaction between species invasions and other human impacts (climate change, landscape conversion), as well as the possible connection of invasions with regime shifts in lakes. Since many invaders become permanent features of the environment, we discuss how humans live with invasive species, and conclude with questions for future research. [ABSTRACT FROM AUTHOR]

The movement of species is one of the most pervasive forms of global change, and few ecosystems remain uninvaded by nonnative species. Studying species interactions is crucial for understanding their distribution and abundance, particularly for nonnative species because interactions may influence the probability of invasion and consequent ecological impact. Interactions among nonnatives are relatively understudied, though the likelihood of nonnative species co-occurrence is high. We quantify and describe the types of interactions among nonnative plants and determine what factors affect interaction outcomes for ecosystems globally. We reviewed 65 studies comprising 201 observations and recorded the interaction type, traits of the interacting species, and study characteristics. We conducted a census of interaction types and a meta-analysis of experiments that tested nonnative competition intensity. Both methods showed that negative and neutral interactions prevailed, and a number of studies reported that the removal of a dominant nonnative led to competitive release of other nonnatives. Positive interactions were less frequently reported and positive mean effect sizes were rare, but the plant characteristics nitrogen fixation, life cycle (annual or perennial), and functional group significantly influenced positive interactions. Positive interactions were three times more frequent when a neighboring nonnative was a nitrogen fixer and 3.5 times lower when a neighboring nonnative was an annual. Woody plants were two or four times more likely to have positive interactions relative to grasses or herbs, respectively. The prevalence of negative interactions suggests that managers should prepare for reinvasion of sites when treating dominant nonnatives. Though positive interactions were infrequent, managers may be able to anticipate positive interactions among nonnatives based upon traits of the co-occurring invaders. Predicting positive nonnative interactions is an important tool for determining habitat susceptibility to a particular invasion and for prioritizing management of nonnatives with a higher likelihood of positive interactions. [ABSTRACT FROM AUTHOR]

The outcome of multiple invasions from a common origin may lead to facilitative interactions because the invaders have co-evolved under similar environmental conditions. This outcome is often referred to as invasional meltdown, with a resultant increase in invasive species and a decline in native species richness and abundance., This study seeks to assess the full scope of the threat posed by a high-risk group of 23 freshwater invaders originating from the Ponto-Caspian region (south-east Europe) across Great Britain. Ponto-Caspian invaders constitute a group of special concern because they have recently caused a large-scale invasion into western Europe., According to a literature review, 76% of reported interactions between Ponto-Caspian invaders are positive (mostly provision of food and commensalism) or neutral (habitat partitioning). Negative interactions (mostly predation) were rare, thus highlighting the ability of Ponto-Caspian invaders to coexist., At least 14 out of the 23 Ponto-Caspian organisms investigated are well-established in the Rhine estuary and Dutch ports. Four of them ( Hemimysis anomala, Dikerogammarus villosus, D. haemobaphes and Hypania invalida) have recently established in Great Britain. Regression models suggest the rest are under a critical risk of being transported, with four species predicted to have arrived already to Great Britain: Echinogammarus ischnus, Jaera istri, Limnomysis benedeni and D. bispinosus., According to species distribution models, the cumulative risk of invasion of multiple Ponto-Caspian species, thus invasional meltdown, is highest in the south-east of England and decreases to the north and west., Great Britain might be on the brink of invasional meltdown, and as a consequence, confronting the problem of Ponto-Caspian invasive species is a vital element for national biosecurity., Synthesis and applications. The predictive models and maps developed in this study provide a means for an evidence-based prioritization of species and habitats for the management of existing and future invasions of Ponto-Caspian species. This integrated approach can be easily applied to risk assess other groups of species and habitats. [ABSTRACT FROM AUTHOR]

Banana passionfruit (Passiflora tripartita var. mollissima) is a noxious vine that is invasive in forest patches in coastal regions throughout New Zealand. We investigated the dispersal mechanisms that facilitate its spread in the Marlborough Sounds. To find out which animals act as dispersers, we monitored tagged fruits in the field. Fruits were removed quickly after ripening. Significantly fewer fruits were wholly removed from off-road locations than locations on road edges, but removal was high in both cases (70% and 93% respectively) indicating likely dispersal by both humans and wild animals. We found no evidence of dispersal by birds, but infrared cameras documented possums and rats consuming fruits in the field. To investigate the effect of fruit handling on germination, we compared germination success between hand-cleaned seeds, fleshy seeds and intact fruits in the field and glasshouse. Seeds germinated readily in all treatments, with no significant difference between treatments, so seeds do not require frugivore handling to germinate. In addition, we measured the germination of seeds extracted from 1.5 kg of feral pig faeces collected from Sus scrofa at Te Weuweu Bay and from faeces from captive possums (Trichosurus vulpecula) and Norway rats (Rattus norvegicus). Seeds extracted from pig and possum droppings readily germinated, with final germination success not significantly different from hand-cleaned or in-flesh seeds. None of the few intact seeds excreted by Norway rats germinated. Banana passionfruit has formed invasive mutualisms for both pollination (with introduced bees) and dispersal (with introduced mammals), exacerbating its spread. Weed management should combine direct weed control with limiting the spread of banana passionfruit, by managing feral pigs and possums. [ABSTRACT FROM AUTHOR]

Mutualisms structure ecosystems and mediate their functioning. They also enhance invasions of many alien species. Invasions disrupt native mutualisms, often leading to population declines, reduced biodiversity, and altered ecosystem functioning. Focusing on three main types of mutualisms (pollination, seed dispersal, and plant-microbial symbioses) and drawing on examples from different ecosystems and from species- and community-level studies, we review the key mechanisms whereby such positive interactions mediate invasions and are in turn influenced by invasions. High interaction generalization is "the norm" in most systems, allowing alien species to infiltrate recipient communities. We identify traits that influence invasiveness (e.g., selfing capacity in plants, animal behavioral traits) or invasibility (e.g., partner choice in mycorrhizas/rhizobia) through mutualistic interactions. Mutualistic disruptions due to invasions are pervasive, and subsequent cascading effects are also widespread. Ecological networks provide a useful framework for predicting tipping points for community collapse in response to invasions and other synergistic drivers of global change. [ABSTRACT FROM AUTHOR]

Facilitation among species may promote non-native plant invasions through alteration of environmental conditions, enemies or mutualists. However, the role of non-trophic indirect facilitation in invasions has rarely been examined., We used a long-term field experiment to test for indirect facilitation by invasions of Microstegium vimineum (stiltgrass) on a secondary invasion of Alliaria petiolata (garlic mustard) by introducing Alliaria seed into replicated plots previously invaded experimentally by Microstegium., Alliaria more readily colonized control plots without Microstegium but produced almost seven times more biomass and nearly four times as many siliques per plant in Microstegium-invaded plots. Improved performance of Alliaria in Microstegium-invaded plots compared to control plots overwhelmed differences in total number of plants such that, on average, invaded plots contained 327% greater total Alliaria biomass and 234% more total siliques compared to control plots., The facilitation of Alliaria in Microstegium-invaded plots was associated with an 85% reduction in the biomass of resident species at the peak of the growing season and significantly greater light availability in Microstegium-invaded than control plots early in the growing season., Synthesis. Our results demonstrate that an initial plant invasion associated with suppression of resident species and increased resource availability can facilitate a secondary plant invasion. Such positive interactions among species with similar habitat requirements, but offset phenologies, may exacerbate invasions and their impacts on native ecosystems. [ABSTRACT FROM AUTHOR]

Ecological interactions among invasive species can affect not only the success of the invaders, but also their impact on ecosystems in the invaded range. In Australia, both dung beetles (subfamily Scarabaeinae) and cane toads ( Rhinella marina) were introduced for biocontrol: the beetles to break down bovine faeces piles (cowpats) that otherwise accumulate and reduce pasture productivity, and the cane toad to consume scarab beetles that eat sugarcane and thus reduce sugar production. The dung beetles have been a success, whereas the toads have been a failure. Our experimental studies show that as well as impacting native fauna directly, cane toads reduce the rate of cowpat breakdown by consuming dung beetles. In the laboratory, dehydrated toads actively sought out cowpats based on scent cues, and in field enclosures, the presence of a cane toad significantly reduced rates of cowpat decomposition. Although toads have benefited from agricultural activities, their spread across Australia likely has reduced the effectiveness of one of the most successful biocontrol programmes ever conducted in that continent. [ABSTRACT FROM AUTHOR]

Biological invasions depend in part on the resistance of native communities. Meta-analyses of terrestrial experiments demonstrate that native primary producers and herbivores generally resist invasions of primary producers, and that resistance through competition strengthens with native producer diversity. To test the generality of these findings, we conducted a meta-analysis of marine experiments. We found that native marine producers generally failed to resist producer invasions through competition unless the native community was diverse, and this diversity effect was weaker in marine than in terrestrial systems. In contrast, native consumers equally resisted invasive producers in both ecosystems. Most marine experiments, however, tested invasive consumers and these invasions were resisted more strongly than were producer invasions. Given these differences between ecosystems and between marine trophic levels, we used a model-selection approach to assess if factors other than the resistance mechanism (i.e. competition vs. consumption) are more important for predicting marine biotic resistance. These results suggest that understanding marine biotic resistance depends on latitude, habitat and invader taxon, in addition to distinguishing between competition with and consumption by native species. By examining biotic resistance within and across ecosystems, our work provides a more complete understanding of the factors that underlie biological invasions. [ABSTRACT FROM AUTHOR]

Abstract: Though biological invasion studies have proliferated in recent decades, a consistent emphasis remains on the study of single-species invasions. Here, we juxtapose the number of invasive plants reported as co-occurring within conservation habitats in one of the most comprehensive global conservation management databases (The Nature Conservancy’s Conservation Projects) with the number of published studies that address impacts of co-occurring invasive plants. We reviewed 86 conservation projects and 153 peer-reviewed publications and found that only one-third of studies mentioned co-occurring invaders, although over two-thirds of habitats were multiply invaded, indicating researchers are more likely to study single invaders, even though conservation managers are more often faced with multiple invaders in a given habitat. Of those studies focused on multiple invasives, the majority did not attempt to differentiate impacts caused by species when found alone or with other invaders and instead either treated co-occurring invaders together as a single, undifferentiated group or compared impacts between invasive plant monocultures. Less than 6% of all studies analyzed invader interactions. The high prevalence of co-occurring invasive plants should encourage more research on multiple invaders, which may better inform prioritization of which species to manage. Specifically, we suggest research on how effects of multiple invaders differ from those of single invaders, what types of interactions (facilitative, competitive, neutral) are most commonly found between invaders, and what effects interactions might have on the overall impact (additive or non-additive) of the individual invader. Though we acknowledge the challenge of studying multiple invaders, there is a critical need to address these questions to make invasion research more relevant to conservation programs. [Copyright &y& Elsevier]

Abstract: In North America, many species of European earthworms have been introduced to northern forests. Facilitative or competitive interactions between these earthworm species may result in non-additive effects on native plant and animal species. We investigated the combined versus individual effects of the litter-dwelling earthworm Dendrobaena octaedra Savigny, 1826 and the deep-burrowing species Lumbricus terrestris L., 1758 on microarthropod assemblages from boreal forest soil by conducting a mesocosm experiment. Soil cores from earthworm-free areas of northern Alberta, Canada, were inoculated with D. octaedra alone, L. terrestris alone, both worm species together, or no earthworms. After 4.5 months, microarthropods were extracted from the soil, counted, and identified to higher taxa. Oribatid mites were further identified to family and genus. Abundance of microarthropods was significantly lower in the treatment containing both species than in the no earthworm treatment and the L. terrestris treatment. Oribatida and Prostigmata/Astigmata differed significantly among treatments and were lowest in the treatment containing both earthworm species, followed by the D. octaedra treatment, although post-hoc pairwise comparisons were not significant. Within the Oribatida, composition differed between the control and L. terrestris treatments as compared to the D. octaedra and both-species treatments, with Suctobelbella and Tectocepheus in particular having higher abundances in the control treatment. Effects of the two earthworm species on microarthropods were neither synergistic nor antagonistic. Our results indicate that earthworms can have strong effects on microarthropod assemblages in boreal forest soils. Future research should examine whether these changes have cascading effects on nutrient cycling, microbial communities, or plant growth. [Copyright &y& Elsevier]

Invasive species represent a major threat with both direct and indirect effects on natural ecosystems, including effects on established and coevolved relationships. In a series of experiments, we examined how the interaction between two native species, a unionid mussel ( Unio pictorum) and the European bitterling ( Rhodeus amarus), a fish that parasitises unionids, was affected by the non-native zebra mussel ( Dreissena polymorpha). The zebra mussel fouls hard substrates, including shells of living unionids, and its presence is often associated with a decrease in population density of native unionid mussels. Bitterling lay their eggs into live unionids and the embryos develop inside their gills. Using a range of zebra mussel densities, we demonstrated that zebra mussel fouling had a negative effect on the number of bitterling eggs inside the mussel host, with abundances of 5-10 zebra mussels (shell size 15-25 mm) per unionid critical for bitterling ability to utilise the host. In a further experiment, we found that bitterling did not discriminate between unfouled unionids and those fouled with five zebra mussels. Most ovipositions into fouled hosts, however, were unsuccessful as eggs failed to reach the unionid gills. We discuss implications of such unsuccessful ovipositions for bitterling recruitment and population dynamics. [ABSTRACT FROM AUTHOR]

Abstract: We evaluated the mechanisms that determine the increasing abundance of Pinus halepensis in mountain pampean grasslands in Argentina that is associated with the continuous presence of feral horses. We hypothesized that direct and indirect effects of feral horse grazing on grassland may affect the establishment of P. halepensis. We conducted a field experiment to evaluate this hypothesis, studying the response of seedling emergence, survival and growth to herbaceous vegetation defoliation and soil disturbance in sites with contrasting grazing histories. We also evaluated the composition and structure of plant communities of each experimental site. Direct effects such as heavily defoliated perennial grasses and indirect effects such as reduced specific and functional richness, diversity and evenness enhanced the emergence, survival and early growth of the seedlings. High bare ground percentage and low grass biomass also positively affected seedling establishment. The experimental evidence demonstrates that the combination of invasional meltdown hypothesis and weakened biotic resistance are the mechanisms involved in P. halepensis invasion. [Copyright &y& Elsevier]

In Mauritius, many of the worst invasive plant species have fleshy fruits and rely on animals for dispersal. The introduced red-whiskered bulbul ( Pycnonotus jocosus) feeds on many fleshy-fruited species, and often moves from invaded and degraded habitats into higher quality native forests, thus potentially acting as a mediator of continued plant invasion into these areas. Furthermore, gut passage may influence seed germination. To investigate this, we fed fleshy fruits of two invasive plant species, Ligustrum robustum and Clidemia hirta, to red-whiskered bulbuls. Gut passage times of seeds were recorded. Gut-passed seeds were sown and their germination rate and germination success compared with that of hand-cleaned seeds, as well as that of seeds in whole fruits. Gut passage and hand-cleaning had significant positive effects on germination of both species. Gut-passed seeds of both C. hirta and L. robustum germinated faster than hand-cleaned seeds. However, for L. robustum, this was only true when compared with hand-cleaned seeds with intact endocarp; when compared with hand-cleaned seeds without endocarp, there was no difference. For overall germination success, there was a positive effect of gut passage for C. hirta, but not for L. robustum. For both C. hirta and L. robustum, no seeds in intact fruits geminated, suggesting that removal of pulp is essential for germination. Our results suggest that, first, the initial invasion of native forests in Mauritius may not have happened so rapidly without efficient avian seed dispersers like the red-whiskered bulbul. Second, the bulbul is likely to be a major factor in the continued re-invasion of C. hirta and L. robustum into weeded and restored conservation management areas. [ABSTRACT FROM AUTHOR]

Invasive woody species frequently change the composition of the established vegetation and the properties of the soil under their canopies. Accordingly, invasion may well affect regenerative phases of the community, especially at the seed bank level, likely influencing community restoration. Pyracantha angustifolia (Rosaceae) is an invasive shrub in central Argentina that affects woody recruitment, particularly enhancing the recruitment of other exotic woody species. There is though no information regarding its effect on the soil seed bank within the invaded community. The present study was set up to gain further insight into the canopy effects of P. angustifolia. We aimed to assess whether the invasive shrub affects seed bank composition, richness and seed density as compared with the dominant native shrub Condalia montana (Rhamnaceae), and to relate the observed seed bank patterns with those of the established vegetation. We evaluated the composition of the germinable seed bank and the established vegetation under the canopy of 16 shrubs of P. angustifolia, 16 shrubs of C. montana, and in 16 control plots (10 m2) without shrub cover. The floristic composition of the seed bank differed among canopy treatments. However, seed bank richness did not differ significantly. There was an overall high seed density of exotic species throughout the study site, though exotic forbs showed significantly lower seed densities under the invasive shrub. Pyracantha angustifolia would not promote the incorporation of new species into the seed bank of the invaded community but rather favour the establishment of woody species that do not depend on seed banks. The absence of dominant woody species in the seed bank, the dominance of exotic forbs, and the high similarity between established exotic species and those present in the seed bank may surely affect community restoration following the main disturbances events observed in the region. [ABSTRACT FROM AUTHOR]

Questions: 1. Is there any post-dispersal positive effect of the exotic shrub Pyracantha angustifolia on the success of Ligustrum lucidum seedlings, as compared to the effect of the native Condalia montana or the open herbaceous patches between shrubs? 2. Is the possible facilitation by Pyracantha and/or Condalia related to differential emergence, growth, or survival of Ligustrum seedlings under their canopies? Location: Córdoba, central Argentina. Methods: We designed three treatments, in which ten mature individuals of Pyracantha, ten of the dominant native shrub Condalia montana, and ten patches without shrub cover were involved. In each treatment we planted seeds and saplings of Ligustrum collected from nearby natural populations. Seedlings emerging from the planted seeds were harvested after one year to measure growth. Survival of the transplanted saplings was recorded every two month during a year. Half of the planted seeds and transplanted saplings were cage-protected from rodents. Results: Ligustrum seedling emergence did not differ among treatments while growth was significantly higher in the absence of shrub cover. Sapling survival was significantly higher under the canopy of Pyracantha, intermediate under Condalia, and lowest in the absence of shrub cover. Caging did not affect growth but enhanced seedling emergence and sapling survival. Conclusion: The differential sapling survival in the shrub canopy treatments is consistent with natural sapling distribution. Pyracantha and, less so, Condalia, has a nurse-plant effect on Ligustrum. This results from contrasting effects of the shrubs on different stages of the life cycle of Ligustrum: no effect on seedling emergence, negative on seedling growth, and positive on sapling survival. This suggests that efforts to control the expansion of Ligustrum over the landscape should tackle Pyracantha as well. [ABSTRACT FROM AUTHOR]

Summary: This study examined an ‘invasional meltdown’, where the invasion of a Midwestern woodland by an exotic shrub (Rhamnus cathartica L.P. Mill) and the invasion by Eurasian earthworms facilitated one another. Using a litterbag approach, we examined mass loss of four substrates (R. cathartica, Acer saccharum, Quercus rubra, and Quercus alba) along a gradient of Eurasian earthworm density and biomass throughout a 40.5ha oak woodland in Glencoe, Illinois. Earthworm densities and biomass were greatest in patches where R. cathartica prevailed, and populations were lowest in an upland forest subcommunity within the woodland. At each of three points along this earthworm gradient, we placed replicated litterbags constructed either to permit or to deny access to the litter by earthworms. The treatments were, therefore, plot treatments (low, medium and high earthworm density and biomass) and litterbag treatments (earthworm access and earthworm excluded). We found that earthworms promoted a very rapid loss of litter from R. cathartica bags. Within 3 months greater than 90% of this litter was lost from the litterbags. Earthworm impacts on other substrates followed the sequence A. saccharum>Q. alba=Q. rubra. Effects of both litterbag and plot treatments were found within 3 months for A. saccharum but Quercus species were affected only after a year. We propose that the impact of earthworms on litter breakdown creates conditions that promote and sustain invasion by R. cathartica. Previous work has demonstrated that R. cathartica may alter soil properties in a way that promotes and sustains invasion by earthworms. These findings have implications for the restoration management of these systems, since the legacy of R. cathartica on soil properties and earthworm populations may persist even after the plant has been physically removed. [Copyright &y& Elsevier]

This study examined the distribution and habitat selection of the invasive black bullhead [ Ameiurus melas (Rafinesque)] in the ditches and surrounded temporary flooded habitats of an artificial wetland in western France. A multiscale approach was used to quantify patterns of A. melas abundance in relation to physical habitat characteristics in the ditch network. Young-of-the-year (YOY) and adult A. melas largely dominated the local fish assemblage but were highly variable among sites. Although we found evidence for some fine-scale habitat differences for YOY and adult individuals, the abundance of A. melas was positively and consistently related to the dominance of reed beds. Furthermore, A. melas preferentially used reed beds as opposed to marsh meadows during the flooding period. The results from this study suggest that the invasion of A. melas has been facilitated by the expansion of reed beds associated with the diminution of agricultural pressure in recent decades. This study represents an unusual example where human activities can have had an unexpected effect by facilitating an invasive fish species. [ABSTRACT FROM AUTHOR]

Biological invasions are a growing aspect of global biodiversity change. In many regions, introduced species richness increases supralinearly over time. This does not, however, necessarily indicate increasing introduction rates or invasion success. We develop a simple null model to identify the expected trend in invasion records over time. For constant introduction rates and success, the expected trend is exponentially increasing. Model extensions with varying introduction rate and success can also generate exponential distributions. We then analyse temporal trends in aquatic, marine and terrestrial invasion records. Most data sets support an exponential distribution (15/16) and the null invasion model (12/16). Thus, our model shows that no change in introduction rate or success need be invoked to explain the majority of observed trends. Further, an exponential trend does not necessarily indicate increasing invasion success or ‘invasional meltdown’, and a saturating trend does not necessarily indicate decreasing success or biotic resistance. [ABSTRACT FROM AUTHOR]

The probability of a bird species going extinct on oceanic islands in the period since European colonization is predicted by the number of introduced predatory mammal species, but the exact mechanism driving this relationship is unknown. One possibility is that larger exotic predator communities include a wider array of predator functional types. These predator communities may target native bird species with a wider range of behavioral or life history characteristics. We explored the hypothesis that the functional diversity of the exotic predators drives bird species extinctions. We also tested how different combinations of functionally important traits of the predators explain variation in extinction probability. Our results suggest a unique impact of each introduced mammal species on native bird populations, as opposed to a situation where predators exhibit functional redundancy. Further, the impact of each additional predator may be facilitated by those already present, suggesting the possibility of "invasional meltdown." [ABSTRACT FROM AUTHOR]

Generalized mutualisms are often predicted to be resilient to changes in partner identity. Variation in mutualism-related traits between native and invasive species however, can exacerbate the spread of invasive species ('invasional meltdown') if invasive partners strongly interact. Here we show how invasion by a seed-dispersing ant (Myrmica rubra) promotes recruitment of a co-introduced invasive over native ant-dispersed (myrmecochorous) plants. We created experimental communities of invasive (M. rubra) or native ants (Aphaenogaster rudis) and invasive and native plants and measured seed dispersal and plant recruitment. In our mesocosms, and in laboratory and field trials, M. rubra acted as a superior seed disperser relative to the native ant. By contrast, previous studies have found that invasive ants are often poor seed dispersers compared with native ants. Despite belonging to the same behavioural guild, seed-dispersing ants were not functionally redundant. Instead, native and invasive ants had strongly divergent effects on plant communities: the invasive plant dominated in the presence of the invasive ant and the native plants dominated in the presence of the native ant. Community changes were not due to preferences for coevolved partners: variation in functional traits of linked partners drove differences. Here, we show that strongly interacting introduced mutualists can be major drivers of ecological change. [ABSTRACT FROM AUTHOR]