Community‐level effects of herbicide‐based restoration treatments: structural benefits but at what cost?

Invasive species alter ecosystem structure, impact biodiversity, and have significant economic costs. In Oregon's Willamette Valley, invasive grasses Arrhenatherum elatius and Schedonorus arundinaceus alter the dynamics of the phenologically paired interaction between an endangered butterfly, Icaricia icarioides fenderi (Fender's blue), and its larval host plant, Lupinus oreganus (Kincaid's lupine). To test methods to restore this interaction, we established a 3‐year experiment where a post‐emergent grass‐specific herbicide, fluazifop‐p‐butyl, was applied to Fender's blue habitat. Plant community data were recorded throughout the growing season at eight paired plots for 1 year prior to treatment and 3 years during treatment. We asked whether annual application of herbicide could reduce the height of invasive grasses to levels at or beneath the height of Kincaid's lupine racemes throughout the Fender's blue flight season. We hypothesized that native forb species, which are critical nectar sources for Fender's blue, would increase in cover and frequency following the release from competitive dominance of invasive grasses. Grass‐specific herbicide reduced grass height during the flight season of Fender's blue, but with several costs. We found no change in nectar and a suppression of lupine growth in plots in response to experimental herbicide treatment. Each study site had multiple secondary invaders; the long‐term impact of these new invaders is unknown. We suggest that herbicide application results in a net negative effect in the context of Fender's blue habitat restoration. That is, the costs to primary resources for Fender's blue and the influx of secondary invaders may be as problematic as the primary invasion by non‐native grasses.     

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.     

A meta-analysis of biotic resistance to exotic plant invasions

Biotic resistance describes the ability of resident species in a community to reduce the success of exotic invasions. Although resistance is a well‐accepted phenomenon, less clear are the processes that contribute most to it, and whether those processes are strong enough to completely repel invaders. Current perceptions of strong, competition‐driven biotic resistance stem from classic ecological theory, Elton's formulation of ecological resistance, and the general acceptance of the enemies‐release hypothesis. We conducted a meta‐analysis of the plant invasions literature to quantify the contribution of resident competitors, diversity, herbivores and soil fungal communities to biotic resistance. Results indicated large negative effects of all factors except fungal communities on invader establishment and performance. Contrary to predictions derived from the natural enemies hypothesis, resident herbivores reduced invasion success as effectively as resident competitors. Although biotic resistance significantly reduced the establishment of individual invaders, we found little evidence that species interactions completely repelled invasions. We conclude that ecological interactions rarely enable communities to resist invasion, but instead constrain the abundance of invasive species once they have successfully established.     

Existing and emerging high impact invasive species are characterized by higher functional responses than natives

Predicting ecological impacts of invasive species and identifying potentially damaging future invaders are research priorities. Since damage by invaders is characterized by their depletion of resources, comparisons of the ‘functional response’ (FR; resource uptake rate as a function of resource density) of invaders and natives might predict invader impact. We tested this by comparing FRs of the ecologically damaging ‘world''s worst’ invasive fish, the largemouth bass (Micropterus salmoides), with a native equivalent, the Cape kurper (Sandelia capensis), and an emerging invader, the sharptooth catfish (Clarias gariepinus), with the native river goby (Glossogobius callidus), in South Africa, a global invasion hotspot. Using tadpoles (Hyperolius marmoratus) as prey, we found that the invaders consumed significantly more than natives. Attack rates at low prey densities within invader/native comparisons reflected similarities in predatory strategies; however, both invasive species displayed significantly higher Type II FRs than the native comparators. This was driven by significantly lower prey handling times by invaders, resulting in significantly higher maximum feeding rates. The higher FRs of these invaders are thus congruent with, and can predict, their impacts on native communities. Comparative FRs may be a rapid and reliable method for predicting ecological impacts of emerging and future invasive species.     

Protected areas offer refuge from invasive species spreading under climate change

Protected areas (PAs) are intended to provide native biodiversity and habitats with a refuge against the impacts of global change, particularly acting as natural filters against biological invasions. In practice, however, it is unknown how effective PAs will be in shielding native species from invasions under projected climate change. Here, we investigate the current and future potential distributions of 100 of the most invasive terrestrial, freshwater, and marine species in Europe. We use this information to evaluate the combined threat posed by climate change and invasions to existing PAs and the most susceptible species they shelter. We found that only a quarter of Europe's marine and terrestrial areas protected over the last 100 years have been colonized by any of the invaders investigated, despite offering climatically suitable conditions for invasion. In addition, hotspots of invasive species and the most susceptible native species to their establishment do not match at large continental scales. Furthermore, the predicted richness of invaders is 11%–18% significantly lower inside PAs than outside them. Invasive species are rare in long‐established national parks and nature reserves, which are actively protected and often located in remote and pristine regions with very low human density. In contrast, the richness of invasive species is high in the more recently designated Natura 2000 sites, which are subject to high human accessibility. This situation may change in the future, since our models anticipate important shifts in species ranges toward the north and east of Europe at unprecedented rates of 14–55 km/decade, depending on taxonomic group and scenario. This may seriously compromise the conservation of biodiversity and ecosystem services. This study is the first comprehensive assessment of the resistance that PAs provide against biological invasions and climate change on a continental scale and illustrates their strategic value in safeguarding native biodiversity.     

Using avatar species to model the potential distribution of emerging invaders

Aim Anticipating the potential distributions of emerging invasive species is complicated by the tendency for species distribution models to perform better when both native and invasive range data are available for model development. If invasive range data are lacking, species models are liable to under‐estimate distributions for emerging invaders, particularly for species that are not at equilibrium with their native range environment due to historical factors, dispersal limitation and/or ecological interactions. We demonstrate the potential to use well‐quantified niche shifts from established ‘avatar’ (i.e. the remote or virtual manifestation of an entity) invaders to develop plausible distributions for data‐poor emerging invaders contingent on niche shifts of similar magnitude or character. Location Global. Methods Using the globally invasive crayfishes Pacifastacus leniusculus and Procambarus clarkii as our avatar invaders, we quantify how niche position, size and structure differs between native and total ranges using Mahalanobis distance (a measure of multivariate similarity) and the climate predictors of annual minimum and maximum air temperature. We then generalize patterns of niche shift from these species to the emerging crayfish invader Cherax quadricarinatus. Results Some patterns of niche shifts were similar for Pacifastacus leniusculus and Procambarus clarkii, but niche shifts were of considerably greater magnitude for P. clarkii. When a native range model for C. quadricarinatus was modified with generalized niche shifts similar to Pacifastacus leniusculus and Procambarus clarkii, the potential global distribution for this species increased considerably, including many areas not identified by the native range model. Main conclusions We illustrate the potential to use avatar invaders to provide cautionary, niche shift‐assuming species distribution models for emerging invaders. Many theoretical and applied implications of the avatar species concept require additional investigation, including the development of frameworks to select appropriate avatar species and evaluate the performance of avatar‐derived models for emerging invaders. Despite these research needs, we believe this concept will have considerable utility for predicting vulnerability to invasion by data‐poor species; this is a critical management need because shifting pathways of introduction and climate change will produce many novel, emerging invasive species in the future.     

Experimental removals reveal dietary niche partitioning facilitates coexistence between native and introduced species

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External morphology explains the success of biological invasions

Biological invasions have become major players in the current biodiversity crisis, but realistic tools to predict which species will establish successful populations are still unavailable. Here we present a novel approach that requires only a morphometric characterisation of the species. Using fish invasions of the Mediterranean, we show that the abundance of non‐indigenous fishes correlates with the location and relative size of occupied morphological space within the receiving pool of species. Those invaders that established abundant populations tended to be added outside or at the margins of the receiving morphospace, whereas non‐indigenous species morphologically similar to resident ones failed to develop large populations or even to establish themselves, probably because the available ecological niches were already occupied. Accepting that morphology is a proxy for a species' ecological position in a community, our findings are consistent with ideas advanced since Darwin's naturalisation hypothesis and provide a new warning signal to identify invaders and to recognise vulnerable communities.     

Niche shift can impair the ability to predict invasion risk in the marine realm: an illustration using Mediterranean fish invaders

Climatic niche conservatism, the tendency of species‐climate associations to remain unchanged across space and time, is pivotal for forecasting the spread of invasive species and biodiversity changes. Indeed, it represents one of the key assumptions underlying species distribution models (SDMs), the main tool currently available for predicting range shifts of species. However, to date, no comprehensive assessment of niche conservatism is available for the marine realm. We use the invasion by Indo‐Pacific tropical fishes into the Mediterranean Sea, the world's most invaded marine basin, to examine the conservatism of the climatic niche. We show that tropical invaders may spread far beyond their native niches and that SDMs do not predict their new distributions better than null models. Our results suggest that SDMs may underestimate the potential spread of invasive species and call for prudence in employing these models in order to forecast species invasion and their response to environmental change.     

Elevated success of multispecies bacterial invasions impacts community composition during ecological succession

Successful microbial invasions are determined by a species’ ability to occupy a niche in the new habitat whilst resisting competitive exclusion by the resident community. Despite the recognised importance of biotic factors in determining the invasiveness of microbial communities, the success and impact of multiple concurrent invaders on the resident community has not been examined. Simultaneous invasions might have synergistic effects, for example if resident species need to exhibit divergent phenotypes to compete with the invasive populations. We used three phylogenetically diverse bacterial species to invade two compositionally distinct communities in a controlled, naturalised in vitro system. By initiating the invader introductions at different stages of succession, we could disentangle the relative importance of resident community structure, invader diversity and time pre‐invasion. Our results indicate that multiple invaders increase overall invasion success, but do not alter the successional trajectory of the whole community.     

Carbohydrate scarcity increases foraging activities and aggressiveness in the ant Prolasius advenus (Hymenoptera: Formicidae)

1. Behavioural responses to varying macronutrient availability are increasingly studied in highly invasive ant species to better understand their ecological success. However, such work is lacking in relation to native ant species confronted with biological invaders. 2. Here the link between diet and behaviour was examined in Prolasius advenus, a native ant from New Zealand facing intense competition for food with invasive insects, including social wasps. A long‐term laboratory experiment was conducted to assess the impact of protein and carbohydrate restriction on several behavioural parameters. 3. Ants shifted their food‐collecting activity towards the least available macronutrient in both protein‐limited and carbohydrate‐limited colonies. But when lacking carbohydrate, they also strongly increased their efforts to discover resources. After 10 weeks, the proportion of the colony patrolling outside of the nest was eight times higher than at the initiation of the experiment. This high patrolling activity was then maintained for several weeks and resulted in a higher efficiency to explore a novel territory. Moreover, ants fed a carbohydrate‐limited diet engaged in longer aggressive acts towards conspecifics. 4. These behavioural responses to carbohydrate scarcity may, in part, enhance the ability of P. advenus to resist a competing invader under natural conditions. While much of the previous research has linked diet and behavioural dominance in invasive ants, the present study shows that conducting similar investigations in the native species to which they are confronted may shed light on the mechanisms behind biotic resistance and the ability of some native species to coexist with highly abundant invaders.     

Predicting the impacts of an introduced species from its invasion history: an empirical approach applied to zebra mussel invasions

1. Quantitative models of impact are lacking for the vast majority of known invasive species, particularly in aquatic ecosystems. Consequently, managers lack predictive tools to help them prioritise invasion threats and decide where they can most effectively allocate limited resources. Predictive tools would also enhance the accuracy of water quality assessments, so that impacts caused by an invader are not erroneously attributed to other anthropogenic stressors. 2. The invasion history of a species is a valuable guide for predicting the consequences of its introduction into a new environment. Regression analysis of data from multiple invaded sites can generate empirical models of impact, as is shown here for the zebra mussel Dreissena polymorpha. Dreissena's impacts on benthic invertebrate abundance and diversity follow predictable patterns that are robust across a range of habitat types and geographic regions. Similar empirical models could be developed for other invaders with a documented invasion history. 3. Because an invader's impact is correlated with its abundance, a surrogate model may be generated (when impact data are unavailable) by relating the invader's abundance to environmental variables. Such a model could help anticipate which habitats will be most affected by invasion. Lack of precision should not be a deterrent to developing predictive models where none exist. Crude predictions can be refined as additional data become available. Empirical modelling is a highly informative and inexpensive, but underused, approach in the management of aquatic invasive species.     

Effects of biological invasions on forest carbon sequestration

There has been a rapidly developing literature on the effects of some of the major drivers of global change on carbon (C) sequestration, particularly carbon dioxide (CO₂) enrichment, land use change, nitrogen (N) deposition and climate change. However, remarkably little attention has been given to one major global change driver, namely biological invasions. This is despite growing evidence that invasive species can dramatically alter a range of aboveground and belowground ecosystem processes, including those that affect C sequestration. In this review, we assess the evidence for the impacts of biological invaders on forest C stocks and C sequestration by biological invaders. We first present case studies that highlight a range of invader impacts on C sequestration in forest ecosystems, and draw on examples that involve invasive primary producers, decomposers, herbivores, plant pathogens, mutualists and predators. We then develop a conceptual framework for assessing the effects of invasive species on C sequestration impacts more generally, by identifying the features of biological invaders and invaded ecosystems that are thought to most strongly regulate C in forests. Finally we assess the implications of managing invasive species on C sequestration. An important principle that emerges from this review is that the direct effects of invaders on forest C are often smaller and shorter‐term than their indirect effects caused by altered nutrient availability, primary productivity or species composition, all of which regulate long‐term C pools and fluxes. This review provides a conceptual basis for improving our general understanding of biological invaders on ecosystem C, but also points to a paucity of primary data that are needed to determine the quantitative effects of invaders on ecosystem processes that drive C sequestration.     

Evaluating the “recovery level” of endangered species without prior information before alien invasion

For maintaining social and financial support for eradication programs of invasive species, quantitative assessment of recovery of native species or ecosystems is important because it provides a measurable parameter of success. However, setting a concrete goal for recovery is often difficult owing to lack of information prior to the introduction of invaders. Here, we present a novel approach to evaluate the achievement level of invasive predator management based on the carrying capacity of endangered species estimated using long‐term monitoring data. In Amami‐Oshima Island, Japan, where the eradication project of introduced small Indian mongoose is ongoing since 2000, we surveyed the population densities of four endangered species threatened by the mongoose (Amami rabbit, the Otton frog, Amami tip‐nosed frog, and Amami Ishikawa's frog) at four time points ranging from 2003 to 2011. We estimated the carrying capacities of these species using the logistic growth model combined with the effects of mongoose predation and environmental heterogeneity. All species showed clear tendencies toward increasing their density in line with decreased mongoose density, and they exhibited density‐dependent population growth. The estimated carrying capacities of three endangered species had small confidence intervals enough to measure recovery levels by the mongoose management. The population density of each endangered species has recovered to the level of the carrying capacity at about 20–40% of all sites, whereas no individuals were observed at more than 25% of all sites. We propose that the present approach involving appropriate monitoring data of native organism populations will be widely applicable to various eradication projects and provide unambiguous goals for management of invasive species.     

Weed risk assessment for aquatic plants: modification of a New Zealand system for the United States

We tested the accuracy of an invasive aquatic plant risk assessment system in the United States that we modified from a system originally developed by New Zealand's Biosecurity Program. The US system is comprised of 38 questions that address biological, historical, and environmental tolerance traits. Values associated with each response are summed to produce a total score for each species that indicates its risk of invasion. To calibrate and test this risk assessment, we identified 39 aquatic plant species that are major invaders in the continental US, 31 species that have naturalized but have no documented impacts (minor invaders), and 60 that have been introduced but have not established. These species represent 55 families and span all aquatic plant growth forms. We found sufficient information to assess all but three of these species. When the results are compared to the known invasiveness of the species, major invaders are distinguished from minor and non-invaders with 91% accuracy. Using this approach, the US aquatic weed risk assessment correctly identifies major invaders 85%, and non-invaders 98%, of the time. Model validation using an additional 10 non-invaders and 10 invaders resulted in 100% accuracy for the former, and 80% accuracy for the latter group. Accuracy was further improved to an average of 91% for all groups when the 17% of species with scores of 31-39 required further evaluation prior to risk classification. The high accuracy with which we can distinguish non-invaders from harmful invaders suggests that this tool provides a feasible, pro-active system for pre-import screening of aquatic plants in the US, and may have additional utility for prioritizing management efforts of established species.     

Species pool size and invasibility of island communities: a null model of sampling effects

The success of alien species on oceanic islands is considered to be one of the classic observed patterns in ecology. Explanations for this pattern are based on lower species richness on islands and the lower resistance of species‐poor communities to invaders, but this argument needs re‐examination. The important difference between islands and mainland is in the size of species pools, not in local species richness; invasibility of islands should therefore be addressed in terms of differences in species pools. Here I examine whether differences in species pools can affect invasibility in a lottery model with pools of identical native and exotic species. While in a neutral model with all species identical, invasibility does not depend on the species pool, a model with non‐zero variation in population growth rates predicts higher invasibility of communities of smaller pools. This is because of species sampling; drawing species from larger pools increases the probability that an assemblage will include fast growing species. Such assemblages are more likely to exclude random invaders. This constitutes a mechanism through which smaller species pools (such as those of isolated islands) can directly underlie differences in invasibility.     

A meta‐analysis of declines in local species richness from human disturbances

There is high uncertainty surrounding the magnitude of current and future biodiversity loss that is occurring due to human disturbances. Here, we present a global meta‐analysis of experimental and observational studies that report 327 measures of change in species richness between disturbed and undisturbed habitats across both terrestrial and aquatic biomes. On average, human‐mediated disturbances lead to an 18.3% decline in species richness. Declines in species richness were highest for endotherms (33.2%), followed by producers (25.1%), and ectotherms (10.5%). Land‐use change and species invasions had the largest impact on species richness resulting in a 24.8% and 23.7% decline, respectively, followed by habitat loss (14%), nutrient addition (8.2%), and increases in temperature (3.6%). Across all disturbances, declines in species richness were greater for terrestrial biomes (22.4%) than aquatic biomes (5.9%). In the tropics, habitat loss and land‐use change had the largest impact on species richness, whereas in the boreal forest and Northern temperate forests, species invasions had the largest impact on species richness. Along with revealing trends in changes in species richness for different disturbances, biomes, and taxa, our results also identify critical knowledge gaps for predicting the effects of human disturbance on Earth's biomes.     

Habitat niche breadth predicts invasiveness in solitary ascidians

A major focus of invasion biology is understanding the traits associated with introduction success. Most studies assess these traits in the invaded region, while only few compare nonindigenous species to the pool of potential invaders in their native region. We focused on the niche breadth hypothesis, commonly evoked but seldom tested, which states that generalist species are more likely to become introduced as they are capable of thriving under a wide set of conditions. Based on the massive introduction of tropical species into the Mediterranean via the Suez Canal (Lessepsian migration), we defined ascidians in the Red Sea as the pool of potential invaders. We constructed unique settlement plates, each representing six different niches, to assess ascidian niche breadth, and deployed them in similar habitats in the native and invaded regions. For each species found on plates, we evaluated its abundance, relative abundance across successional stages, and niche breadth, and then compared (1) species in the Red Sea known to have been introduced into the Mediterranean (Lessepsian species) and those not known from the Mediterranean (non‐Lessepsian); and (2) nonindigenous and indigenous species in the Mediterranean. Lessepsian species identified on plates in the Red Sea demonstrated wider niche breadth than non‐Lessepsian species, supporting the niche breadth hypothesis within the native region. No differences were found between Lessepsian and non‐Lessepsian species in species abundance and successional stages. In the Mediterranean, nonindigenous species numerically dominated the settlement plates. This precluded robust comparisons of niche breadth between nonindigenous and indigenous species in the invaded region. In conclusion, using Red Sea ascidians as the pool of potential invaders, we found clear evidence supporting the niche breadth hypothesis in the native region. We suggest that such patterns may often be obscured when conducting trait‐based studies in the invaded regions alone. Our findings indicate that quantifying the niche breadth of species in their native regions will improve estimates of invasiveness potential.     

It takes one to know one: Similarity to resident alien species increases establishment success of new invaders

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.

     

The times are changing: temporal shifts in patterns of fish invasions in central European fresh waters

This study examines the invasion history of alien fish species based on exhaustive national data sets on fish invasions of two contiguous central European countries (Germany and Austria). Fifteen alien fish species are currently established in both countries, constituting 14 and 17% of the total freshwater fish fauna of Germany and Austria, respectively. In both countries, six alien species are present, but not established. The status of five alien species in Germany and three species in Austria remains unknown. Accumulation rates of alien fish species have increased in recent decades with >50% of them reported after 1971. North America and Asia were the primary sources of alien fish species in Germany and Austria up to the 1980s, whereas European species of Ponto‐Caspian origin dominate now. Fisheries (including aquaculture) and the animal trade were responsible for most earlier introductions, whereas waterways were the main pathway for recent invaders. The extent of the spatial distribution of alien species was positively correlated with residence time, i.e. the time elapsed since the first national record. Different thermal preferences of early invaders (mostly coldwater species) and new invaders (typically warmwater adapted) may benefit the latter in the face of climate change. It is concluded that new challenges for alien fish management arise and that ecosystem‐based approaches as endorsed by the E.U. Water Framework Directive (maintaining or restoring good ecological status of rivers and streams) should become the centrepiece of river management in Europe.