Invading predatory crustacean Dikerogammarus villosus eliminates bath native and exotic species

As the tempo of biological invasions increases, explanations and predictions of their impacts become more crucial. Particularly with regard to biodiversity, we require elucidation of interspecific behavioural interactions among invaders and natives. In freshwaters in The Netherlands, we show that the invasive Ponto-Caspian crustacean amphipod Dikerogammarus villosus is rapidly eliminating Gammarus duebeni, a native European amphipod, and Gammarus tigrinus, until now a spectacularly successful invader from North America. In the laboratory, survival of single (unguarded) female G. duebeni was significantly lower when male D. villosus were free to roam as compared with isolated within microcosms. In addition, survival of paired (guarded) female G. duebeni was significantly lower when male D. villosus as compared with male G. duebeni were present. D. villosus killed and consumed both recently moulted and, unusually, intermoult victims. Survival of G. tigrinus was significantly lower when D. villosus were free to roam as compared with isolated within microcosms and, again, both moulted and intermoult victims were preyed upon. Male D. villosus were significantly more predatory than were females, while female G. tigrinus were significantly more often preyed upon than were males. Predation by D. villosus on both species occurred over a range of water conductivities, an environmental feature previously shown to promote amphipod coexistence. This predatory invader is predicted to reduce further the amphipod diversity in a range of freshwater habitats in Europe and North America.

European hare (Lepus europaeus) invasion ecology: Implication for the conservation of the endemic Irish hare (Lepus timidus hibernicus)

European hare Lepus europaeus populations have undergone recent declines but the species has successfully naturalised in many countries outside its native range. It was introduced to Ireland during the mid-late nineteenth century for field sport and is now well established in Northern Ireland. The native Irish hare Lepus timidus hibernicus is an endemic subspecies of mountain hare L. timidus and has attracted major conservation concern following a long-term population decline during the twentieth century and is one of the highest priority species for conservation action in Ireland. Little is known about the European hare in Ireland or whether it poses a significant threat to the native mountain hare subspecies by compromising its ecological security or genetic integrity. We review the invasion ecology of the European hare and examine evidence for interspecific competition with the mountain hare for habitat space and food resources, interspecific hybridisation, disease and parasite transmission and possible impacts of climate change. We also examine the impact that introduced hares can have on native non-lagomorph species. We conclude that the European hare is an emerging and significant threat to the conservation status of the native Irish hare. Invasive mammal species have been successfully eradicated from Ireland before and immediate action is often the only opportunity for cost-effective eradication. An urgent call is issued for further research whilst the need for a European hare invasive Species Action Plan (iSAP) and Eradication strategy are discussed.

Interactions between invasive and native crustaceans: differential functional responses of intraguild predators towards juvenile hetero-specifics

Intraguild predation (IGP) between invasive and native species can lead to species exclusions or co-existence, dependent on the direction and strength of the interaction. Recently, derivation of 'functional responses' has been identified as a means of comparing the community impacts of invasive and native species. Here, we employ a novel use of this functional response methodology to evaluate any IGP asymmetries between the invasive Ponto-Caspian amphipod Echinogammarus ischnus and the North American native Gammarus fasciatus. The direction and magnitude of intraguild predation of adult males on hetero-specific adult females has previously been shown to reverse across a water conductivity gradient. This partially explains field patterns, but does not predict the co-existence of the two species observed in many habitats and locations. Here, we compared intraguild predation by both species on each other's juveniles in high- and low- conductivity water. G. fasciatus has a higher type II functional response towards E. ischnus juveniles compared to the reciprocal interaction. Conductivity did not influence the predation rate on juveniles of either E. ischnus or G. fasciatus. Thus, the male/female IGP advantage to the native G. fasciatus in low conductivity water is compounded by a juvenile IGP asymmetry, which also counteracts the male/female IGP advantage to E. ischnus in high conductivity waters, helping to explain field patterns of exclusion and co-existence. Thus, complex asymmetries in mutual IGP associated with inherent species differences, environmental modulation, and life-history effects can help us understand and predict the population and community level outcomes of species invasions.

Trophic interactions and consequent impacts of the invasive fish Pseudorasbora parva in a native aquatic foodweb: a field investigation in the UK

Introduction of the invasive Asian cyprinid fish Pseudorasbora parva into a 0.3 ha pond in England with a fish assemblage that included Cyprinus carpio, Rutilus rutilus and Scardinius erythrophthalmus resulted in their establishment of a numerically dominant population in only 2 years; density estimates exceeded 60 ind. m(-2) and they comprised > 99% of fish present. Stable isotope analysis (SIA) revealed significant trophic overlap between P. parva, R. rutilus and C. carpio, a shift associated with significantly depressed somatic growth in R. rutilus. Despite these changes, fish community composition remained similar between the ponds. Comparison with SIA values collected from an adjacent pond free of P. parva revealed a simplified food web in P. parva presence, but with an apparent trophic position shift for several fishes, including S. erythrophthalmus which appeared to assimilate energy at a higher trophic level, probably through P. parva consumption. The marked isotopic shifts shown in all taxa in the P. parva invaded pond (C-13-enriched, N-15 depleted) were indicative of a shift to a cyanobacteria-dominated phytoplankton community. These findings provide an increased understanding of the ecological consequences of the ongoing P. parva invasion of European freshwater ecosystems.

Range expansion in an invasive small mammal: influence of life-history and habitat quality.

Invasive species pose a major threat to biodiversity but provide an opportunity to describe the processes that lead to changes in a species’ range. The bank vole (Myodes glareolus) is an invasive rodent that was introduced to Ireland in the early twentieth century. Given its continuing range expansion, the substantial empirical data on its spread thus far, and the absence of any eradication program, the bank vole in Ireland represents a unique model system for studying the mechanisms influencing the rate of range expansion in invasive small mammals. We described the invasion using a reaction–diffusion model informed by empirical data on life history traits and demographic parameters. We subsequently modelled the processes involved in its range expansion using a rule-based spatially explicit simulation. Habitat suitability interacted with density-dependent parameters to influence dispersal, most notably the density at which local populations started to donate emigrating individuals, the number of dispersing individuals and the direction of dispersal. Whilst local habitat variability influenced the rate of spread, on a larger scale the invasion resembled a simple reaction–diffusion process. Our results suggest a Type 1 range expansion where the rate of expansion is generally constant over time, but with some evidence for a lag period following introduction. We demonstrate that a two-parameter empirical model and a rule-based spatially explicit simulation are sufficient to accurately describe the invasion history of a species that exhibits a complex, density-dependent pattern of dispersal.

Ecological impacts of an invasive predator explained and predicted by comparative functional responses

Forecasting the ecological impacts of invasive species is a major challenge that has seen little progress, yet the development of robust predictive approaches is essential as new invasion threats continue to emerge. A common feature of ecologically damaging invaders is their ability to rapidly exploit and deplete resources. We thus hypothesized that the 'functional response' (the relationship between resource density and consumption rate) of such invasive species might be of consistently greater magnitude than those of taxonomically and/or trophically similar native species. Here, we derived functional responses of the predatory Ponto-Caspian freshwater 'bloody red' shrimp, Hemimysis anomala, a recent and ecologically damaging invader in Europe and N. America, in comparison to the local native analogues Mysis salemaai and Mysis diluviana in Ireland and Canada, respectively. This was conducted in a novel set of experiments involving multiple prey species in each geographic location and a prey species that occurs in both regions. The predatory functional responses of the invader were generally higher than those of the comparator native species and this difference was consistent across invaded regions. Moreover, those prey species characterized by the strongest and potentially de-stabilizing Type II functional responses in our laboratory experiments were the same prey species found to be most impacted by H. anomala in the field. The impact potential of H. anomala was further indicated when it exhibited similar or higher attack rates, consistently lower prey handling times and higher maximum feeding rates compared to those of the two Mysis species, formerly known as 'Mysis relicta', which itself has an extensive history of foodweb disruption in lakes to which it has been introduced. Comparative functional responses thus merit further exploration as a methodology for predicting severe community-level impacts of current and future invasive species and could be entered into risk assessment protocols.

Four new exotic red seaweeds on European shores

We report the occurrence of four red macroalgae new to Europe. Two species were unambiguously determined to the species level with a DNA barcoding approach, while the remaining two species could only be assigned to a genus. Gelidium vagum was found in the Oosterschelde estuary (the Netherlands). Gracilariopsis chorda, Chondracanthus sp. and Solieria sp. were found in the Gulf of Morbihan in Brittany (France); Solieria sp. was also subsequently observed in the Thau Lagoon (France). Gelidium vagum and Gracilariopsis chorda are species originating from the north-western Pacific, around the Japanese archipelago. Phylogenetic analyses also show a likely Pacific origin for Chondracanthus sp. and Solieria sp. Three of these species are likely to have been introduced after 2008, indicating some active transport pathways between the Pacific and the north-eastern Atlantic. These findings also underline the importance of consistent and continuous local expertise (versus rapid assessment) in early warning systems.

Incorporating uncertainty in predictive species distribution modelling

Motivated by the need to solve ecological problems (climate change, habitat fragmentation and biological invasions), there has been increasing interest in species distribution models (SDMs). Predictions from these models inform conservation policy, invasive species management and disease-control measures. However, predictions are subject to uncertainty, the degree and source of which is often unrecognized. Here, we review the SDM literature in the context of uncertainty, focusing on three main classes of SDM: niche-based models, demographic models and process-based models. We identify sources of uncertainty for each class and discuss how uncertainty can be minimized or included in the modelling process to give realistic measures of confidence around predictions. Because this has typically not been performed, we conclude that uncertainty in SDMs has often been underestimated and a false precision assigned to predictions of geographical distribution. We identify areas where development of new statistical tools will improve predictions from distribution models, notably the development of hierarchical models that link different types of distribution model and their attendant uncertainties across spatial scales. Finally, we discuss the need to develop more defensible methods for assessing predictive performance, quantifying model goodness-of-fit and for assessing the significance of model covariates.

Advancing impact prediction and hypothesis testing in invasion ecology using a comparative functional response approach

Invasion ecology urgently requires predictive methodologies that can forecast the ecological impacts of existing, emerging and potential invasive species. We argue that many ecologically damaging invaders are characterised by their more efficient use of resources. Consequently, comparison of the classical ‘functional response’ (relationship between resource use and availability) between invasive and trophically analogous native species may allow prediction of invader ecological impact. We review the utility of species trait comparisons and the history and context of the use of functional responses in invasion ecology, then present our framework for the use of comparative functional responses. We show that functional response analyses, by describing the resource use of species over a range of resource availabilities, avoids many pitfalls of ‘snapshot’ assessments of resource use. Our framework demonstrates how comparisons of invader and native functional responses, within and between Type II and III functional responses, allow testing of the likely population-level outcomes of invasions for affected species. Furthermore, we describe how recent studies support the predictive capacity of this method; for example, the invasive ‘bloody red shrimp’ Hemimysis anomala shows higher Type II functional responses than native mysids and this corroborates, and could have predicted, actual invader impacts in the field. The comparative functional response method can also be used to examine differences in the impact of two or more invaders, two or more populations of the same invader, and the abiotic (e.g. temperature) and biotic (e.g. parasitism) context-dependencies of invader impacts. Our framework may also address the previous lack of rigour in testing major hypotheses in invasion ecology, such as the ‘enemy release’ and ‘biotic resistance’ hypotheses, as our approach explicitly considers demographic consequences for impacted resources, such as native and invasive prey species. We also identify potential challenges in the application of comparative functional responses in invasion ecology. These include incorporation of numerical responses, multiple predator effects and trait-mediated indirect interactions, replacement versus non-replacement study designs and the inclusion of functional responses in risk assessment frameworks. In future, the generation of sufficient case studies for a meta-analysis could test the overall hypothesis that comparative functional responses can indeed predict invasive species impacts.

Fortune favours the bold: a higher predator reduces the impact of a native but not an invasive intermediate predator

Emergent multiple predator effects (MPEs) might radically alter predictions of predatory impact that are based solely on the impact of individuals. In the context of biological invasions, determining if and how the individual-level impacts of invasive predators relates to their impacts in multiple-individual situations will inform understanding of how such impacts might propagate through recipient communities. Here, we use functional responses (the relationship between prey consumption rate and prey density) to compare the impacts of the invasive freshwater mysid crustacean Hemimysis anomala with a native counterpart Mysis salemaai when feeding on basal cladoceran prey (i) as individuals, (ii) in conspecific groups and (iii) in conspecific groups in the presence of a higher fish predator, Gasterosteus aculeatus. In the absence of the higher predator, the invader consumed significantly more basal prey than the native, and consumption was additive for both mysid species - that is, group consumption was predictable from individual-level consumption. Invaders and natives were themselves equally susceptible to predation when feeding with the higher fish predator, but an MPE occurred only between the natives and higher predator, where consumption of basal prey was significantly reduced. In contrast, consumption by the invaders and higher predator remained additive. The presence of a higher predator serves to exacerbate the existing difference in individual-level consumption between invasive and native mysids. We attribute the mechanism responsible for the MPE associated with the native to a trait-mediated indirect interaction, and further suggest that the relative indifference to predator threat on the part of the invader contributes to its success and impacts within invaded communities.

Parasites that change predator or prey behaviour can have keystone effects on community composition

Parasites play pivotal roles in structuring communities, often via indirect interactions with non-host species. These effects can be density-mediated (through mortality) or trait-mediated (behavioural, physiological and developmental), and may be crucial to population interactions, including biological invasions. For instance, parasitism can alter intraguild predation (IGP) between native and invasive crustaceans, reversing invasion outcomes. Here, we use mathematical models to examine how parasite-induced trait changes influence the population dynamics of hosts that interact via IGP. We show that trait-mediated indirect interactions impart keystone effects, promoting or inhibiting host coexistence. Parasites can thus have strong ecological impacts, even if they have negligible virulence, underscoring the need to consider trait-mediated effects when predicting effects of parasites on community structure in general and biological invasions in particular.

Tackling Invasive Alien Species in Europe: the Top 20 Issues

Globally, Invasive Alien Species (IAS) are considered to be one of the major threats to native biodiversity, with the World Conservation Union (IUCN) citing their impacts as ?immense, insidious, and usually irreversible?. It is estimated that 11% of the c. 12,000 alien species in Europe are invasive, causing environmental, economic and social damage; and it is reasonable to expect that the rate of biological invasions into Europe will increase in the coming years. In order to assess the current position regarding IAS in Europe and to determine the issues that were deemed to be most important or critical regarding these damaging species, the international Freshwater Invasives - Networking for Strategy (FINS) conference was convened in Ireland in April 2013. Delegates from throughout Europe and invited speakers from around the world were brought together for the conference. These comprised academics, applied scientists, policy makers, politicians, practitioners and representative stakeholder groups. A horizon scanning and issue prioritization approach was used by in excess of 100 expert delegates in a workshop setting to elucidate the Top 20 IAS issues in Europe. These issues do not focus solely on freshwater habitats and taxa but relate also to marine and terrestrial situations. The Top 20 issues that resulted represent a tool for IAS management and should also be used to support policy makers as they prepare European IAS legislation.

Habitat simplification increases the impact of a freshwater invasive fish

Biodiversity continues to decline at a range of spatial scales and there is an urgent requirement to understand how multiple drivers interact in causing such declines. Further, we require methodologies that can facilitate predictions of the effects of such drivers in the future. Habitat degradation and biological invasions are two of the most important threats to biodiversity and here we investigate their combined effects, both in terms of understanding and predicting impacts on native species. The predatory largemouth bass Micropterus salmoides is one of the World’s Worst Invaders, causing declines in native prey species, and its introduction often coincides with habitat simplification. We investigated the predatory functional response, as a measure of ecological impact, of juvenile largemouth bass in artificial vegetation over a range of habitat complexities (high, intermediate, low and zero). Prey, the female guppy Poecilia reticulata, were representative of native fish. As habitats became less complex, significantly more prey were consumed, since, even although attack rates declined, reduced handling times resulted in higher maximum feeding rates by bass. At all levels of habitat complexity, bass exhibited potentially population destabilising Type II functional responses, with no emergence of more stabilising Type III functional responses as often occurs in predator-prey relationships in complex habitats. Thus, habitat degradation and simplification potentially exacerbate the impact of this invasive species, but even highly complex habitats may ultimately not protect native species. The utilisation of functional responses under varying environmental contexts provides a method for the understanding and prediction of invasive species impacts.