Ecological Impacts of Alien Species: Quantification, Scope, Caveats, and Recommendations

Despite intensive research during the past decade on the effects of alien species, invasion science still lacks the capacity to accurately predict the impacts of those species and, therefore, to provide timely advice to managers on where limited resources should be allocated. This capacity has been limited partly by the context-dependent nature of ecological impacts, research highly skewed toward certain taxa and habitat types, and the lack of standardized methods for detecting and quantifying impacts. We review different strategies, including specific experimental and observational approaches, for detecting and quantifying the ecological impacts of alien species. These include a four-way experimental plot design for comparing impact studies of different organisms. Furthermore, we identify hypothesis-driven parameters that should be measured at invaded sites to maximize insights into the nature of the impact. We also present strategies for recognizing high-impact species. Our recommendations provide a foundation for developing systematic quantitative measurements to allow comparisons of impacts across alien species, sites, and time.

Predation in the marine intertidal amphipod Echinogammarus marinus Leach: implications of inter- and intra-individual variation

Studies of competition, predator–prey dynamics and food webs typically consider conspecifics as equal, however, individuals from the same population that are seemingly identical can show considerable variation with regards to a number of processes. Such phenomena may be demonstrated in terms of diet, and the quantities and types of resources that are consumed are commonly considered. The marine amphipod Echinogammarus marinus, a recently demonstrated predator on intertidal rocky shores, has been shown to consume a wide range of food types but it is unknown how this may vary between individuals. Here, we investigated the variation that occurs both among and within individuals of a population of E. marinus with respect to the mean numbers consumed of a common prey item, the isopod Jaera nordmanni. First, by comparing the length of starvation times, used as a proxy for hunger level, individuals maintained without food for up to 24 h consumed significantly less prey during feeding trials than those starved for 48 h and longer. The degree of inter-individual variation within each starvation period was also found to differ, with greater variation among individuals starved for shorter periods of time than those starved for longer time periods. Secondly, we tested whether individual amphipods tracked over time consumed consistently similar numbers of prey or whether they showed intra-individual variation, and if so, to what degree. We found that the numbers of prey consumed per individual could be predicted in the short-term between consecutive feeding trials, however over the long-term this relationship broke down. These results are discussed with respect to potential physiological and behavioural mechanisms, as well as the implications that such variation may have for stability of prey populations in the field.

Stressor intensity determines antagonistic interactions between species invasion and multiple stressor effects on ecosystem functioning

Biological invasions, nutrient enrichment and ocean warming are known to threaten biodiversity and ecosystem functioning. The independent effects of these ecological stressors are well studied, however, we lack understanding of their cumulative effects, which may be additive, antagonistic or synergistic. For example, the impacts of biological invasions are often determined by environmental context, which suggests that the effects of invasive species may vary with other stressors such as pollution or climate change. This study examined the effects of an invasive seaweed (Sargassum muticum) on the structure and functioning of a benthic marine assemblage and tested explicitly whether these effects varied with nutrient enrichment and ocean warming. Overall, the presence of Sargassum muticum increased assemblage productivity rates and warming altered algal assemblage structure, which was characterised by a decrease in kelp and an increase in ephemeral green algae. The effects of Sargassum muticum on total algal biomass accumulation, however, varied with nutrient enrichment and warming producing antagonistic cumulative effects on total algal biomass accumulation. These findings show that the nature of stressor interactions may vary with stressor intensity and among response variables, which leads to less predictable consequences for the structure and functioning of communities.

A genome-wide association study of anorexia nervosa

Anorexia nervosa (AN) is a complex and heritable eating disorder characterized by dangerously low body weight. Neither candidate gene studies nor an initial genome-wide association study (GWAS) have yielded significant and replicated results. We performed a GWAS in 2907 cases with AN from 14 countries (15 sites) and 14 860 ancestrally matched controls as part of the Genetic Consortium for AN (GCAN) and the Wellcome Trust Case Control Consortium 3 (WTCCC3). Individual association analyses were conducted in each stratum and meta-analyzed across all 15 discovery data sets. Seventy-six (72 independent) single nucleotide polymorphisms were taken forward for in silico (two data sets) or de novo (13 data sets) replication genotyping in 2677 independent AN cases and 8629 European ancestry controls along with 458 AN cases and 421 controls from Japan. The final global meta-analysis across discovery and replication data sets comprised 5551 AN cases and 21 080 controls. AN subtype analyses (1606 AN restricting; 1445 AN binge-purge) were performed. No findings reached genome-wide significance. Two intronic variants were suggestively associated: rs9839776 (P=3.01 × 10(-7)) in SOX2OT and rs17030795 (P=5.84 × 10(-6)) in PPP3CA. Two additional signals were specific to Europeans: rs1523921 (P=5.76 × 10(-)(6)) between CUL3 and FAM124B and rs1886797 (P=8.05 × 10(-)(6)) near SPATA13. Comparing discovery with replication results, 76% of the effects were in the same direction, an observation highly unlikely to be due to chance (P=4 × 10(^-6)), strongly suggesting that true findings exist but our sample, the largest yet reported, was underpowered for their detection. The accrual of large genotyped AN case-control samples should be an immediate priority for the field.

Ultrahigh Brilliance Multi-MeV γ-Ray Beams from Nonlinear Relativistic Thomson Scattering

We report on the generation of a narrow divergence (θγ<2.5mrad), multi-MeV (Emax≈18MeV) and ultrahigh peak brilliance (>1.8×1020photonss-1mm-2mrad-2 0.1% BW) γ-ray beam from the scattering of an ultrarelativistic laser-wakefield accelerated electron beam in the field of a relativistically intense laser (dimensionless amplitude a0≈2). The spectrum of the generated γ-ray beam is measured, with MeV resolution, seamlessly from 6 to 18 MeV, giving clear evidence of the onset of nonlinear relativistic Thomson scattering. To the best of our knowledge, this photon source has the highest peak brilliance in the multi-MeV regime ever reported in the literature.

Particle-in-cell simulation study of the interaction between a relativistically moving leptonic micro-cloud and ambient electrons

Context. The jets of compact accreting objects are composed of electrons and a mixture of positrons and ions. These outflows impinge on the interstellar or intergalactic medium and both plasmas interact via collisionless processes. Filamentation (beam-Weibel) instabilities give rise to the growth of strong electromagnetic fields. These fields thermalize the interpenetrating plasmas. 

Aims. Hitherto, the effects imposed by a spatial non-uniformity on filamentation instabilities have remained unexplored. We examine the interaction between spatially uniform background electrons and a minuscule cloud of electrons and positrons. The cloud size is comparable to that created in recent laboratory experiments and such clouds may exist close to internal and external shocks of leptonic jets. The purpose of our study is to determine the prevalent instabilities, their ability to generate electromagnetic fields and the mechanism, by which the lepton micro-cloud transfers energy to the background plasma. 

Methods. A square micro-cloud of equally dense electrons and positrons impinges in our particle-in-cell (PIC) simulation on a spatially uniform plasma at rest. The latter consists of electrons with a temperature of 1 keV and immobile ions. The initially charge- and current neutral micro-cloud has a temperature of 100 keV and a side length of 2.5 plasma skin depths of the micro-cloud. The side length is given in the reference frame of the background plasma. The mean speed of the micro-cloud corresponds to a relativistic factor of 15, which is relevant for laboratory experiments and for relativistic astrophysical outflows. The spatial distributions of the leptons and of the electromagnetic fields are examined at several times. 

Results. A filamentation instability develops between the magnetic field carried by the micro-cloud and the background electrons. The electromagnetic fields, which grow from noise levels, redistribute the electrons and positrons within the cloud, which boosts the peak magnetic field amplitude. The current density and the moduli of the electromagnetic fields grow aperiodically in time and steadily along the direction that is anti-parallel to the cloud's velocity vector. The micro-cloud remains conjoined during the simulation. The instability induces an electrostatic wakefield in the background plasma. 

Conclusions. Relativistic clouds of leptons can generate and amplify magnetic fields even if they have a microscopic size, which implies that the underlying processes can be studied in the laboratory. The interaction of the localized magnetic field and high-energy leptons will give rise to synchrotron jitter radiation. The wakefield in the background plasma dissipates the kinetic energy of the lepton cloud. Even the fastest lepton micro-clouds can be slowed down by this collisionless mechanism. Moderately fast charge- and current neutralized lepton micro-clouds will deposit their energy close to relativistic shocks and hence they do not constitute an energy loss mechanism for the shock.

A spatio-temporal contrast of the predatory impact of an invasive freshwater crustacean

Aim: Impacts of invasive species may vary across invasion gradients, owing to trait-based sorting of individuals through dispersal: those aggregating at invasion fronts may be more aggressive and voracious. We examine, in the field and laboratory, variation in the predatory impacts of an invasive Ponto-Caspian crustacean Hemimysis anomala G.O. Sars, 1907 at two sites along a spatio-temporal gradient of invasion.

Location: Republic of Ireland.

Methods: We used reciprocal transplant field-deployed mesocosms to compare predation rates of invasion front and well-established H. anomala on natural zooplankton assemblages. In the laboratory, we measured the functional response (relationship between predation rate and prey supply) of H. anomala from both sites, for a per capita mechanistic comparison of predation efficiency. We also assessed prey selectivity of H. anomala in the mesocosm experiments to further compare feeding behaviour. Finally, we used a correlative approach to assess the community impact of H. anomala across sites, including a nearby uninvaded site, by comparing zooplankton diversities and densities.

Results: Invasion front H. anomala had higher predation rates than well-established H. anomala at high in situ zooplankton densities. Invasion front H. anomala also had higher functional responses - in particular showing higher 'attack rates' - indicating a heightened ability to locate and capture prey. Prey selectivity was consistent across the spatio-temporal contrast, with positive selection for cladocerans. Zooplankton diversity and density declined with time since H. anomala invasion, both being maximal at the uninvaded site.

Main conclusions: Our study, for the first time, (1) reveals differences in predatory per capita effects and associated behavioural traits between two sites along a spatio-temporal invasion gradient and (2) shows a negative community-level impact of the invasive H. anomala in natural water bodies. Further spatio-temporal comparisons of predatory per capita effects of invaders are needed to assess the generality of these results.

The paradox of invasion: Reeves’ muntjac deer invade the British Isles from a limited number of founding females

High levels of genetic diversity and high propagule pressure are favoured by conservation biologists as the basis for successful reintroductions and ensuring the persistence of populations. However, invasion ecologists recognize the ‘paradox of invasion’, as successful species introductions may often be characterized by limited numbers of individuals and associated genetic bottlenecks. In the present study, we used a combination of high-resolution nuclear and mitochondrial genetic markers to investigate the invasion history of Reeves' muntjac deer in the British Isles. This invasion has caused severe economic and ecological damage, with secondary spread currently a concern throughout Europe and potentially globally. Microsatellite analysis based on eight loci grouped all 176 introduced individuals studied from across the species' range in the UK into one genetic cluster, and seven mitochondrial D-loop haplotypes were recovered, two of which were present at very low frequency and were related to more common haplotypes. Our results indicate that the entire invasion can be traced to a single founding event involving a low number of females. These findings highlight the fact that even small releases of species may, if ignored, result in irreversible and costly invasion, regardless of initial genetic diversity or continual genetic influx.

Alien aquatics in Europe: assessing the relative environmental and socio-economic impacts of invasive aquatic macroinvertebrates and other taxa

Invasive alien aquatic species, including marine and freshwater macroinvertebrates, have become increasingly important in terms of both environmental and socio-economic impacts. In order to assess their environmental and economic costs, we applied the Generic Impact Scoring System (GISS) and performed a comparison with other taxa of invaders in Europe. Impacts were scored into six environmental and six socio-economic categories, with each category containing five impact levels. Among 49 aquatic macroinvertebrates, the most impacting species were the Chinese mitten crab, Eriocheir sinensis (Milne-Edwards, 1853) and the zebra mussel, Dreissena polymorpha (Pallas, 1771). The highest impacts found per GISS impact category were, separately; on ecosystems, through predation, as competitors, and on animal production. Eleven species have an impact score > 10 (high impact) and seven reach impact level 5 in at least one impact category (EU blacklist candidates), the maximum score that can be given is 60 impact points. Comparisons were drawn between aquatic macroinvertebrates and vertebrate invaders such as fish, mammals and birds, as well as terrestrial arthropods, revealing invasive freshwater macroinvertebrates to be voracious predators of native prey and damaging to native ecosystems compared with other taxa. GISS can be used to compare these taxa and will aid policy making and targeting of invasive species for management by relevant agencies, or to assist in producing species blacklist candidates.

Interspecific comparison of estrogen and testosterone concentrations in three species of amphipods (Gammarus duebeni celticus, G. pseudolimnaeus, and G. pulex)

The presence and biological significance of vertebrate-related steroid sex hormones in aquatic invertebrates are poorly understood. We compared the concentrations of estrogen (17β-estradiol) and testosterone between amplexing male and female freshwater amphipods of three species from two continents: Gammarus duebeni celticusLiljeborg, 1852 and G. pulex(L., 1758) from Europe, and G. pseudolimnaeusBousfield, 1958 from North America. All three species were found to have measureable concentrations of both hormones in whole body lysate samples but the concentrations differed between species, with testosterone differing significantly between species only for male amphipods and estradiol differing significantly between species only for female amphipods. Concentrations of both testosterone and estrogen differed between males and females in two of the three species ( G. duebeni celticusand G. pseudolimnaeus). Females had the highest concentration of both hormones in G. duebeni celticusand the lowest concentration of both hormones in G. pseudolimnaeus. These results contribute to a growing body of evidence that these hormones are endogenously produced and biologically relevant in amphipods. Such evidence is particularly important in light of increasing prevalence of endocrine-disrupting compounds in the environment and the central role played by amphipods in aquatic ecosystems.

Predator-free space, functional responses and invasions.

1. Predator–prey interactions are mediated by the structural complexity of habitats, but disentangling the many facets of structure that contribute to this mediation remains elusive. In a world replete with altered landscapes and biological invasions, determining how structure mediates the interactions between predators and novel prey will contribute to our understanding of invasions and predator–prey dynamics in general.
2. Here, using simplified experimental arenas, we manipulate predator-free space, whilst holding surface area and volume constant, to quantify the effects on predator–prey interactions between two resident gammarid predators and an invasive prey, the Ponto-Caspian corophiid Chelicorophium curvispinum.
3. Systematically increasing predator-free space alters the functional responses (the relationship between prey density and consumption rate) of the amphipod predators by reducing attack rates and lengthening handling times. Crucially, functional response shape also changes subtly from destabilizing Type II towards stabilizing Type III, such that small increases in predator-free space to result in significant reductions in prey consumption at low prey densities.
4. Habitats with superficially similar structural complexity can have considerably divergent consequences for prey population stability in general and, particularly, for invasive prey establishing at low densities in novel habitats.

Eaten alive: cannibalism is enhanced by parasites.

Cannibalism is ubiquitous in nature and especially pervasive in consumers with stage-specific resource utilization in resource-limited environments. Cannibalism is thus influential in the structure and functioning of biological communities. Parasites are also pervasive in nature and, we hypothesize, might affect cannibalism since infection can alter host foraging behaviour. We investigated the effects of a common parasite, the microsporidian Pleistophora mulleri, on the cannibalism rate of its host, the freshwater amphipod Gammarus duebeni celticus. Parasitic infection increased the rate of cannibalism by adults towards uninfected juvenile conspecifics, as measured by adult functional responses, that is, the rate of resource uptake as a function of resource density. This may reflect the increased metabolic requirements of the host as driven by the parasite. Furthermore, when presented with a choice, uninfected adults preferred to cannibalize uninfected rather than infected juvenile conspecifics, probably reflecting selection pressure to avoid the risk of parasite acquisition. By contrast, infected adults were indiscriminate with respect to infection status of their victims, probably owing to metabolic costs of infection and the lack of risk as the cannibals were already infected. Thus parasitism, by enhancing cannibalism rates, may have previously unrecognized effects on stage structure and population dynamics for cannibalistic species and may also act as a selective pressure leading to changes in resource use.

Aquatic invasive alien species – top issues for their management. Outcomes from the IFI/EIFAAC conference “Freshwater Invasives – Networking for Strategy”

In November 2014, a new EU Regulation to address Invasive Alien Species (IAS) and protect biodiversity was published. This entered into force across the EU in January 2015. The aim of the Regulation is to ‘prevent the introduction of, control or eradicate alien species which threaten ecosystems, habitats or species’. In an effort to provide focus to the Regulation prior to its publishing and to identify the major issues relating to Invasive Alien Species in Europe, the views of invasive species experts from around the world were sought. These were consolidated at an international conference (Freshwater Invasives - Networking for Strategy (FINS)) that was held in Ireland in April 2013. A major outcome from this meeting of experts was the production of the Top 20 IAS issues that relate primarily to freshwater habitats but are also directly relevant to marine and terrestrial ecosystems. This list will support policy makers throughout the EU as preparations are made to implement this important piece of legislation. A further outcome from the conference was the formation of an expert IAS Advisory Group to support EIFAAC in its work on invasive species

Differential ecological impacts of an invader and a native under environmental change are revealed by comparative functional responses.

Predicting the ecological impacts of damaging invasive species under relevant environmental contexts is a major challenge, for which comparative functional responses (the relationship between resource availability and consumer uptake rate) have great potential. Here, the functional responses of Gammarus pulex, an ecologically damaging invader in freshwaters in Ireland and other islands, were compared with those of a native trophic equivalent Gammarus duebeni celticus. Experiments were conducted at two dissolved oxygen concentrations (80 and 50 % saturation), representative of anthropogenic water quality changes, using two larval prey, blackfly (Simuliidae spp.) and mayfly (Baetis rhodani). Overall, G. pulex had higher Type II functional responses and hence predatory impacts than G. d. celticus and the functional responses of both predators were reduced by lowered oxygen concentration. However, this reduction was of lower magnitude for the invader as compared to the native. Further, the invader functional response at low oxygen was comparable to that of the native at high oxygen. Attack rates of the two predators were similar, with low oxygen reducing these attack rates, but this effect occurred more strongly for blackfly than mayfly prey. Handling times were significantly lower for the invader compared with the native, and significantly higher at low oxygen, however, the effect of lowered oxygen on handling times was minimal for the invader and pronounced for the native. Maximum feeding rates were significantly greater for the invader compared with the native, and significantly reduced at low oxygen, with this effect again lesser for the invader as compared to the native. The greater functional responses of the invader corroborate with its impacts on recipient macroinvertebrate communities when it replaces the native. Further, our experiments predict that the impact of the invader will be less affected than the native under altered oxygen regimes driven by anthropogenic influences.