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In Northern Ireland, the amphipods Gammarus duebeni celticus (native) and G. pulex (invasive) coexist in some places, whilst in others the native species has been replaced by the invader. We explored the role of parasites in mediating interactions between these amphipods, which demonstrate mutual intraguild predation (IGP: predation between animals that also compete for prey). IGP and cannibalism can be important factors in structuring populations and communities. We investigated the effects of parasitism on rates of IGP between G. d. celticus and G. pulex and on cannibalism within each species by comparing functional responses (FRs: relationships between the use of a prey resource and its availability). Infection with the microsporidian Pleistophora mulleri caused an increase in IGP and cannibalism by G. d. celticus, which showed increased attack rates and reduced prey handling times. In contrast, infection with the acanthocephalan parasite Echinorhynchus truttae did not alter IGP or cannibalism by G. pulex. A prey preference experiment revealed that both amphipods were more likely to feed on heterospecific rather than conspecific prey, and this was also corroborated by the fact that overall IGP FRs were higher than cannibalism FRs. This may be selectively advantageous, as feeding on heterospecific prey removes possible competitors without the risk of consuming juvenile kin or acquiring parasites from infected conspecifics. Infection of the native G. d. celticus with P. mulleri enhanced IGP on the invasive G. pulex, which is likely to facilitate the coexistence of the 2 species.

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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.

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Species distributions are changing at an unprecedented rate owing to human activity. We examine how two key processes of redistribution - biological invasion and disease emergence - are interlinked. There are many parallels between invasion and emergence processes, and invasions can drive the spread of new diseases to wildlife. We examine the potential impacts of invasion and disease emergence, and discuss how these threats can be countered, focusing on biosecurity. In contrast with international policy on emerging diseases of humans and managed species, policy on invasive species and parasites of wildlife is fragmented, and the lack of international cooperation encourages individual parties to minimize their input into control. We call for international policy that acknowledges the strong links between emerging diseases and invasion risk.

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Predatory functional responses play integral roles in predator-prey dynamics, and their assessment promises greater understanding and prediction of the predatory impacts of invasive species. Other interspecific interactions, however, such as parasitism and higher-order predation, have the potential to modify predator-prey interactions and thus the predictive capability of the comparative functional response approach. We used a four-species community module (higher-order predator; focal native or invasive predators; parasites of focal predators; native prey) to compare the predatory functional responses of native Gammarus duebeni celticus and invasive Gammarus pulex amphipods towards three invertebrate prey species (Asellus aquaticus, Simulium spp., Baetis rhodani), thus, quantifying the context dependencies of parasitism and a higher-order fish predator on these functional responses. Our functional response experiments demonstrated that the invasive amphipod had a higher predatory impact (lower handling time) on two of three prey species, which reflects patterns of impact observed in the field. The community module also revealed that parasitism had context-dependent influences, for one prey species, with the potential to further reduce the predatory impact of the invasive amphipod or increase the predatory impact of the native amphipod in the presence of a higher-order fish predator. Partial consumption of prey was similar for both predators and occurred increasingly in the order A. aquaticus, Simulium spp. and B. rhodani. This was associated with increasing prey densities, but showed no context dependencies with parasitism or higher-order fish predator. This study supports the applicability of comparative functional responses as a tool to predict and assess invasive species impacts incorporating multiple context dependencies.

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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.

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Emerging infectious diseases (EIDs) are recognized as having significant social, economic and ecological costs, threatening human health, food security, wildlife conservation and biodiversity. We review the processes underlying the emergence of infectious disease, focusing on the similarities and differences between conceptual models of disease emergence and biological invasions in general.Study of the IUCN's list of the world's worst invaders reveals that disease is cited as a driver behind the conservation, medical or economic impact of nearly a quarter of the species on the data base.The emergence of novel diseases in new host species are, in essence, examples of invasions by parasites. Many of the ecological and anthropogenic drivers of disease emergence and classical invasions are also shared, with environmental change and global transport providing opportunities for the introduction and spread of invaders and novel parasites.The phases of disease emergence and biological invasions have many parallels; particularly the early and late phases, where demographic and anthropogenic factors are key drivers. However, there are also differences in the intermediate phases, where host-parasite co-evolution plays a crucial role in determining parasite establishment in novel hosts.Similar opportunities and constraints on control and management occur at the different phases of invasions and disease emergence. However, exploitation of host immune responses offers additional control opportunities through contact control and vaccination against EIDs. We propose that cross-fertilization between the disciplines of disease emergence and invasion biology may provide further insights into their prediction, control and management.

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BACKGROUND: Sex-ratio distorting parasites are of interest due to their effects upon host population dynamics and their potential to influence the evolution of host sex determination systems. In theory, the ability to distort host sex-ratios allows a parasite with efficient vertical (hereditary) transmission to dispense completely with horizontal (infectious) transmission. However, recent empirical studies indicate that some sex-ratio distorting parasites have retained the capability for horizontal transmission. RESULTS: Numerical simulations using biologically realistic parameters suggest that a feminising parasite is only likely to lose the capability for horizontal transmission if its host occurs at low density and/or has a male-biased primary sex ratio. It is also demonstrated that even a small amount of horizontal transmission can allow multiple feminising parasites to coexist within a single host population. Finally it is shown that, by boosting its host's rate of population growth, a feminising parasite can increase its own horizontal transmission and allow the invasion of other, more virulent parasites. CONCLUSIONS: The prediction that sex-ratio distorting parasites are likely to retain a degree of horizontal transmission has important implications for the epidemiology and host-parasite interactions of these organisms. It may also explain the frequent co-occurrence of several sex-ratio distorting parasite species in nature.

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Invasive species can have profound impacts on communities and it is increasingly recognized that such effects may be mediated by parasitism. The 'enemy release' hypothesis posits that invaders may be successful and have high impacts owing to escape from parasitism. Alternatively, we hypothesize that parasites may increase host feeding rates and hence parasitized invaders may have increased community impacts. Here, we investigate the influence of parasitism on the predatory impact of the invasive freshwater amphipod Gammarus pulex. Up to 70 per cent of individuals are infected with the acanthocephalan parasite Echinorhynchus truttae, but parasitized individuals were no different in body condition to those unparasitized. Parasitized individuals consumed significantly more prey (Asellus aquaticus; Isopoda) than did unparasitized individuals. Both parasitized and unparasitized individuals displayed Type-II functional responses (FRs), with the FR for parasitized individuals rising more steeply, with a higher asymptote, compared with unparasitized individuals. While the parasite reduced the fitness of individual females, we predict a minor effect on population recruitment because of low parasite prevalence in the peak reproductive period. The parasite thus has a large per capita effect on predatory rate but a low population fitness effect, and thus may enhance rather than reduce the impact of this invader.

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Intraguild predation (IGP) is common in communities, yet theory suggests it should not often persist and coexistence of participating species should be rare. As parasitism can play keystone roles in interactions between competitors, and between predators and prey, here we examine the role of parasites in maintaining IGP. We used numerical exploration of population dynamic equations to determine coexistence and exclusion zones for two species engaged in IGP with shared parasitism. We demonstrate that parasitism increases the range of conditions leading to coexistence when the parasite exerts a greater deleterious effect on the 'stronger' species in terms of the combined effects of competition and predation. Such a parasite can enable an inferior competitor that is also the less predatory to persist, and may actually lead to numerical dominance of this species.

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We present a synthesis of empirical and theoretical work investigating how parasites influence competitive and predatory interactions between other species. We examine the direct and indirect effects of parasitism and discuss examples of density and parasite-induced trait-mediated effects. Recent work reveals previously unrecognized complexity in parasite-mediated interactions. In addition to parasite-modified and apparent competition leading to species exclusion or enabling coexistence, parasites and predators interact in different ways to regulate or destablize the population dynamics of their joint prey. An emerging area is the impact of parasites on intraguild predation (IGP). Parasites can increase vulnerability of infected individuals to cannibalism or predation resulting in reversed species dominance in IGP hierarchies. We discuss the potential significance of parasites for community structure and biodiversity, in particular their role in promoting species exclusion or coexistence and the impact of emerging diseases. Ongoing invasions provide examples where parasites mediate native/invader interactions and play a key role in determining the outcome of invasions. We highlight the need for more quantitative data to assess the impact of parasites on communities, and the combination of theoretical and empirical studies to examine how the effects of parasitism scale up to community-level processes.

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Microsporidia are unusual amongst eukaryotic parasites in that they utilize both vertical and horizontal transmission and vertically transmitted species can cause sex ratio distortion in their host. Here we study vertical transmission in two species of feminising microsporidia, Nosema granulosis and Dictyocoela duebenum, infecting a single population of the crustacean host Gammarus duebeni and measure the effect of temperature on parasite transmission and replication. N. granulosis was vertically transmitted to 82% of the host embryos and D. duebenum was transmitted to 72% of host embryos. For both parasites, we report relatively low parasite burdens in developing host embryos. However, the parasites differ in their pattern of replication and burden within developing embryos. Whilst N. granulosis undergoes replication during host development, the burden of D. duebenum declines, leading us to propose that parasite dosage and feminisation efficiency underlie the different parasite frequencies in the field. We also examine the effect of temperature on parasite transmission and replication. Temperature does not affect the percentage of young that inherit the infection. However, low temperatures inhibit parasite replication relative to host cell division, resulting in a reduction in parasite burden in infected embryos. The reduced parasite burden at low temperatures may underpin reduced feminization at low temperatures and so limit the spread of sex ratio distorters through the host population.

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Local adaptation theory predicts that, on average, most parasite species should be locally adapted to their hosts (more suited to hosts from local than distant populations). Local adaptation has been studied for many horizontally transmitted parasites, however, vertically transmitted parasites have received little attention. Here we present the first study of local adaptation in an animal/parasite system where the parasite is vertically transmitted. We investigate local adaptation and patterns of virulence in a crustacean host infected with the vertically transmitted microsporidian Nosema granulosis. Nosema granulosis is vertically transmitted to successive generations of its crustacean host, Gammarus duebeni and infects up to 46% of adult females in natural populations. We investigate local adaptation using artificial horizontal infection of different host populations in the UK. Parasites were artificially inoculated from a donor population into recipient hosts from the sympatric population and into hosts from three allopatric populations in the UK. The parasite was successfully established in hosts from all populations regardless of location, infecting 45% of the recipients. Nosema granulosis was vertically (transovarially) transmitted to 39% of the offspring of artificially infected females. Parasite burden (intensity of infection) in developing embryos differed significantly between host populations and was an order of magnitude higher in the sympatric population, suggesting some degree of host population specificity with the parasite adapted to its local host population. In contrast with natural infections, artificial infection with the parasite resulted in substantial virulence, with reduced host fecundity (24%) and survival (44%) of infected hosts from all the populations regardless of location. We discuss our findings in relation to theories of local adaptation and parasite-host coevolution.

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In its freshwater amphipod host Gammarus duebeni celticus, the microsporidian parasite Pleistophora mulleri showed 23% transmission efficiency when uninfected individuals were fed infected tissue, but 0% transmission by water-borne and coprophagous routes. Cannibalism between unparasitised and parasitised individuals was significantly in favour of the former (37% compared to 0%). In addition, cannibalism between parasitised individuals was significantly higher than between unparasitised individuals (27% compared to 0%). Thus, parasitised individuals were more likely to be cannibalised by both unparasitised and parasitised individuals. We discuss the conflicting selective forces within this host/parasite relationship, the implications of parasite mediated cannibalism for host population structure and the impacts this may have on the wider aquatic community.

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Parasites can structure biological communities directly through population regulation and indirectly by processes such as apparent competition. However, the role of parasites in the process of biological invasion is less well understood and mechanisms of parasite mediation of predation among hosts are unclear. Mutual predation between native and invading species is an important factor in determining the outcome of invasions in freshwater amphipod communities. Here, we show that parasites mediate mutual intraguild predation among native and invading species and may thereby facilitate the invasion process. We find that the native amphipod Gammarus duebeni celticus is host to a microsporidian parasite, Pleistophora sp. (new species), with a frequency of infection of 0-90%. However, the parasite does not infect three invading species, G. tigrinus, G. pulex and Crangonyx pseudogracilis. In field and laboratory manipulations, we show that the parasite exhibits cryptic virulence: the parasite does not affect host fitness in single-species populations, but virulence becomes apparent when the native and invading species interact. That is, infection has no direct effect on G. d. celticus survivorship, size or fecundity; however, in mixed-species experiments, parasitized natives show a reduced capacity to prey on the smaller invading species and are more likely to be preyed upon by the largest invading species. Thus, by altering dominance relationships and hierarchies of mutual predation, parasitism strongly influences, and has the potential to change, the outcome of biological invasions.

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In a river survey, Gammarus pulex amphipods both unparasitised and parasitised with the acanthocephalan Echinorhynchus truttae were distributed similarly with respect to flow regimen, tending to be more abundant in faster, shallower, riffle patches. However, there was a higher prevalence of parasitism in faster, shallower areas than in slower, deeper areas and abundance correlated with macrophyte coverage for unparasitised but not parasitised amphipods, indicating subtle differences in habitat usage. A laboratory 'patch' simulation indicated that parasitism influenced micro-distribution. There were higher proportions of unparasitised amphipods in/under stone substrates and within weed. In contrast, there were higher proportions of parasitised amphipods in the water column and at the water surface. As the experiment progressed, unparasitised but not parasitised amphipod habitat usage shifted from those micro-habitats above the substrate and in the water column to those in/under the substrates. Experiments also demonstrated that parasitised amphipods were more active and had a greater preference for illumination. Previous studies of the effects of acanthocephalan parasitism of amphipod hosts have focussed on how drift behaviour is altered, now we show that subtle differences in micro-habitat usage could translate to greatly increased vulnerability to fish predation. We discuss how aggregation of parasitised individuals within specific habitats could promote parasite transmission.

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