Parasite mediated predation between native and invasive amphipods.

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.

Factors influencing the distribution of native and introduced Gammarus spp. in Irish river systems

Populations of Gammarus duebeni celticus, previously the only amphipod species resident in the rivers of the Lough Neagh catchment, N. Ireland, have been subjected to invasion by G. pulex from the British mainland. Numerous previous studies have investigated the potential behavioural mechanisms, principally differential mutual predation, underlying the replacement of G. d. celticus by G. pulex in Irish waters, and the mutually exclusive distributions of these species in Britain and mainland Europe. However, the relative degree of influence of abiotic versus biotic factors in structuring these amphipod communities remains unresolved. This study used principal component analysis (PCA) to distinguish physico-chemical parameters that have significant roles in determining the current distribution of G. pulex relative to G. d. celticus in L. Neagh rivers. We show that the original domination of rivers by the native G. d, celticus has changed radically, with many sites in several rivers containing either both species or only G. pulex. G. pulex was more abundant than the G. d. celticus in sites with low dissolved oxygen levels. This was reflected in the macroinvertebrate assemblages associated with G. pulex in these sites, which tended to be those tolerant of low biological water quality. The present study thus emphasizes the importance of the habitat template, particularly water quality, for Gammarus spp. interactions. If rivers become increasingly stressed by organic pollution, it is probable the range expansion of G. pulex will continue. Because these two species are not ecological equivalents, the outcomes of G. pulex incursions into G. d. celticus sites may ultimately depend on the prevailing physico-chemical regimes in each site.

Predatory nature of the littoral amphipod Echinogammarus marinus: gut content analysis and effects of alternative food and substrate heterogeneity

Interspecific interactions are major structuring forces in marine littoral communities; however, it is unclear which of these interactions are exhibited by many key-component species. Gut content analysis showed that the ubiquitous rocky/cobble shore amphipod Echinogammarus marinas, often ascribed as a mesograzer, consumes both algae and macroinvertebrates. Further, laboratory experiments showed that E. marinus is an active predator of such macroinvertebrates, killing and consuming the isopod Jaera nordmanni and the oligochaete Tubificoides benedii. Predatory impacts of E. marinus were not alleviated by the presence of alternative food in the form of alga discs. However, in the presence of prey, consumption of alga by E. marinus was significantly reduced. Further, survival of prey was significantly higher when substrate was provided, but predation remained significant and did not decline with further increases in substrate heterogeneity. We conclude that such amphipods can have pervasive predatory impacts on a range of species, with implications for community structure, diversity and functioning.

Invasion by the amphipod Gammarus pulex alters community composition of native freshwater macroinvertebrates

Assessing the effects of invading species on native community structure is often confounded by environmental factors and weakened by lack of replicated, long-term pre- and post-invasion monitoring. Here, we uncouple the community effects of a freshwater amphipod invader from environmental differences. In Irish rivers, the introduced Gammarus pulex replaces the native Gammarus duebeni celticus. However, the River Lissan in Northern Ireland is dissected by a weir that has slowed the upstream invasion by G. pulex. This allowed us in 2000 to sample three contiguous 150-m reaches that were (1) G. pulex dominated; (2) mixed Gammarus spp.; and (3) G. duebeni celticus only. In 2003, we resampled these reaches and one additional of mixed Gammarus species and one with only G. duebeni celticus further upstream. In temperature, conductivity, and pH, there were statistically significant but no biologically relevant differences among the five reaches of 2003, and between the three reaches surveyed in both years. Although there was evidence of recovery in macroinvertebrate diversity and richness in invaded reaches between years, continued upstream invasion was associated with sustained reductions in these community metrics as compared to un-invaded sites. Community ordination indicated (1) different associations of community composition attributed to the distribution, abundance, and biomass of the invader; and (2) increasing similarity of invaded communities over time. The impact mechanisms of G. pulex on macroinvertebrate community composition may include predation and competition. The consequences of the observed community changes for ecosystem functioning require further investigation.

Differences in composition of macroinvertebrate communities with invasive and native Gammarus spp. (Crustacea : Amphipoda)

1. Assessing the effects on communities of invasive species is often confounded by environmental factors. In Irish rivers, the introduced amphipod Gammarus pulex replaces the native G. duebeni celticus in lowland stretches. The two amphipods are associated with different macroinvertebrate communities, which may in part be the result of natural longitudinal physicochemical change. However, this hinders assessment of any direct community impacts of the invasive as compared with the native species. Here, we report on a fortuitous circumstance that allowed us to uncouple the community effects of Gammarus species from environmental differences.

Functional responses of the intertidal amphipod Echinogammarus marinus: effects of prey supply, model selection and habitat complexity.

The influence of predation in structuring ecological communities can be informed by examining the shape and magnitude of the functional response of predators towards prey. We derived functional responses of the ubiquitous intertidal amphipod Echinogammarus marinus towards one of its preferred prey species, the isopod Jaera nordmanni. First, we examined the form of the functional response where prey were replaced following consumption, as compared to the usual experimental design where prey density in each replicate is allowed to deplete. E. marinus exhibited Type II functional responses, i.e. inversely density-dependent predation of J. nordmanni that increased linearly with prey availability at low densities, but decreased with further prey supply. In both prey replacement and non-replacement experiments, handling times and maximum feeding rates were similar. The non-replacement design underestimated attack rates compared to when prey were replaced. We then compared the use of Holling’s disc equation (assuming constant prey density) with the more appropriate Rogers’ random predator equation (accounting for prey depletion) using the prey non-replacement data. Rogers’ equation returned significantly greater attack rates but lower maximum feeding rates, indicating that model choice has significant implications for parameter estimates. We then manipulated habitat complexity and found significantly reduced predation by the amphipod in complex as opposed to simple habitat structure. Further, the functional response changed from a Type II in simple habitats to a sigmoidal, density-dependent Type III response in complex habitats, which may impart stability on the predator−prey interaction. Enhanced habitat complexity returned significantly lower attack rates, higher handling times and lower maximum feeding rates. These findings illustrate the sensitivity of the functional response to variations in prey supply, model selection and habitat complexity and, further, that E. marinus could potentially determine the local exclusion and persistence of prey through habitat-mediated changes in its predatory functional responses.

How simple grazing rules can lead to persistent boundaries in vegetation communities.

Natural landscape boundaries between vegetation communities are dynamically influenced by the selective grazing of herbivores. Here we show how this may be an emergent property of very simple animal decisions, without the need for any sophisticated choice rules etc., using a model based on biased diffusion. Animal grazing intensity is coupled with plant competition, resulting in reaction-diffusion dynamics, from which stable boundaries spontaneously emerge. In the model, animals affect their resources by both consumption and trampling. It is assumed that forage consists of two heterogeneously distributed competing resource species, one that is preferred (grass) over the other (heather) by the animals. The solutions to the resulting system of differential equations for three cases a) optimal foraging, b) random walk foraging and c) taxis-diffusion are presented. Optimal and random foraging gave unrealistic results, but taxis-diffusion accorded well with field observations. Persistent boundaries between patches of near-monoculture vegetation were predicted, with these boundaries drifting in response to overall grazing pressure (grass advancing with increased grazing and vice versa). The reaction-taxis-diffusion model provides the first mathematical explanation for such vegetation mosaic dynamics and the parameters of the model are open to experimental testing.