Distribution and host range of the microsporidian Pleistophora mulleri

ABSTRACT. Microsporidia of the genus Pleistophora are important parasites of fish and crustacea. Pleistophora mulleri has been described previously as a parasite of the gammarid amphipod crustacean Gammarus duebeni celticus in Irish freshwater habitats. Through a survey of European G. duebeni populations, P. mulleri was found to be widely distributed in the western British Isles (Wales, Scotland, and the Isle of Man), and populations of the subspecies Gammarus duebeni duebeni as well as G. d. celticus were infected. Pleistophora infections were also detected in G. d. duebeni sampled from the Bay of Gdansk on Poland's Baltic coast, indicating a wide distribution of Pleistophora in European G. duebeni . Sequencing and phylogenetic analysis of the 16S rRNA , 18S rRNA , and Rpb1 genes of P. mulleri suggest that this species may be synonymous with P. typicalis , a parasite of fish. These findings suggest that amphipod crustaceans may act as intermediate or reservoir hosts for microsporidian parasites of fish.     

Distribution and microhabitats of native and non-native gammarids (Amphipoda, Crustacea) in Brittany, with particular reference to the endangered endemic sub-species Gammarus duebeni celticus

Aim  To assess temporal changes in gammarid distribution in Brittany and microhabitat-use overlap between the endangered endemic Gammarus duebeni celticus Stock & Pinkster, 1970 , the expanding natives G. pulex (Linnaeus, 1758) and Echinogammarus berilloni (Catta, 1878), and the introduced G. tigrinus Sexton, 1939.
Location  Brittany and adjacent regions in western France.
Methods  The spatial and temporal patterns in distribution of gammarids at the scale of Brittany were studied using 351 sites. Longitudinal distributions (from the source to the estuary of the river) and microhabitat-use (substratum type and water velocity) were also considered in selected rivers.
Results  At the regional scale, all species occurred together less often than expected statistically, with significant deviations from expected for G. pulex vs. both G. duebeni celticus and G. tigrinus , and for E. berilloni vs. both G. duebeni celticus and G. tigrinus . However, at the microhabitat scale, E. berilloni occurred significantly more often than expected with the endemic G. duebeni celticus , and this appears to be due to similar substratum and water velocity preferences, although at both the regional and microhabitat scales E. berilloni prefers wider streams than G. duebeni celticus . This study reveals a decline in the endangered G. duebeni celticus since 1970.
Main conclusions  The longitudinal and local distributions of G. duebeni celticus , and the higher-than-expected co-occurrence of the species with G. pulex , suggest that the decline of the endemic species may be due to changes in the environment and/or interference from native G. pulex , which is expanding its range in Brittany. The results are discussed as regards to the consequences for regional biodiversity.     

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.     

Differential drift and parasitism in invading and native Gammarus spp. (Crustacea: Amphipoda)

Invading and native species often interact directly, such as by predation, producing patterns of exclusion and coexistence. Less direct factors, such as interactions with the broader abiotic and biotic environment, may also contribute to such patterns, but these have received less recognition. In Northern Ireland, the North American Gammarus tigrinus has invaded freshwaters populated with the native Gammarus duebeni celticus , with intraguild predation between the two implicated in their relative success. However, these species also engage in day and night "drifting", an activity that subjects amphipods to intense predation from fish and wildfowl. Sampling of two rivers where the invader and native co-occur showed that, compared with the benthos, G. tigrinus was underrepresented and G. d. celticus overrepresented in the drift. In addition, G. tigrinus were free from parasites, whereas some G. d. celticus harboured the acanthocephalans Polymorphus minutus and Echinorhynchus truttae and the muscle wasting microsporidian Pleistophora sp. (new species). Compared with the benthos, G. d. celticus parasitized with P. minutus were overrepresented and unparasitized individuals underrepresented in the drift. The opposite pattern was found with Pleistophora sp . In laboratory experiments, G. tigrinus were less positively phototropic and less "active" than G. d. celticus (unparasitized animals). Polymorphus minutus increased G. d. celticus positive phototropism and activity, while Pleistophora sp. increased positive phototropism but decreased activity. Previous studies show that the invader G. tigrinus is more disadvantaged by intraguild predation from the native G. d. celticus than vice versa. However, the native appears more disadvantaged with respect to drift, parasitism and the interaction of the two. These factors may mitigate direct interactions and help explain complex patterns of coexistence between these invader and native species.     

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

SUMMARY 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.
2. The lowland stretch of the River Lissan is dissected by a weir, which has slowed the upstream invasion by G. pulex . We took physicochemical measurements and macroinvertebrate samples from three contiguous 150 m reaches of this stretch with G. pulex only, mixed Gammarus and G. d. celticus only communities.
3. We found no biologically significant differences in physicochemistry among the three reaches. Overall invertebrate densities did not differ among reaches. However, G. pulex numerically dominated its reach, whilst G. d. celticus abundance was relatively low in its reach. The G. pulex reach had significantly higher overall biomass because of high invader abundance. In addition, both diversity and species richness of macroinvertebrate communities were lower in the G. pulex than the G. d. celticus reach, with the mixed Gammarus reach intermediate.
4. Ordination indicated distinctly different associations of invertebrate community samples and taxa that were best explained by the distributions of the Gammarus species. Again, the mixed Gammarus species samples were intermediate.
5. This study indicates that the invasive G. pulex has a greater impact on macroinvertebrate community composition than the native G. d. celticus , probably through biotic interactions such as predation.     

Microtubule systems associated with the septate junctions of the gill cells of four gammarid amphipods

The microtubular systems associated with the septate junctions of the gill epithelial cells of four species of gammarid amphipod are described. The four species examined included two relatively stenohaline marine forms, Chaetogammarus marinus and Gammarus locusta; a highly euryhaline species, Gammarus duebeni, and a stenohaline freshwater species, Gammarus pulex. Of these amphipods, G. locusta and C. marinus maintain only a limited osmotic gradient between their haemolymph and the medium and have a poorly developed junctional microtubular system; G. pulex has haemolymph which is some 300 mOsmol hypertonic to freshwater and has a well ordered system of microtubules on both sides of fairly long septate junctions; G. duebeni from brackish water tend to have a somewhat shorter length of septate junctions lined by one or occasionally by a double row of microtubules. The most complex junctional microtubular systems are shown by specimens of the freshwater race of G. duebeni celticus which have been acclimated to seawater. These can take the form of multiple arrays in which some microtubules are linked to the plasma membrane by dense strands. It is suggested that these findings are consistent with the hypothesis that one role of these microtubules is to provide mechanical stability to enable the integrity of the septate junctions to be maintained during osmotic stress.     

Electrophoretic approach to the biochemical systematics of gammarids

By utilizing the techniques for electrophoretic separation of proteins by vertical starch gels, the biochemical systematics of 10 Gammaridae species obtained from marine, brackish and freshwater habitats was studied. They includedChaetogammarus marinus, Gammarus zaddachi, G. salinus, G. oceanicus, G. tigrinus, G. chevreuxi, G. locusta, G. duebeni duebeni, G. d. celticus, G. pulex pulex, andG. fossarum. For comparison of electrophoretic mobilities selected enzymes (phosphoglucose isomerase, glutamate oxalacetate transaminase, arginine phosphokinase, hexokinase, leucine amino peptidase, mannose 6-phosphate isomerase) were assayed. They were used as diagnostic characters in terms of electrophoretic identities or diversities of most frequent alleles at polymorphic gene loci. These criteria could be applied to estimate intrageneric enzymic variation and degrees of genetic relatedness between the crustacean amphipod species under consideration, thereby complementing traditional morphological classification.     

The nature and implications of differential predation between Gammarus pulex and G. duebeni celticus (Crustacea: Amphipoda)

The mutual predatory behaviour between Gammarus pulex (Linnaeus, 1758) and Gammarus duebeni celticus (Stock & Pinkster, 1970) is described. The implications of the predatory nature of these species are discussed in terms of the feeding ecology of Gammarus and other amphipods. Predation rates by males on moulted congeneric females are assessed and compared. When males are present at the moult of a congeneric female, 100%of females are eaten in both species/sex combinations. When females are allowed to recover from moulting prior to presentation to a congeneric male, predation rates are significantly reduced in both species. However, significantly more G. d. celticus females are eaten by G. pulex males compared with the reciprocal situation. The predatory 'clumping' behaviour of both species is described and shown to be a form of feeding frenzy upon congenerics. Clumping behaviour also results in significantly higher rates of congeneric predation on G. d. celticus females than on G. pulex females. Support is thus given to the hypothesis that differential predation by males on moulted females may be the primary mechanism by which the introduced G. pulex has displaced the native Irish species G. d. celticus . The implications of amphipod congeneric predation are discussed in terms of observed patterns of interspecific coexistence and exclusion.     

Predation on mayfly nymph, Baetis rhodani, by native and introduced Gammarus: direct effects and the facilitation of predation by salmonids

1. In a series of laboratory experiments, we assessed the predatory nature of the native Irish amphipod, Gammarus duebeni celticus , and the introduced G. pulex , towards the mayfly nymph Baetis rhodani . We also investigated alterations in microhabitat use and drift behaviour of B . rhodani in the presence of Gammarus , and indirect predatory interactions with juvenile Atlantic salmon, Salmo salar .
2. In trials with single predators and prey, B. rhodani survival was significantly lower when Gammarus were free to interact with nymphs as than when Gammarus were isolated from them. The invader G. pulex reduced the survival of B. rhodani more rapidly than did the native G. d. celticus . Both Gammarus spp. were active predators.
3. In `patch' experiments, B. rhodani survival was significantly lower both when G. pulex and G. d. celticus were present, although the effect of the two Gammarus species did not differ. Again, active predation of nymphs by Gammarus was observed. Significantly more nymphs occurred on the top and sides of a tile, and per capita drifts were significantly higher, when Gammarus were present. Baetis rhodani per capita drift was also significantly higher in the presence of the introduced G. pulex than with the native G. d. celticus .
4.  Gammarus facilitated predation by salmon parr of B. rhodani by significantly increasing fish–nymph encounters on exposed gravel and in the drift. There were no differential effects of the two Gammarus spp. on fish – B . rhodani encounters or consumption.
5. We conclude that Gammarus as a predator can have lethal, nonlethal, direct and indirect effects in freshwaters. We stress the need for recognition of this predatory role when assigning Gammarus spp. to a `Functional Feeding Group'.     

The cannibalistic behaviour of two Gammarus species (Crustacea: Amphipoda)

This study examines the cannibalistic behaviour of the freshwater amphipods Gammarus duebeni celticus Stock & Pinkster, 1970 and G. pulex (L., 1758). In the first experiment, interactions were staged among all combinations of single adult males, single adult females, adults in the precopulatory mate-guarding phase and juveniles. Cannibalism by inter-moult individuals on newly moulted conspecifics occurred in all interaction categories in both species. Gammarus d. celticus , however, were significantly more cannibalistic than G. pulex. Cannibalism between and within sex and size categories (males > females > juveniles) was facilitated by the vulnerability of individuals at moult. Individuals of smaller size categories, however, did not cannibalize newly moulted conspecifics of larger size categories. Males were less cannibalistic on newly moulted females than on newly moulted males and juveniles and, when in the precopulatory condition, appeared to defend females from cannibalistic attacks. In a second experiment, stream conditions were simulated in the laboratory and replicated populations monitored for nine weeks. High levels of cannibalism, and the species and sex differences in cannibalism identified in the first experiment, were confirmed under these heterogeneous conditions. Cannibalism by males on their newly moulted female mating partners, termed 'reversed' sexual cannibalism, was further investigated. When males were deprived of foraging opportunities, cannibalism of precopulatory partners was significantly more frequent. The occurrence of 'reversed' sexual cannibalism is thus interpreted as a conflict between motivation to feed and motivation to mate.     

Introduction of the non-indigenous amphipod Gammarus pulex alters population dynamics and diet of juvenile trout Salmo trutta

Summary 1. The amphipod Gammarus pulex , introduced to Irish rivers with the aim of enhancing trout feeding, is displacing the native Gammarus duebeni celticus . These two species are generally associated with different environmental conditions and macroinvertebrate communities, confounding assessment of effects of the invader as compared with the native on fish populations. Here, we uncouple effects of the two Gammarus species from environmental gradients.
2. A weir dissects a lowland stretch of the River Lissan, slowing the upstream invasion by G. pulex and resulting in contiguous G. pulex , mixed species and G. d. celticus reaches. Total invertebrate abundance and biomass in the benthos were significantly higher in the G. pulex reach, driven by high invader abundance, with low abundance of other taxa. Gammarus pulex was particularly prominent in night-time drift.
3. Correspondingly, densities and biomass of 0+ trout were significantly higher in the G. pulex reach, while instantaneous loss rates were lower. Fish growth rates were similar among the three reaches.
4. In the G. pulex reach, this invader dominated the diet of 0+ trout, leading to ingestion of significantly higher invertebrate biomass than fish in the other reaches. Fish generally preyed on Gammarus in proportion to its abundance, but exhibited some positive selection for G. pulex in the invaded reach.
5. The negative effects of the invader on native invertebrates are contrasted with positive effects on juvenile trout. This indicates changes in energy flux after invasion, with differential resource use or assimilation by G. pulex probable underlying mechanisms. As the frequency of amphipod invasions increases globally, investigations of their role as strong interactors at multiple levels of ecological organisation is required if the consequences of deliberate and unintentional introductions are to be predicted, and ultimately, prevented.     

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.     

Parasite transmission and cannibalism in an amphipod (Crustacea)

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.     

Phylogenetic Relationships of Five Species of the Amphipod Genus Gammarus

The phylogenetic relationships of five amphipod species of the genus Gummarus: G. oceanicus, G. locusta, G. salinus, G. zaddachi and G. duebeni are presented based on starch gel electrophoresis of 12 enzymes determined by 17 loci. The five species separate into three groups: G. locusta and G. duebeni each represent a distinct group while G. oceanicus, G. salinus and G. zaddachi together form a third group. This grouping results when the number of diagnostic loci, as well as when genetic distance between species pairs, are considered. The results compare well with morphological and, to some extent, ecological data.     

Quantitative analysis of sexual dimorphism and sex ratio in Hyphydrus ovatus (Coleoptera: Dytiscidae)

Sexual dimorphism in size, shape, and mass relative to length, and sex ratios are quantified for populations of Hyphydrus ovatus. a dytiscid beetle Males of H ovatus are not only longer than females, but also significantly larger in elytron length, thorax width, head width, leg length, total width, total depth, and abdominal segment length Two local populations differ slightly but significantly in total depth and abdominal segment length, but sexual dimorphism in size is similar for the two populations Hyphydrus ovatus are also sexually dimorphic in shape, with males having relatively broader heads and thoraxes than females The two populations differed slightly but significantly in relative abdominal segment length, but as with size, sexual dimorphism in shape is similar for the two populations Males are relatively heavier than females, although the slope of the log mass vs log length relationship is the same for the two sexes Sex ratios in field samples vary significantly over the summer, with percent females declining from c 50% to c 15% Sex ratios are significantly below 50% females m two of five monthly samples and in the total pooled set of samples Sexual dimorphism in size, shape, and relative mass, combined with male-biased sex ratios suggest that larger size of male H ovatus is a product of sexual selection     

Reflectance of sexually dichromatic UV‐blue patches varies during the breeding season and between two subspecies of Gallotia galloti (Squamata: Lacertidae)

Body coloration is sexually dimorphic in many vertebrate species, including lizards, in which males are often more conspicuous than females. A detailed analysis of the relative size of coloured patches and their reflectance, including the ultraviolet (UV) range, has rarely been performed. In the present work we quantified sexual dimorphism in body traits and surface area of all lateral patches from adult females and males of two subspecies of Gallotia galloti (G. g. galloti and G. g. eisentrauti). We also analysed the magnitude of sexual dichromatism in the UV‐visible reflectance of such patches and the changes in patch size and brightness during the reproductive season (April–July). Males had significantly larger patch areas (relative to their snout‐vent length) and higher brightness (mainly in the UV‐blue range) than did females in both subspecies. The comparison of relative patch areas among months did not reach statistical significance. However, patch brightness significantly changed during the breeding season: that of the UV‐blue (300–495 nm) range from lizards of the two subspecies was significantly larger in June than in April, while brightness in the 495–700 nm range in G. g. galloti was larger in May, June, and July than in April. A different pattern of dichromatism was also detected in the two populations, with G. g. eisentrauti being more sexually dichromatic than G. g. galloti. We discuss the results in terms of possible evolutionary causes for the sexual dichromatism related to different ecological characteristics of the habitats where each subspecies live. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 556–569.     

Cytogenetic and biometric observations on members of the Anopheles gambiae complex in Mozambique

Four species of the Anopheles gambiae complex were identified in Mozambique (East Africa) by chromosomal analysis. They were An. merus, An. gambiae s.s., An. arabiensis and An. quadriannulatus. An. merus was observed in coastal zones as well as in inner areas where the rivers are tidal and brackish and/or the soil is salty. An. gambiae s.s. is present in the central-northern regions (north of Save river) from the coast to the western mountains. On the coast it is often sympatric with An. merus. It is apparently absent south of Save river. An. arabiensis was observed in samples from the north-western hilly and mountainous areas, sympatrically with An. gambiae s.s., as well as south of Save river where often it is sympatric with An. merus. Only one specimen of An. quadriannulatus was observed. It was from a small sample collected feeding on bovid in a southern locality (Bela Vista-Maputo area). No inversion polymorphism was observed in the 446 An. merus identified. A quite low degree of inversion polymorphism was shown by both An. gambiae s.s. and An. arabiensis, involving 2Rb, 2La and 2Rb inversion systems respectively. By morphological analysis of cytogenetically identified samples of three species of the complex, number of sensilla coeloconica and palpal ratio were confirmed to be useful to distinguish An. merus from An. gambiae s.s./An. arabiensis. The overlapping areas between brackish and freshwater species become smaller when both characters are considered together, 1.5% and 3.5% being the probabilities of confusing An. merus with An. gambiae s.s. and An. arabiensis respectively.     

Correlation between rheologic properties and in vitro ciliary transport of rat nasal mucus.

The relationship between mucus rheologic variables and in vitro ciliary transport was investigated in mucus samples collected from the upper airways of 30 Wistar rats. In vitro mucus transportability was determined by means of the frog palate preparation. Rheologic evaluation was done by measuring the rigidity modulus (log G*, representing the vectorial sum of viscosity and elasticity) and the loss tangent (tan delta, i.e. the ratio between viscosity and elasticity) at 1 and 100 radian/s using a magnetic microrheometer. The correlation between the rheologic variables and in vitro mucus transportability was made by stepwise multiple linear regression analysis, with frog palate transport rate considered as the dependent variable. A significant relationship was obtained between the rheologic parameters (log G* and tan delta) measured at 1 radian/s and the frog palate transport ratio. The relative speed of mucus samples was related to rheology according to the following relationship: rat/frog speed ratio = 1.666-0.434 log G*-0.331 tan delta, for G* and delta determined at 1 radian/s (multiple r = 0.666, p < 0.001). Transport rates predicted from the above formula gave a satisfactory fit to those observed in a second set of 30 rats. The present results indicate that the overall mucus impedance, as well as the ratio between viscosity and elasticity, are important in determining the efficiency of clearance. In addition, it was shown that measurements performed by applying relatively low frequency deformations are preferable for predicting ciliary transport.     

Sexual dimorphism and allometry in two seed beetles (Coleoptera: Bruchidae)

Male Callosobruchus chinensis (Coleoptera: Bruchidae) have elaborated, pectinate antennae, which are absent from conspecific females and both sexes of a congener, Callosobruchus maculatus. To begin to unravel the mechanisms producing this striking dimorphism, we examined which morphological traits best explain body size variation in bruchid beetles and quantified sexual dimorphism of antenna size through allometric analyses. Using principal component analyses, we found that elytron length and pronotum width were significantly correlated with the first principal component, which was interpreted as explaining variation in body size. Regressions of log‐transformed body size measures on log‐transformed antenna length revealed that males of both species had longer antennae than conspecific females for any given body size, although most of this effect was attributable to higher intercepts, rather than increased allometry, in males. Comparisons among heterospecific males revealed that C. maculatus males have noticeably longer antennae than C. chinensis males at large body sizes. Callosobruchus chinensis males, thus, appear to have increased the receptive area of their antennae by adding to the width of, rather than further elongating, their antennae. Finally, we found evidence for positive allometry between log‐transformed antenna length and log‐transformed antenna width in C. chinensis males. We discuss our results in the context of evidence supporting the presence of an additional, and potentially unique, sex pheromone in C. chinensis females.     

Parasitism and epibiosis in native and non-native gammarids in freshwater in Ireland

The introduced amphipod crustaceans Gammarus pulex and G. tigrinus are displacing the native G. ducheni celticus in a number of freshwater sites in Northern Ireland. We investigated parasite and epibiont infection in populations in the Rivet Lagan and Lough Neagh where both native and invading species occur. Prevalence of the four parasites and epibionts observed was higher in the native G. d. celticus than in the invading amphipods at both field sites. In Lough Neagh. G. d. celticus individuals suffered higher burdens of the rotifer Embata parasitica and the protozoan Epistylis in comparison with the invading species. These patterns may reflect host specificity by the parasites or may result from different susceptibilities of the native and invading host species. We consider the influence of parasitism on host invasions and resulting species distributions.