In disturbed environments, infected organisms have to face both parasitic and chemical stresses. Although this situation is common, few studies have been devoted to the effects of infection on hosts' energy reserves and antitoxic defence capacities, while parasite survival depends on host survival. In this study, we tested the consequences of an infection by Polymorphus minutus on the energy reserves (protein, lipid and glycogen) and antioxidant defence capacities (reduced glutathione, γ-glutamylcysteine ligase activity) of Gammarus roeseli males and females, in the absence of chemical stress. Moreover, malondialdehyde concentration was used as a toxicity biomarker. The results revealed that in infected G. roeseli, whatever their gender and the sampling month, protein and lipid contents were lower, but glycogen contents were higher. This could be explained by the fact that the parasite diverts part of the host's energy for its own development. Moreover, glutathione concentrations and γ-glutamylcysteine ligase activity were both lower, which could lead to lower antitoxic defence in the host. These results suggest negative effects on individuals in the case of additional stress (e.g. pollutant exposure). In the absence of chemical stress, the lower malondialdehyde level in infected gammarids could imply a probable protective effect of the parasite.

Here we report direct underwater observations of the pom pom anemone Liponema multicornis at sites in the western Gulf of Maine. The paucity of records from historic collections as well as recent studies suggest this species is widespread but numerically rare across the region. We found this species hosts high numbers of the commensal amphipod Stenothoe brevicornis amongst the tentacles. In addition, pandalid shrimp were observed using the anemone for shelter by occupying space around the distal edges of the tentacles and as a food source by feeding on bits of mucous. Krill were also observed feeding on amphipods occurring amongst the tentacles. A long history of using bottom contact fishing gear in the region may have reduced abundance, thereby reducing the functional role of L. multicornis as a source for refuge and prey of several crustacean species.

Factors constraining host specificity are poorly understood. Intraspecific variation in host preferences in generalist parasites may reveal which factors affect patterns of host use, and thus the evolution of specialization. Here, laboratory experiments examined genetic variation in host preferences and the effect of a refugium against infection on host use. Firstly, 6 cercarial clones of the trematode Maritrema novaezealandensis (ranging widely in heterozygosities) were exposed simultaneously to 2 alternative hosts, the amphipods Heterophoxus stephenseni and Paracalliope novizealandiae, to assess host preferences and fitness correlations with parasite heterozygosity. All clones showed a distinct preference for H. stephenseni, though the extent of this preference varied among clones. No clear association was found between heterozygosity and either parasite infection success or preference for a particular host. Secondly, cercariae were exposed to the same 2 amphipods in both the presence and absence of sand (refugium for H. stephenseni). Without sand, infection levels were significantly higher in H. stephenseni than in P. novizealandiae. With sand, H. stephenseni was able to hide, offsetting the parasite's intrinsic preferences for this host. These results demonstrate the existence of genetic variation in host preferences, as well as the effect of environmental variables on observed patterns of host use.

The amphipod (Crustacea: Peracarida) epibionts of the sea turtles Caretta caretta and Chelonia mydas were studied in the Gulf of Gabès. A total of eight amphipod species were identified, with Hyale grimaldii and Caprella andreae by far the most common. Among these species, Ericthonius punctatus and Ampithoe riedli are recorded for the first time as epibionts of Caretta caretta and Chelonia mydas, respectively, in the Mediterranean Sea.

Comparisons were made between the protein concentrations and protein profiles of uninfected Gammarus pulex and those infected with cystacanths of Pomphorhynchus laevis. Comparisons of the extractable haemolymph volumes from uninfected and infected G. pulex, when related to host tissue weight, showed that there was no difference in the host tissue/haemolymph volume ratio between infected and uninfected individuals. SDS-PAGE of haemolymph samples resolved 36 bands of protein, the major components being band 6(19 kDa) and 21(33 kDa). Densitometric analysis of haemolymph samples showed that infected gammarids had total protein concentrations elevated by 84%. The (88 kDa) subunit was identified as the monomer of the respiratory protein haemocyanin. This band was significantly elevated in infected individuals (one-way ANOVA P < 0.001); increasing from 30 to 45% of the total haemolymph proteins.

Characteristics of the second gnathopod are traditionally used to distinguish between species of caprellid amphipods. However, these distinctions are often subjective and can be variable within a species. Geometric morphometrics were used to quantitatively assess shape variation of the second gnathopod propodus of three species of caprellids in North America, including the non-native Caprella mutica. Gnathopod shapes of C. mutica specimens from different latitudes revealed distinct morphologies; the factors responsible for the shape variations are unknown. Allometric change of propodus shape was observed in C. mutica. Larger individuals showed a wide array of possible propodus morphologies. Despite this variability, there were clear differences between large specimens of C. mutica and two species native to North America: C. alaskana and C. kennerlyi. The use of geometric morphometrics and the thin-plate spline method can serve to both complement descriptions using traditional keys and aid in identification of non-native species in novel geographical regions.

In this paper, we describe the occurrence of a microsporidian parasite in female-biased populations of an intertidal amphipod, Corophium volutator (Pallas), at mudflat sites in the Bay of Fundy, Canada. Sequence data for the parasite's 16S rDNA indicate that it is a novel microsporidian species. This parasite was found principally in female host gonads, indicating that it might be a vertically transmitted, sex-distorting microparasite. At 4 sites each sampled in early and mid-summer, parasite prevalence varied from 0 to 21%. In the lab, infected mothers gave rise to more female-biased broods, than did uninfected mothers. Infection was not associated with size of females or with lowered survivorship of their young. Surprisingly, infected mothers actually had higher fertility controlling for body length than did uninfected mothers. Taken together, our results suggest that this novel microsporidian is likely a feminizing microparasite and is a contributing factor to local and widespread sex ratio distortion in C. volutator.

Vertically transmitted parasites may have positive, neutral or negative effects on host fitness, and are also predicted to exhibit sex-specific virulence to increase the proportion or fitness of the transmitting sex. We investigated these predictions in a study on the survival and sex ratio of offspring of the amphipod Gammarus roeseli from females infected by the vertically transmitted microsporidia Nosema granulosis. We found, to our knowledge, the first evidence for a positive relationship between N. granulosis infection and host survival. Infection was associated with sex ratio distortion, not by male-killing, but probably by parasite-induced feminization of putative G. roeseli males. This microsporidia also feminizes another amphipod host, Gammarus duebeni, which is phylogenetically and biogeographically distant from G. roeseli. Our study suggests that the reproductive system of gammarids is easily exploited by these vertically-transmitted parasites, although the effects of infections on host fitness may depend on specific host-parasite species interactions.

Predation between invading and native species can produce patterns of exclusion and coexistence. Dikerogammarus villosus, a Ponto-Caspian amphipod species, has invaded many central European freshwaters in the past decade, replacing native Gammarus amphipod species. For instance, the arrival of D. villosus in Holland has been accompanied by the decline of Gammarus duebeni and G. tigrinus populations within invaded systems. This study examined what may happen when D. villosus eventually encounters native Dutch populations of Gammarus pulex, and how factors such as microhabitat and food resource availability could contribute to a future species replacement or coexistence. A laboratory simulation of a lake/pooled area of river indicated that G. pulex and D. villosus differed in distribution within the same habitat, and showed that although the distribution of the native differed in the presence of the invader, the presence of the native had no effect on the distribution of the invader. Gammarus pulex suffered severe intraguild predation (IGP) from D. villosus in mixed species treatments with no reciprocal predation of D. villosus by G. pulex. This IGP occurred regardless of whether no alternative food resource was available (91% of the G. pulex population eliminated after 7 days), or alternative foods/prey were available to excess, such as leaf material (85%), chironomids (77%) or fish food flakes (74%). We conclude that although differential microdistribution of the two species could promote coexistence, the presence of alternative foods/prey resources, merely slow the rate of IGP and replacement of the native by the invader. Our study joins one of an increasing number emphasizing the potential damaging impacts of D. villosus on native communities.

The microsporidian parasite, Pleistophora mulleri, infects the abdominal muscle of the freshwater amphipod Gammarus duebeni celticus. We recently showed that P. mulleri infection was associated with G. d. celticus hosts being more vulnerable to predation by the invasive amphipod Gammarus pulex. Parasitized G. d. celticus also had a reduced ability to prey upon other co-occurring amphipods. We suggested the parasite may have pervasive influences on host ecology and behaviour. Here, we examine the association between P. mulleri parasitism and parameters influencing individual host fitness, behaviour and interspecific interactions. We also investigate the relationship between parasite prevalence and host population structure in the field. In our G. d. celticus study population, P. mulleri prevalence was strongly seasonal, ranging from 8·5% in summer to 44·9% in winter. The relative abundance of hosts with the heaviest parasite burden increased during summer, which coincided with high host mortality, suggesting that parasitism may regulate host abundance to some degree. Females were more likely to be parasitized than males and parasitized males were paired with smaller females than unparasitized males. Parasitism was associated with reduction in the host's activity level and reduced both its predation on the isopod Asellus aquaticus and aggression towards precopula pairs of the invasive G. pulex. We discuss the pervasive influence of this parasite on the ecology of its host.

Dikerogammarus villosus, a freshwater invasive amphipod, exhibits conspicuous pigmentation polymorphism. This polymorphism is documented in two recently colonized areas, the Saône and Moselle rivers (north-eastern France), and some of the mechanisms by which pigmentation polymorphism can arise and be maintained are addressed. Body size, reproductive status, fecundity and mate choice are compared among morphs of D. villosus in field samples collected in summer 2001. Body size and female gonad developmental stage were comparable among the different morphs, suggesting that polymorphism is not the result of changes in pigmentation with age or moult-cycle. Fecundity and reproductive status (paired vs non-paired) were not affected by pigmentation morph either. A random combination of morphs in pre-copula pairs was observed, showing that the colour pattern does not play a major role in mate choice. A strong size-assortative pairing was found, and this pattern was similar among male morphs. Overall, our study shows that morph polymorphism is not related to different stages in the moult cycle or the life cycle, and is not maintained by morph-assortative pairing. Alternative explanations to the mate choice hypothesis for the maintenance of pigmentation polymorphism are discussed.

The amphipod Gammarus pulex is an intermediate host to the acanthocephalan fish parasite Echinorhynchus truttae. Gammarus pulex has a wide trophic repertoire, feeding as a herbivore, detritivore and predator. In this study an examination was made of the effects of E. truttae parasitism on components of the G. pulex diet: stream-conditioned leaves, dead chironomids and live juvenile isopods Asellus aquaticus. Over 21 days, parasitism had no effect on daily feeding rates or wet weights of G. pulex fed on leaves or chironomids. Parasitism had a significant effect on the number of A. aquaticus killed by G. pulex, with parasitized individuals killing significantly fewer than their unparasitized counterparts. In addition, unparasitized amphipods killed all size classes of A. aquaticus indiscriminately, whereas parasitized animals tended to kill the smaller size classes. The impacts of the parasitism of G. pulex throughout the wider freshwater community are discussed.

The complexity of the life-cycle of many parasitic helminths has driven the evolution of several well-documented adaptations serving to facilitate the completion of each difficult step in the cycle. In some trematode taxa, selection has even favoured the truncation of the life-cycle: progenetic larval stages (metacercariae) are capable of producing eggs inside their usual intermediate hosts, eliminating the need to be transmitted by predation to a definitive host. In some species, progenesis is shown by all individuals, whereas in other species both the normal and truncated life-cycles coexist in the same populations. Here, the strategies chosen by normal and progenetic metacercariae of the trematode Coitocaecum parvum are investigated and compared. Normal metacercariae, i.e. metacercariae that only develop into adults following ingestion by a suitable fish definitive host, were not capable of manipulating the behaviour of their amphipod intermediate host in ways that could facilitate their capture by fish. These metacercariae were associated with increased mortality in amphipods. Progenetic metacercariae, on the other hand, showed lower virulence levels than normal metacercariae. At the time of their death, amphipods harbouring progenetic metacercariae contained close to the maximum number of eggs that can be produced by progenetic metacercariae, suggesting that the parasite lowers its virulence to keep its host alive long enough to maximize egg output. The maintenance of the 2 strategies in the C. parvum population suggests that they may have equal fitness payoffs: progenesis and low virulence guarantee the production of at least some eggs, whereas the normal life-cycle and its associated higher virulence in the intermediate host provides a low probability of much higher fecundity.

Larval helminths often share individual intermediate hosts with other larval worms of the same or different species. In the case of immature acanthocephalans capable of altering the phenotype of their intermediate hosts, the benefits or costs of host sharing can be evaluated in terms of increased or decreased probability of transmission to a suitable definitive host. Competitive interactions among the immature stages of acanthocephalans within the intermediate host could create additional costs of host sharing, however. The effects of intraspecific and interspecific interactions were measured in 3 sympatric species of acanthocephalans exploiting a population of the amphipod Echinogammarus stammeri in the River Brenta, Italy. The strength of interactions was assessed from differences in the size achieved by infective cystacanths in the intermediate host. The size of Pomphorhynchus laevis cystacanths was not correlated with host size, whereas the size of Acanthocephalus clavula and Polymorphus minutus cystacanths increased with host size. Reductions in cystacanth size caused by intraspecific competition were only detected in P. laevis, but may also occur in both A. clavula and P. minutus. When co-occurring in the same amphipod with cystacanths of A. clavula, cystacanths of P. laevis attained a smaller size than when they occurred on their own. This effect was not reciprocal, with the size of A. clavula cystacanths not being affected. This supports earlier suggestions that it is adaptive for A. clavula to associate with P. laevis in amphipod intermediate hosts, with both species going to the same fish definitive hosts. In contrast, cystacanths of P. laevis achieved their largest size when they co-occurred in an amphipod with a cystacanth of P. minutus, which has a different definitive host (i.e. birds). These findings suggest that the net benefits of sharing an intermediate host can only be estimated by taking into account both the effects on transmission success and the consequences for cystacanth development.