Leeches Baicalobdella torquata feed on hemolymph but have a low effect on the cellular immune response of amphipod Eulimnogammarus verrucosus from Lake Baikal.

Lake Baikal is one of the largest and oldest freshwater reservoirs on the planet with a huge endemic diversity of amphipods (Amphipoda, Crustacea). These crustaceans have various symbiotic relationships, including the rarely described phenomenon of leech parasitism on amphipods. It is known that leeches feeding on hemolymph of crustacean hosts can influence their physiology, especially under stressful conditions. Here we show that leeches Baicalobdella torquata (Grube, 1871) found on gills of Eulimnogammarus verrucosus (Gerstfeldt, 1858), one of the most abundant amphipods in the Baikal littoral zone, indeed feed on the hemolymph of their host. However, the leech infection had no effect on immune parameters such as hemocyte concentration or phenoloxidase activity and also did not affect glycogen content. The intensity of hemocyte reaction to foreign bodies in a primary culture was identical between leech-free and leech-infected animals. Artificial infection with leeches also had only a subtle effect on the course of a model microbial infection in terms of hemocyte concentration and composition. Despite we cannot fully exclude deleterious effects of the parasites, our study indicates a low influence of a few leeches on E. verrucosus and shows that leech-infected amphipods can be used at least for some types of ecophysiological experiments.

Possible seasonal and diurnal modulation of Gammarus pulex (Crustacea, Amphipoda) drift by microsporidian parasites.

In lotic freshwater ecosystems, the drift or downstream movement of animals (e.g., macroinvertebrates) constitutes a key dispersal pathway, thus shaping ecological and evolutionary patterns. There is evidence that macroinvertebrate drift may be modulated by parasites. However, most studies on parasite modulation of host drifting behavior have focused on acanthocephalans, whereas other parasites, such as microsporidians, have been largely neglected. This study provides new insight into possible seasonal and diurnal modulation of amphipod (Crustacea: Gammaridae) drift by microsporidian parasites. Three 72 h drift experiments were deployed in a German lowland stream in October 2021, April, and July 2022. The prevalence and composition of ten microsporidian parasites in Gammarus pulex clade E varied seasonally, diurnally, and between drifting and stationary specimens of G. pulex. Prevalence was generally higher in drifting amphipods than in stationary ones, mainly due to differences in host size. However, for two parasites, the prevalence in drift samples was highest during daytime suggesting changes in host phototaxis likely related to the parasite's mode of transmission and site of infection. Alterations in drifting behavior may have important implications for G. pulex population dynamics and microsporidians' dispersal. The underlying mechanisms are more complex than previously thought.

Disentangling the determinants of symbiotic species richness in native and invasive gammarids (Crustacea, Amphipoda) of the Baltic region.

Dispersal of alien species is a global problem threatening native biodiversity. Co-introduction of non-native parasites and pathogens adds to the severity of this threat, but this indirect impact has received less attention. To shed light on the key factors determining the richness of microorganisms in native and invasive host species, we compared symbiotic (parasitic and epibiotic) communities of gammarids across different habitats and localities along the Baltic coast of Poland. Seven gammarid species, two native and five invasive, were sampled from 16 freshwater and brackish localities. Sixty symbiotic species of microorganisms of nine phyla were identified. This taxonomically diverse species assemblage of symbionts allowed us to assess the effect of host translocation and regional ecological determinants driving assembly richness in the gammarid hosts. Our results revealed that (i) the current assemblages of symbionts of gammarid hosts in the Baltic region are formed by native and co-introduced species; (ii) species richness of the symbiotic community was higher in the native Gammarus pulex than in the invasive hosts, probably reflecting a process of species loss by invasive gammarids in the new area and the distinct habitat conditions occupied by G. pulex and invasive hosts; (iii) both host species and locality were key drivers shaping assembly composition of symbionts, whereas habitat condition (freshwater versus brackish) was a stronger determinant of communities than geographic distance; (iv) the dispersion patterns of the individual species richness of symbiotic communities were best described by Poisson distributions; in the case of an invasive host, the dispersion of the rich species diversity may switch to a right-skewed negative binomial distribution, suggesting a host-mediated regulation process. We believe this is the first analysis of the symbiotic species richness in native and invasive gammarid hosts in European waters based on original field data and a broad range of taxonomic groups including Microsporidia, Choanozoa, Ciliophora, Apicomplexa, Platyhelminthes, Nematoda, Nematomorha, Acanthocephala and Rotifera, to document the patterns of species composition and distribution.

Intermediate host patterns of acanthocephalans in the Weser river system: co-invasion vs host capture.

Anthropogenic interference is a major driver of ecological change in freshwater ecosystems. Pollution and the introduction of new species not only alter macrozoobenthic community structures, but can also affect their respective parasite communities. The ecology of the Weser river system experienced a drastic decline in biodiversity over the past century due to salinization caused by the local potash industry. As a response, the amphipod Gammarus tigrinus was released into the Werra in 1957. A few decades after the introduction and subsequent spread of this North American species, its natural acanthocephalan Paratenuisentis ambiguus was recorded in the Weser in 1988, where it had captured the European eel Anguilla anguilla as a novel host. To assess the recent ecological changes in the acanthocephalan parasite community, we investigated gammarids and eel in the Weser river system. In addition to P. ambiguus, 3 Pomphorhynchus species and Polymorphus cf. minutus were discovered. The introduced G. tigrinus serves as a novel intermediate host for the acanthocephalans Pomphorhynchus tereticollis and P. cf. minutus in the tributary Werra. Pomphorhynchus laevis is persistent in the tributary Fulda in its indigenous host Gammarus pulex. Pomphorhynchus bosniacus colonized the Weser with its Ponto-Caspian intermediate host Dikerogammarus villosus. This study highlights the anthropogenically driven changes in ecology and evolution in the Weser river system. Based on morphological and phylogenetic identification, the shifts in distribution and host usage described here for the first time contribute to the puzzling taxonomy of the genus Pomphorhynchus in times of ecological globalization.

The trematode Podocotyle atomon modulates biochemical responses of Gammarus locusta to thermal stress but not its feeding rate or survival.

Although parasitism is one of the most common species interactions in nature, the role of parasites in their hosts' thermal tolerance is often neglected. This study examined the ability of the trematode Podocotyle atomon to modulate the feeding and stress response of Gammarus locusta towards temperature. To accomplish this, infected and uninfected females and males of Gammarus locusta were exposed to temperatures (2, 6, 10, 14, 18, 22, 26, 30 °C) for six days. Shredding (change in food biomass) and defecation rates (as complementary measure to shredding rate) were measured as proxies for feeding activity. Lipid and glycogen concentrations (energy reserves), catalase (oxidative stress indicator), and phenoloxidase (an immunological response in invertebrates) were additionally measured. Gammarid survival was optimal at 10 °C as estimated by the linear model and was unaffected by trematode infection. Both temperature and sex influenced the direction of infection effect on phenoloxidase. Infected females presented lower phenoloxidase activity than uninfected females at 14 and 18 °C, while males remained unaffected by infection. Catalase activity increased at warmer temperatures for infected males and uninfected females. Higher activity of this enzyme at colder temperatures occurred only for infected females. Infection decreased lipid content in gammarids by 14 %. Infected males had significantly less glycogen than uninfected, while infected females showed the opposite trend. The largest infection effects were observed for catalase and phenoloxidase activity. An exacerbation of catalase activity in infected males at warmer temperatures might indicate (in the long-term) unsustainable, overwhelming, and perhaps lethal conditions in a warming sea. A decrease in phenoloxidase activity in infected females at warmer temperatures might indicate a reduction in the potential for fighting opportunistic infections. Results highlight the relevance of parasites and host sex in organismal homeostasis and provide useful insights into the organismal stability of a widespread amphipod in a warming sea.

Infection with an acanthocephalan helminth reduces anxiety-like behaviour in crustacean host.

Trophically transmitted heteroxenous parasites of diverse clades can decrease or reverse antipredator behaviours in their intermediate hosts, thereby increasing their chances of reaching their final hosts. Such behavioural alterations could result from compromised cognitive abilities affecting fear- or more generally stress-related neurophysiological pathways. We tested this hypothesis in a key model system in the study of parasitic manipulation, the fish acanthocephalan parasite Pomphorhynchus tereticollis and its intermediate crustacean host Gammarus fossarum, using the 'threat of electric shock' paradigm. We exposed uninfected and infected G. fossarum to chronic and/or acute electric shock programs at two different intensities (voltage), and then quantified their sheltering behaviour as a proxy for anxiety-like state. Infected gammarids did not express anxiety-like response to electric shocks, while uninfected gammarids hid more when exposed to acute treatments, and when exposed to the high intensity chronic treatment. Interestingly, the lack of response in infected gammarids depended on parasite developmental stage. Our results support the hypothesis that this acanthocephalan parasite impacts the general anxiety-like circuitry of their intermediate host. Further studies are needed to investigate whether it involves inappropriate processing of information, impaired integration, or altered activation of downstream pathways initiating behavioural action.

Histopathological survey for parasite groups in Gammarus varsoviensis (Amphipoda).

Invasive non-native amphipods (Crustacea) are becoming a model system in which to explore the impact and diversity of invasive parasites-parasites that are carried along an invasion route with their hosts. Gammarus varsoviensis is a freshwater amphipod species that has a recently explored invasion history. We provide a histopathological survey for a putatively invasive non-native population of this amphipod, identifying 8 symbiotic groups: Acanthocephala, Rotifera, Digenea, ciliated protozoa, Haplosporidia, Microsporidia, 'Candidatus Aquirickettsiella', and a putative nudivirus, at various prevalence. Our survey indicates that the parasites have no sex bias and that each has the potential to be carried in either sex along an invasion route. We discuss the pathology and prevalence of the above symbiotic groups and whether those that are parasitic may pose a risk if G. varsoviensis were to carry them to novel locations.

Txikispora philomaios n. sp., n. g., a micro-eukaryotic pathogen of amphipods, reveals parasitism and hidden diversity in Class Filasterea.

This study provides a morphological, ultrastructural, and phylogenetic characterization of a novel micro-eukaryotic parasite (2.3-2.6 µm) infecting amphipod genera Echinogammarus and Orchestia. Longitudinal studies across two years revealed that infection prevalence peaked in late April and May, reaching 64% in Echinogammarus sp. and 15% in Orchestia sp., but was seldom detected during the rest of the year. The parasite infected predominantly hemolymph, connective tissue, tegument, and gonad, although hepatopancreas and nervous tissue were affected in heavier infections, eliciting melanization and granuloma formation. Cell division occurred inside walled parasitic cysts, often within host hemocytes, resulting in hemolymph congestion. Small subunit (18S) rRNA gene phylogenies including related environmental sequences placed the novel parasite as a highly divergent lineage within Class Filasterea, which together with Choanoflagellatea represent the closest protistan relatives of Metazoa. We describe the new parasite as Txikispora philomaios n. sp. n. g., the first confirmed parasitic filasterean lineage, which otherwise comprises four free-living flagellates and a rarely observed endosymbiont of snails. Lineage-specific PCR probing of other hosts and surrounding environments only detected T. philomaios in the platyhelminth Procerodes sp. We expand the known diversity of Filasterea by targeted searches of metagenomic datasets, resulting in 13 previously unknown lineages from environmental samples.

Two parasites in one host: spatiotemporal dynamics and co-occurrence of Microsporidia and Rickettsia in an amphipod host.

Biological interactions can greatly influence the abundance of species. This is also true for parasitic species that share the same host. Microsporidia and Rickettsia are widespread intracellular parasites in populations of Paracalliope fluviatilis, the most common freshwater amphipods in New Zealand. Although both parasites coexist in many populations, it is unclear whether they interact with each other. Here, we investigated spatial−temporal dynamics and co-occurrence of the two parasites, Microsporidia and Rickettsia in P. fluviatilis hosts, across one annual cycle and in three different locations. Prevalence of both Microsporidia and Rickettsia changed over time. However, while the prevalence of Rickettsia varied significantly between sampling times, that of Microsporidia did not change significantly and remained relatively low. The two parasites therefore followed different temporal patterns. Also, the prevalence of both parasites differed among locations, though the two species reached their highest prevalence in different locations. Lastly, there was no evidence for positive or negative associations between the two parasite species; the presence of one parasite in an individual host does not appear to influence the probability of infection by the other parasite. Their respective prevalence may follow different patterns among populations on a larger spatial scale due to environmental heterogeneity across locations.

Dictyocoela microsporidia diversity and co-diversification with their host, a gammarid species complex (Crustacea, Amphipoda) with an old history of divergence and high endemic diversity.

BACKGROUND: Although the processes of co-evolution between parasites and their hosts are well known, evidence of co-speciation remains scarce. Microsporidian intracellular parasites, due to intimate relationships with their hosts and mixed mode of transmission (horizontal but also vertical, from mother to offspring), may represent an interesting biological model for investigating co-speciation. Amphipod crustaceans, especially gammarids, are regular hosts of microsporidian parasites, in particular the Dictyocoela spp., which have so far been found limited to these amphipods and are known to use a vertical mode of transmission. The amphipod genus Gammarus has a diversification history spanning the last 50-60 Mya and an extensive cryptic diversity in most of the nominal species. Here, we investigated the degree of co-diversification between Dictyocoela and Gammarus balcanicus, an amphipod with high degrees of ancient cryptic diversification and lineage endemism, by examining the genetic diversity of these parasites over the entire geographic range of the host. We hypothesised that the strong host diversification and vertical transmission of Dictyocoela would promote co-diversification. RESULTS: Using the parasite SSU rDNA as a molecular marker, analyzing 2225 host specimens from 88 sites covering whole host range, we found 31 haplogroups of Dictyocoela, 30 of which were novel, belonging to four Dictyocoela species already known to infect other Gammarus spp. The relationships between Dictyocoela and gammarids is therefore ancient, with the speciation in parasites preceding those of the hosts. Each novel haplogroup was nevertheless specific to G. balcanicus, leaving the possibility for subsequent co-diversification process during host diversification. A Procrustean Approach to Co-phylogeny (PACo) analysis revealed that diversification of Dictyocoela was not random with respect to that of the host. We found high degrees of congruence between the diversification of G. balcanicus and that of Dictyocoela roeselum and D. muelleri. However, we also found some incongruences between host and Dictyocoela phylogenies, e.g. in D. duebenum, probably due to host shifts between different G. balcanicus cryptic lineages. CONCLUSION: The evolutionary history of Dictyocoela and Gammarus balcanicus represents an example of an overall host-parasite co-diversification, including cases of host shifts.

Application of DNA barcoding and morphometric analysis in differentiating cystacanths of Polymorphus species (Acanthocephala: Polymorphidae) from central Alberta, Canada.

Acanthocephalans are multi-host endoparasites, many of which use freshwater amphipods as intermediate hosts for their larval stages (e.g., cystacanths) while adults live in the intestines of vertebrates, including waterfowl. In central Alberta, Canada, several co-occurring species of the acanthocephalan genus Polymorphus use the amphipod Gammarus lacustris Sars, 1863 as an intermediate host. We applied DNA barcoding and morphometric analysis to differentiate cystacanth larvae from G. lacustris sampled from 17 Albertan water bodies. We slide-mounted specimens and measured morphological traits relating to proboscis hooks. We sequenced the standard DNA barcoding region of the mitochondrial cytochrome c oxidase subunit I gene (COI). Morphometric analysis suggested that the acanthocephalans we collected belonged to four morphologically different groups that keyed to Polymorphus contortus (Bremser, 1821) Travassos, 1926; P. marilis Van Cleave, 1939; P. paradoxus Connel et Corner, 1957; and P. strumosoides (Lundström, 1942) Amin, 2013. Our Bayesian tree based on COI sequences generally corroborated the morphological results and supported that the specimens assigned to P. cf. contortus and P. cf. strumosoides belong to two distinct species. In contrast, the Bayesian tree showed that specimens of P. cf. marilis were nested as a cluster within the P. cf. paradoxus clade. Similarly, small pairwise genetic distance (< 2%) between specimens identified as P. cf. contortus and P. cf. strumosoides suggests that they are conspecific. Future studies should use morphology and sequence data from adult acanthocephalans to assess the taxonomic identity of the cystacanth-based Polymorphus taxa. Our study is the first to provide genetic information for the four Polymorphus taxa and emphasizes the importance of applying multiple approaches to differentiate parasite species.

Increased temperature has no consequence for behavioral manipulation despite effects on both partners in the interaction between a crustacean host and a manipulative parasite.

Parasites alter many traits of their hosts. In particular, parasites known as "manipulative" may increase their probability of transmission by inducing phenotypic alterations in their intermediate hosts. Although parasitic-induced alterations can modify species' ecological roles, the proximate factors modulating this phenomenon remain poorly known. As temperature is known to affect host-parasite associations, understanding its precise impact has become a major challenge in a context of global warming. Gammarids are ecologically important freshwater crustaceans and serve as intermediate hosts for several acanthocephalan species. These parasites induce multiple effects on gammarids, including alterations of their behavior, ultimately leading to modifications in their functional role. Here, experimental infections were used to assess the effect of two temperatures on several traits of the association between Gammarus pulex and its acanthocephalan parasite Pomphorhynchus laevis. Elevated temperature affected hosts and parasites in multiple ways (decreased host survival, increased gammarids activity, faster parasites development and proboscis eversion). However, behavioral manipulation was unaffected by temperature. These results suggest that predicted change in temperature may have little consequences on the trophic transmission of parasites through changes in manipulation, although it may modify it through increased infection success and faster parasites development.

Origin of the natural variation in the storage of dietary carotenoids in freshwater amphipod crustaceans.

Carotenoids are diverse lipophilic natural pigments which are stored in variable amounts by animals. Given the multiple biological functions of carotenoids, such variation may have strong implications in evolutionary biology. Crustaceans such as Gammarus amphipods store large amounts of these pigments and inter-population variation occurs. While differences in parasite selective pressure have been proposed to explain this variation, the contribution of other factors such as genetic differences in the gammarid ability to assimilate and/or store pigments, and the environmental availability of carotenoids cannot be dismissed. This study investigates the relative contributions of the gammarid genotype and of the environmental availability of carotenoids in the natural variability in carotenoid storage. It further explores the link of this natural variability in carotenoid storage with major crustacean immune parameters. We addressed these aspects using the cryptic diversity in the amphipod crustacean Gammarus fossarum and a diet supplementation protocol in the laboratory. Our results suggest that natural variation in G. fossarum storage of dietary carotenoids results from both the availability of the pigments in the environment and the genetically-based ability of the gammarids to assimilate and/or store them, which is associated to levels of stimulation of cellular immune defences. While our results may support the hypothesis that carotenoids storage in this crustacean may evolve in response to parasitic pressure, a better understanding of the specific roles of this large pigment storage in the crustacean physiology is needed.

The diversity of microsporidian parasites infecting the Holarctic amphipod Gammarus lacustris from the Baikal region is dominated by the genus Dictyocoela.

Microsporidia are a highly diverse group of single-celled eukaryotic parasites related to fungi and infecting hosts belonging to all groups of eukaryotes, including some protists, invertebrate and vertebrate animals. We investigated the diversity of microsporidia in the Holarctic amphipod species Gammarus lacustris from mostly, but not limited to, water bodies in the Lake Baikal region. Ribosomal DNA sequencing and host transcriptome sequencing data from various works show that this species is predominantly infected by representatives of the genus Dictyocoela and probably has some features underlying this specific interaction.

Morphological comparison of genetically differentiated Polymorphus cf. minutus types.

In the present study, we analyzed the morphology of three genetic types of the bird-infecting acanthocephalan Polymorphus cf. minutus (PspT1, PspT2, PspT3), mainly based on the cystacanth-stage obtained from amphipods (Gammarus fossarum, Gammarus pulex, Gammarus roeselii, Echinogammarus spp.). Males and females were pooled as there was no considerable difference between the sexes concerning the hook measurements. Additionally, we conducted a laboratory infection of one domestic duck for each Polymorphus type, to compare their performance and localization in this host species, and to obtain adult specimens for morphological comparison. The recovery rate from the ducks 4 weeks after infection was 16% for PspT1, 23.8% for PspT2, and 25% for PspT3. The adult worms were gravid, and the females contained mature eggs. Hook size did not differ considerably between cystacanths and adults of the respective type. The three Polymorphus types could be distinguished based on the cystacanth stage by a linear discriminant analysis that included hook measurements, proboscis length, proboscis width, and number of longitudinal hook rows and hooks per row. Furthermore, PspT3 was more different from PspT1 and PspT2 than the latter types from each other. Mainly the number of longitudinal hook rows differed in PspT3 from the existing descriptions of P. minutus (mainly 14 vs. mainly 16 rows). Potentially, PspT3 could be a non-indigenous parasite that was introduced with G. roeselii and that adapted to use the indigenous G. pulex as a host, while PspT2 might have been introduced to central Europe together with Echinogammarus spp.

A molecular war: convergent and ontogenetic evidence for adaptive host manipulation in related parasites infecting divergent hosts.

Mermithids (phylum Nematoda) and hairworms (phylum Nematomorpha) somehow drive their arthropod hosts into water, which is essential for the worms' survival after egression. The mechanisms behind this behavioural change have been investigated in hairworms, but not in mermithids. Establishing a similar mechanistic basis for host behavioural change between these two distantly related parasitic groups would provide strong convergent evidence for adaptive manipulation and insight into how these parasites modify and/or create behaviour. Here, we search for this convergence, and also contrast changes in physiology between hosts infected with immature and mature mermithids to provide the first ontogenetic evidence for adaptive manipulation by disentangling host response and pathology from the parasite's apparent manipulative effects. We used SWATH-mass spectrometry on brains of Forficula auricularia (earwig) and Bellorchestia quoyana (sandhopper), infected with the mermithids Mermis nigrescens and Thaumamermis zealandica, respectively, at both immature and mature stages of infection, to quantify proteomic changes resulting from mermithid infection. Across both hosts (and hairworm-infected hosts, from earlier studies), the general function of dysregulated proteins was conserved. Proteins involved in energy generation/mobilization were dysregulated, corroborating reports of erratic/hyperactive behaviour in infected hosts. Dysregulated proteins involved in axon/dendrite and synapse modulation were also common to all hosts, suggesting neuronal manipulation is involved in inducing positive hydrotaxis. Furthermore, downregulation of CamKII and associated proteins suggest manipulation of memory also contributes to the behavioural shift.

Amphipod disease: model systems, invasions and systematics-Introduction to DAO Special 8.

Amphipods are a group of globally abundant Crustacea present throughout terrestrial, marine and freshwater ecosystems. These organisms host a highly diverse systematic assemblage of parasites and pathogens, which are closely linked to the host's evolution and ecological niche. Such symbioses have been found to affect the behaviour, physiology and overall health of amphipod hosts; including effects at both the individual and population scale, altering aquatic trophic structure and possibly representing far reaching consequences for fisheries species and predatory species. Amphipod diseases explored in this Special have been linked with biological invasions, systematics, behavioural ecology, ecotoxicology, epidemiology, host physiology and cannibalistic tendencies. These studies exemplify the importance of amphipod research and provide keystone studies for the use of these animals as model systems for understanding the effects of disease in crustacean assemblages.

Pathogens and other symbionts of the Amphipoda: taxonomic diversity and pathological significance.

With over 10000 species of Amphipoda currently described, this order is one of the most diverse groups of freshwater and marine Crustacea. Members of this group are globally distributed, and many are keystone species and ecosystem engineers within their respective ecologies. As with most organisms, disease is a key factor that can alter population size, behaviour, survival, invasion potential and physiology of amphipod hosts. This review explores symbiont diversity and pathology in amphipods by coalescing a range of current and historical literature to provide the first full review of our understanding of amphipod disease. The review is broken into 2 parts. The first half explores amphipod microparasites, which include data pertaining to viruses, bacteria, fungi, oomycetes, microsporidians, dinoflagellates, myxozoans, ascetosporeans, mesomycetozoeans, apicomplexans and ciliophorans. The second half reports the metazoan macroparasites of Amphipoda, including rotifers, trematodes, acanthocephalans, nematodes, cestodes and parasitic Crustacea. In all cases we have endeavoured to provide a complete list of known species that cause disease in amphipods, while also exploring the effects of parasitism. Although our understanding of disease in amphipods requires greater research efforts to better define taxonomic diversity and host effects of amphipod symbionts, research to date has made huge progress in cataloguing and experimentally determining the effects of disease upon amphipods. For the future, we suggest a greater focus on developing model systems that use readily available amphipods and diseases, which can be comparable to the diseases in other Crustacea that are endangered, economically important or difficult to house.

Fluctuating asymmetry, parasitism and reproductive fitness in two species of gammarid crustacean.

Fluctuating asymmetry (FA), defined as random deviations from perfect bilateral symmetry, is assumed to reflect developmental instability. FA is predicted to increase in response to environmental stress, including parasite infection. In addition, based on theory we predict a higher FA in sexually selected traits, due to their greater sensitivity to stress. We investigated the relationships between FA, parasitism and reproductive fitness in 2 species of gammarid crustacean, incorporating both sexual and non-sexual traits. We tested the hypothesis that gammarids infected by vertically transmitted Microsporidia will display higher levels of FA than those infected by horizontally transmitted trematodes, because vertically transmitted Microsporidia can be present at the earliest stages of host development. We found little evidence for a relationship between FA and fecundity in Gammarus spp.; however, egg diameter for infected female Gammarus duebeni was significantly smaller than uninfected female G. duebeni. FA was not correlated with brood size in females or with sperm number in males. In contrast to our prediction, we report a lower relative FA in response to sexual traits than non-sexual traits. However, FA in sexual traits was found to be higher in males than females, supporting the theory that sexual selection leads to increased FA. Additionally, we report a negative correlation between FA and both trematode (Podocotyle atomon) and PCR-positive microsporidian (Nosema granulosis and Dictyocoela duebenum) infections and interpret these results in the context of the parasites' transmission strategies. FA in G. duebeni and G. zaddachi appears to associate with trematode and microsporidian presence, although reproductive fitness is less altered by infection.

Podocotyle atomon (Trematoda: Digenea) impacts reproductive behaviour, survival and physiology in Gammarus zaddachi (Amphipoda).

The Trematoda are a group of phylogenetically diverse metazoan parasites that exhibit complex life cycles that often pass through invertebrate and vertebrate hosts. Some trematodes influence their host's behaviour to benefit transmission. Their parasitic influence may impact host population size by inhibiting an individual's reproductive capacity. We assessed the impact of infection by Podocotyle atomon on the reproductive behaviour and fecundity of its amphipod intermediate host, Gammarus zaddachi, using laboratory and field studies. Parasite prevalence was high in the field, with males more likely to be infected (prevalence in males 64%, in females 39%). Males also suffered a higher parasite burden than females. Infected females were less active, but we found no evidence for a reduction in female reproductive success. Infected females also had comparable pairing success to uninfected females. In males, infection reduced survival and fecundity, with mortality being highest, and sperm numbers lowest, in heavily infected individuals. Trematode parasites are sometimes associated with altered host fecundity, but studies often lack the relevant experimental data to explore the evolution of the trait. We discuss this among information specific to the effect of P. atomon infection in G. zaddachi.