Facilitative and competitive interactions among coexisting parasite species, as well as among alternative host species, produce considerable ecological and evolutionary responses to host-parasite relationships. Such effects can be illuminated by impacts of non-native species on relationships in local communities. We used the association between parasitic European bitterling fish (Rhodeus amarus) and unionid mussels (which host bitterling eggs in their gills) to test the effects of the invasive Chinese pond mussel (Sinanodonta woodiana) and the presence of non-bitterling mussel macroparasites on the pattern of host mussel use by the bitterling across 12 unionid mussel communities with the absence or presence of S. woodiana (and variation in duration of coexistence with local species). While all six European mussel species were used by the bitterling (with the prevalence of > 30% in Unio spp.), no S. woodiana individual was infected by the bitterling. The presence of S. woodiana did not affect bitterling eggs distribution in native mussels. Large native mussels hosted more bitterling. Infection by non-bitterling parasites, mostly water mites (prevalence 47%) and trematodes (25%), did not affect rates of bitterling parasitism. We discuss our results in the context of the rapid evolution of non-native species in their new range and its implication on mussel conservation.

• Keywords
• Freshwater mussel, Host selection, Host-parasite interactions, Invasive species, Mussel parasitism, Parasite facilitation,
• MeSH
• Host-Parasite Interactions * MeSH
• Bivalvia * parasitology   physiology   MeSH
• Ovum * parasitology   MeSH
• Introduced Species * MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH

Embryonic development is often considered shielded from the effects of natural selection, being selected primarily for reliable development. However, embryos sometimes represent virulent parasites, triggering a coevolutionary "arms race" with their host. We have examined embryonic adaptations to a parasitic lifestyle in the bitterling fish. Bitterlings are brood parasites that lay their eggs in the gill chamber of host mussels. Bitterling eggs and embryos have adaptations to resist being flushed out by the mussel. These include a pair of projections from the yolk sac that act as an anchor. Furthermore, bitterling eggs all adopt a head-down position in the mussel gills which further increases their chances of survival. To examine these adaptations in detail, we have studied development in the rosy bitterling (Rhodeus ocellatus) using molecular markers, X-ray tomography, and time-lapse imaging. We describe a suite of developmental adaptations to brood parasitism in this species. We show that the mechanism underlying these adaptions is a modified pattern of blastokinesis-a process unique, among fish, to bitterlings. Tissue movements during blastokinesis cause the embryo to do an extraordinary "front-flip" on the yolk. We suggest that this movement determines the spatial orientation of the other developmental adaptations to parasitism, ensuring that they are optimally positioned to help resist the ejection of the embryo from the mussel. Our study supports the notion that natural selection can drive the evolution of a suite of adaptations, both embryonic and extra-embryonic, via modifications in early development.

• Keywords
• Eco-Evo-Devo, egg shape, gastrulation, hatching, parasitism,
• MeSH
• Cyprinidae * MeSH
• Host-Parasite Interactions MeSH
• Parasites * MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH

Parasitic infections elicit host defences that pose energetic trade-offs with other fitness-related traits. Bitterling fishes and unionid mussels are involved in a two-way parasitic interaction. Bitterling exploit mussels by ovipositing into their gills. In turn, mussel larvae (glochidia) develop on the epidermis and gills of fish. Hosts have evolved behavioural responses to reduce parasite load, suggesting that glochidia and bitterling parasitism are costly. We examined the energetic cost of parasitism on both sides of this relationship. We used intermittent flow-through respirometry to measure (1) standard metabolic rate (SMR) of individual duck mussels Anodonta anatina (a common bitterling host) before and during infection by embryos of the European bitterling Rhodeus amarus, and (2) SMR and maximum oxygen uptake (MO2max) of individual R. amarus before and during infection with glochidia of the Chinese pond mussel Sinanodonta woodiana (a mussel species that successfully infects bitterling). As predicted, we observed an increase in mussel SMR when infected by bitterling embryos and an increased SMR in glochidia-infected bitterling, though this was significantly mediated by the time post-infection. Contrary to our predictions, glochidia infection did not impair MO2max and the number of glochidia attached to gills positively (rather than negatively) correlated with MO2max. The results suggest that tolerance is the prevailing coping mechanism for both fish and mussels when infected, while resistance mechanisms appear to be confined to the behavioural level.

• Keywords
• Acheilognathinae, Coevolution, Evolutionary arms race, Host–parasite relationship, Unionidae,
• MeSH
• Cyprinidae * MeSH
• Host-Parasite Interactions MeSH
• Oxygen MeSH
• Parasites * MeSH
• Oxygen Consumption MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH
• Names of Substances
• Oxygen MeSH

Generalist parasites have the capacity to infect multiple hosts. The temporal pattern of host specificity by generalist parasites is rarely studied, but is critical to understanding what variables underpin infection and thereby the impact of parasites on host species and the way they impose selection on hosts. Here, the temporal dynamics of infection of four species of freshwater mussel by European bitterling fish (Rhodeus amarus) was investigated over three spawning seasons. Bitterling lay their eggs in the gills of freshwater mussels, which suffer reduced growth, oxygen stress, gill damage and elevated mortality as a result of parasitism. The temporal pattern of infection of mussels by European bitterling in multiple populations was examined. Using a Bernoulli Generalized Additive Mixed Model with Bayesian inference it was demonstrated that one mussel species, Unio pictorum, was exploited over the entire bitterling spawning season. As the season progressed, bitterling showed a preference for other mussel species, which were inferior hosts. Temporal changes in host use reflected elevated density-dependent mortality in preferred hosts that were already infected. Plasticity in host specificity by bitterling conformed with the predictions of the host selection hypothesis. The relationship between bitterling and their host mussels differs qualitatively from that of avian brood parasites.

• Keywords
• Brood parasite, Host–parasite co-evolution, Oviposition, Spawning site, Superparasitism,
• MeSH
• Bayes Theorem MeSH
• Cyprinidae MeSH
• Host Specificity * MeSH
• Host-Parasite Interactions * MeSH
• Parasites MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH

Predicting the impacts of non-native species remains a challenge. As populations of a species are genetically and phenotypically variable, the impact of non-native species on local taxa could crucially depend on population-specific traits and adaptations of both native and non-native species. Bitterling fishes are brood parasites of unionid mussels and unionid mussels produce larvae that parasitize fishes. We used common garden experiments to measure three key elements in the bitterling-mussel association among two populations of an invasive mussel (Anodonta woodiana) and four populations of European bitterling (Rhodeus amarus). The impact of the invasive mussel varied between geographically distinct R. amarus lineages and between local populations within lineages. The capacity of parasitic larvae of the invasive mussel to exploit R. amarus was higher in a Danubian than in a Baltic R. amarus lineage and in allopatric than in sympatric R. amarus populations. Maladaptive oviposition by R. amarus into A. woodiana varied among populations, with significant population-specific consequences for R. amarus recruitment. We suggest that variation in coevolutionary states may predispose different populations to divergent responses. Given that coevolutionary relationships are ubiquitous, population-specific attributes of invasive and native populations may play a critical role in the outcome of invasion. We argue for a shift from a species-centred to population-centred perspective of the impacts of invasions.

• Keywords
• Anodonta woodiana, alien species, glochidia, host–parasite dynamics, intraspecific variation, symbiosis,
• MeSH
• Anodonta genetics   growth & development   physiology   MeSH
• Cyprinidae parasitology   physiology   MeSH
• Host-Parasite Interactions MeSH
• Oviposition * MeSH
• Larva genetics   growth & development   physiology   MeSH
• Reproduction MeSH
• Avoidance Learning * MeSH
• Introduced Species * MeSH
• Animals MeSH
• Check Tag
• Female MeSH
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH
• Geographicals
• Europe MeSH