Freshwater mussels in the order Unionida are highly adapted to parasitize fish for the primary purpose of dispersal. The parasitic larval stage affixes itself to the gills or fins of the host where it becomes encysted in the tissue, eventually excysting to develop into a free-living adult. Research on the parasitic interactions between unionids and their host fishes has garnered attention recently due to the increase in worldwide preservation efforts surrounding this highly endangered and ecologically significant order. With the exception of heavy infestation events, these mussels cause minor effects to their hosts, typically only observable effect in combination with other stressors. Moreover, the range of effect intensities on the host varies greatly with the species involved in the interaction, an effect that may arise from different evolutionary strategies between long- and short-infesting mussels; a distinction not typically made in conservation practices. Lower growth and reduced osmotic potential in infested hosts are commonly observed and correlated to infestation load. These effects are typically also associated with increases in metabolic rate and behaviour indicative of stress. Host fish seem to compensate for this through a combination of rapid wound healing in the parasitized areas and higher ventilation rates. The findings are heavily biased towards Margaritifera margaritifera, a unique mussel not well suited for cross-species generalizations. Furthermore, the small body of molecular and genetic studies should be expanded as many conclusions are drawn from studies on the ultimate effects of glochidiosis rather than proximate studies on the underlying mechanisms.

Freshwater mussels in the order Unionida are a highly endangered taxon of ecosystem engineers with a variety of conservation and reintroduction efforts around the world attempting to increase wild populations. The unionid lifecycle involves a parasitic larval life-stage (glochidia), primarily adapted for dispersal rather than growth. Little is known about the behavioural effects of the glochidia on their host fish. We investigated the shoaling behaviour and habitat preference of the Eurasian minnow (Phoxinus phoxinus) while infested with the glochidia of one of Europe’s most endangered unionid mussels, the thick-shelled river mussel (Unio crassus). Behavioural assays were carried out in a hydrologically complex flow-through aquaria at two time points of the infestation period, a carrying stage and an excystment stage (14 and 28 days post infestation). We demonstrate that glochidiosis reduces minnow shoaling frequency and increases the number of isolated individuals. However, minnow shoal area and nearest neighbour distance remained unaffected. Infested minnows demonstrated a significant preference for slower flowing and less turbulent habitat. These results suggest that glochidiosis reduces host swimming efficiency, which likely increases predatory pressure on infested minnows. Simultaneously, this preference for calmer habitats may benefit the encysted mussels, as these habitats are better suited for juvenile mussel excystment. Our results highlight the importance of low-flow and low-turbulence habitats for unionid recruitment, as parasitized host individuals tolerate these conditions well.

Parasitic freshwater mussels are endangered ecosystem engineers with an array of impacts on multiple trophic levels and life stages. While the impacts of adult mussels on separate trophic levels have been studied, few have directly tested how adult mussels can impact trophic interactions, or investigated the impacts of the parasitic mussel larvae (glochidia) on such interactions. We present a laboratory study which mimics two-stream substrates for the endangered thick-shelled river mussel (Unio crassus): one dominated by gravel and one by cobbles. First, the preference of a gammarid (Gammarus pulex) for mussel-dominated habitats was tested in the presence/absence of chemical cues from the predator bullhead (Cottus gobio). Second, the preference of bullhead for mussel-dominated habitats was tested under or without glochidia infestation. Third, the effect of infestation on bullhead predation on gammarids was assessed in the presence of adult mussels. Gammarids only significantly preferred mussel habitats in the absence of predator cues, whereas infested bullhead tended to prefer mussel habitats in cobble substrates. The presence of adult mussels only significantly reduced bullhead predation on gammarids in the gravel habitat, whereas infestation did not affect bullhead predation. Despite gammarids not preferring mussel habitat in the presence of predator cues, mussel beds represent valuable habitat to gammarids as mussel presence can facilitate a reduction of predation by bullhead. Infestation did not affect the rate of bullhead predation on gammarids but did attract bullhead to mussel habitat in cobble substrates. Our results suggest that mussel beds may be valuable habitat for both their host fish and the prey of their hosts, attracting both and increasing predator–prey interactions. This study highlights the cross-trophic mechanisms by which multiple life stages of parasitic mussels can impact the interactions of their surrounding benthic community, underscoring their importance as ecosystem engineers.

Parasites often have a large impact on their hosts and can alter host phenotype to increase their own fitness, a phenomenon known as extended phenotype. Studies demonstrating extended phenotype for non-trophically transmitted parasites are scarce. Unionid mussels have a parasitic life stage adapted to parasitize fish which can affect host behavior, habitat use and growth rates, raising the question if parasitic freshwater mussels can also manipulate their host fish to compensate for downstream dispersal and to reach habitats favorable for newly excysted juvenile mussels. Wild-caught, parasite-na & iuml;ve juvenile brown trout (Salmo trutta) were PIT-tagged, and half of the individuals were infested with parasitic larvae from the freshwater pearl mussel (Margaritifera margaritifera), all individuals were then returned to their home stream. During the following year, trout were tracked to investigate movement and habitat use, and also periodically recaptured to measure growth and body condition factor. The infested trout showed significantly higher upstream movement than non-infested trout and were more often recaptured in stream sections with slow-moving shallow water, particularly during the parasite excystment period (270 d post infestation). These data suggest that the juvenile mussels were successfully transported an average of 170 m upstream from the host trout release points to stream sections favorable for adult mussels. Infested trout survived as well as the non-infested, but had a significantly lower specific growth rate than non-infested trout. These results indicate a first example of extended phenotype in unionid mussels and highlight the importance of understanding glochidia-induced changes to host fish behavioral ecology. Parasites often manipulate host behavior to increase their reproductive success, but studies on this phenomenon typically focus on parasites which lead to the eventual death of their host, an effect that parasitic freshwater mussels generally do not have. Here, we show that parasitic freshwater mussel larvae manipulate host fish into swimming further upriver to habitats well suited for the next stage of their lifecycle.