Freshwater mussels are declining throughout their range. Their important ecological functions along with insufficient levels of natural recruitment have prompted captive breeding for population augmentation and questions about the usefulness and applicability of such measures. This article reviews the current state of captive breeding and rearing programmes for freshwater mussels in Europe. It considers the various species, strategies, and techniques of propagation, as well as the different levels of effort required according to rearing method, highlighting the key factors of success. Within the last 30 years, 46 breeding activities in 16 European countries have been reported, mainly of Margaritifera margaritifera and Unio crassus. Some facilities propagate species that are in a very critical situation, such as Pseudunio auricularius, Unio mancus, and Unio ravoisieri, or multiple species concurrently. In some streams, the number of released captive-bred mussels already exceeds the size of the remaining natural population. Rearing efforts range from highly intensive laboratory incubation to lower intensity methods using in-river mussel cages or silos. Most breeding efforts are funded by national and EU LIFE(+) grants, are well documented, and consider the genetic integrity of the propagated mussels. Limited long-term funding perspectives, the availability of experienced staff, water quality, and feeding/survival during early life stages are seen as the most important challenges. Successful captive breeding programmes need to be combined with restoration of the habitats into which the mussels are released. This work will benefit from an evidence-based approach, knowledge exchange among facilities, and an overall breeding strategy comprising multiple countries and conservation units.

Management decisions in conservation of threatened species require trading off social needs against biodiversity values, including the prioritization of conservation locations, i.e. where conservation efforts should take place. To improve conservation decisions for the thick-shelled river mussel, Unio crassus, a highly threatened temporary parasite on fish, we performed a field study on how mussel and host fish density (European bullhead, Cottus gobio, and common minnow, Phoxinus phoxinus) affect reproduction potential of the mussel at different sites along a river. We assumed that the proportions of gravid mussels would be higher at high mussel density, and result in enhanced glochidia (mussel larvae) encapsulation rates on fish. We also expected the highest ‘glochidia density’—a proxy for the potential number of recruits per stream area, assessed by multiplying glochidia encapsulation rates on fish by fish density, to occur at high mussel density sites. Such river sites, producing many offspring and conveying important conservation values, may thus be prioritized. However, contrary to our assumptions, higher glochidia density and higher proportions of gravid mussels occurred at lower density mussel sites. We also found that P. phoxinus had higher glochidia encapsulation rates than C. gobio, possibly related to species-specific behavioural and life-history traits. Even so, glochidia density was similar for both fish species, reflecting comparable ecological functions in hosts. The results of this study suggest that mussel and host fish densities should be considered along with glochidia density in conservation prioritization and management trade-offs.

Adaptation to temperature regimes and host presence may enhance fitness in parasites. In an experimental study, we evaluated the timing of glochidia release by Unio crassus subjected to three spring water temperature regimes in the presence and absence of the host fish Cottus gobio. The timing of glochidia release was delayed at (i) constantly low temperatures (<10°C), in contrast to earlier and pronounced releases at (ii) natural temperature increases that level off at intermediate temperatures (10–15°C), and (iii) higher-than-normal temperatures (10–20°C). Mussels from treatment (i) that had not released glochidia during the experiment did so soon after being moved to the temperature in (ii), indicating a temperature threshold for glochidia release. Neither host fish presence nor the combined effect of temperature and host fish presence significantly affected the timing of glochidia release. The treatment with natural spring water temperatures indicated possible fitness benefits for U. crassus through combined effects of high intensities of glochidia releases and high survival of released glochidia. The furthered understanding of climate change effects on mussel and host phenology in seasonal environments, potentially inducing temporal mismatches of glochidia release to host availability, is key to mussel conservation

Unionoid mussels are globally threatened and their conservation requires species-specific knowledge on their ecology and parasite-host interaction. Unio crassus is one of Europe’s most threatened unionoid species and has a temporary obligate parasitic life stage (glochidia) on fish. A lack of suitable hosts is probably a major limitation for mussel recruitment, but host species composition, suitability and availability in time and space have yet to be fully explored. This thesis examines different aspects of the host fish species, including their composition, suitability and ecological importance, in relation to U. crassus, using both field and laboratory studies. The effects of mussel and host density on mussel reproductive potential were considered, as were aspects of evolutionary adaptations between mussels and fish and how climate change may affect their interaction.

The results show that U. crassus is a host generalist, parasitizing a variety of fish species. Host suitability and density, which varied among fish species and rivers, affected the level of glochidia encapsulation, hence mussel reproductive potential, more so than the density of mussels taking part in reproduction. Ecologically important hosts included both highly suitable primary hosts, and less suitable hosts that were highly abundant. Whether or not U. crassus has specific adaptations to its hosts to enhance juvenile transformation remains unclear. No distinct pattern of local adaptation was found, nor was there an effect of host fish presence on the timing of glochidia release by adult mussels. Instead, temperature played a major role, with results suggesting that changes in spring water temperature regimes can cause temporal and spatial mismatches in the mussel-host interaction. This thesis indicates that investigations of local mussel-host interactions help in identifying mechanisms important for unionoid conservation management and prioritization.

Många sötvattenmusslor har en komplex livscykel där larverna (glochidier) under sin utveckling till frilevande musslor parasiterar på gälarna hos lämpliga värdfiskar. Flera av våra musslor, såsom den tjockskaliga målarmusslan (Unio crassus), är globalt hotade och för att kunna bevara och förvalta dessa arter på bästa sätt behöver vi lära oss mer om deras ekologi och samspelet mellan musslan och dess värdfiskar. Avsaknaden av värdfiskar innebär förmodligen en stor begränsning för rekryteringen av juvenila musslor, men det finns trots detta en begränsad kunskap om hur artsammansättningen i fisksamhället och dess tillgänglighet påverkar musselpopulationer. 

Min avhandling undersöker olika aspekter av interaktioner mellan U. crassus och dess värdfiskar, som hur värdfisksamhällen och fiskarters värdlämplighet påverkar musslans reproduktionspotential. Jag har även studerat hur tätheter av olika fiskarter och vuxna musslor påverkar rekryteringen, eventuella evolutionära anpassningar samt om en förhöjd temperatur skulle kunna påverka interaktionen mellan U. crassus och dess värdfiskar.

Resultaten visar att U. crassus är en generalist som parasiterar på en mängd olika fiskarter. Jag fann dock en stor variation i dominerande fiskarter och lämpliga värdar mellan olika åar, vilket påverkade reproduktionspotentialen hos musslorna mer än vad tätheten vuxna musslor som deltog i reproduktionen gjorde. Som ekologiskt viktiga värdar fanns således både särskilt lämpliga, primära värdarter, men också mindre lämpliga arter som förekom i höga tätheter. Ingen tydlig lokal anpassning kunde observeras, och fiskens närvaro påverkade inte tidpunkten för när de vuxna musslorna släppte sina glochidielarver. Däremot fann jag att temperaturen spelade en viktig roll för musslans reproduktion, där ökad temperatur föreslås ha negativa effekter på interaktionen mellan musslan och dess värdfiskar.

Avhandlingen visar på vikten av att studera interaktioner mellan den tjockskaliga målarmusslan och dess värdar på lokal skala för att bättre kunna identifiera och prioritera viktiga naturvårdsåtgärder.

1.Parasite–host interactions can involve strong reciprocal selection pressure, and may lead to locally adapted specializations. The highly threatened unionoid mussels are temporary parasites on fish, but local adaptation has not yet been investigated for many species. 2.Patterns of local adaptation of one of Europe's most threatened unionoids, the thick‐shelled river mussel (Unio crassus) were investigated. Eurasian minnows (Phoxinus phoxinus) from two rivers (separate drainage areas) were cross‐infested in the laboratory with sympatric and allopatric mussel larvae, while bullheads (Cottus gobio), inhabiting only one of the rivers, were infested with sympatric or allopatric mussel larvae. Larval encystment, juvenile mussel excystment and survival were measured. 3.For one river, but not the other, juvenile excystment from P. phoxinus was highest when infested with sympatric mussels. The opposite pattern was found for C. gobio in this river, where juvenile excystment and post‐parasitic juvenile survival from allopatric C. gobio were highest. The results thus cannot confirm local adaptation of U. crassus to P. phoxinus in the study rivers, as excystment was not consistently higher in all sympatric mussel–host combinations, whereas there were potential maladaptive signs of U. crassus in relation to C. gobio. There was no loss of encysted larvae 3 days after infestation until juvenile excystment. Most juveniles were excysted between 17 and 29 days after infestation, and the numbers of excysted juveniles increased with fish size. 4.The results have implications for parasite–host ecology and conservation management with regard to unionoid propagation and re‐introduction. This includes the need to (1) test suitability and adaptation patterns between U. crassus and multiple host fish species, (2) evaluate the suitability of certain unionoids and host fish strains after more than 3 days, and (3) determine whether large fish produce more juvenile mussels than smaller fish

Surface ice in rivers and lakes buffers the thermal environment and provides overhead cover, protecting aquatic animals from terrestrial predators. We tested if surface ice influenced the behavior (swimming activity, aggressive encounters, and number of food items eaten) and stress level (coloration of eyes and body) of stream-living brown trout Salmo trutta at temperatures of 3–4 °C in indoor experimental flumes. We hypothesized that an individual’s resting metabolic rate (RMR, as measured by resting ventilation rate) would affect winter behavior. Therefore, groups of 4 trout, consisting of individuals with high, low, or mixed (2 individuals each) RMR, were exposed to experimental conditions with or without ice cover. Ice cover reduced stress responses, as evaluated by body coloration. Also, trout in low RMR groups had a paler body color than those in both mixed and high RMR groups. Trout increased their swimming activity under ice cover, with the highest activity found in high RMR groups. Ice cover increased the number of aggressive encounters but did not influence the number of drifting food items taken by each group. In mixed RMR groups, however, single individuals were better able to monopolize food than in the other groups. As the presence of surface ice increases the activity level and reduces stress in stream-living trout, ice cover should influence their energy budgets and production. The results should be viewed in light of ongoing global warming that reduces the duration of ice cover, especially at high latitudes and altitudes.

Nature-like fishways are designed to imitate the characteristics of natural streams, thereby providing both fish passage and habitat for a variety of aquatic organisms. To date, however, the potential for habitat rehabilitation of nature-like fishways has not been fully realized. To develop the concept of how to design a nature-like fishway, a 500-m long nature-like fishway, termed the biocanal, was constructed at the Eldforsen hydroelectric facility, Sweden. It included four habitat types: riffle, pool, floodplain and braided (i.e. with islands), each replicated three times. The riffle sections were considered controls for typical Swedish nature-like fishways. Thus the biocanal had a more varied in-stream environment than those of conventional fishways. To test the prediction that the biocanal had a positive effect on biodiversity, we compared the physical habitat and benthic fauna composition of the more diverse habitat types in the biocanal to the riffle habitats. We also made comparisons between the biocanal and six natural reference streams in the area. After two years, 63% of the benthic fauna families found in the reference streams had colonized the biocanal. Families present in the reference streams, but not in the biocanal, were predominantly slow colonizers or taxa linked to riparian vegetation, which was scarce and in an early successional stage along the biocanal. In the biocanal, pool and floodplain habitats contained the highest number of families, the highest family diversity (Shannon-Weaver) and the highest densities of Ephemeroptera, Plecoptera and Trichoptera. Since these habitats contained more families and had higher diversities than the riffle habitats which are typical of conventional nature-like fishways, we suggest that the construction of biocanals indeed possesses the potential for high biodiversity. (C) 2013 Elsevier B.V. All rights reserved.