Freshwater mussels of the Order Unionida provide important ecosystem functions and services, yet many of their populations are in decline. We comprehensively review the status of the 16 currently recognized species in Europe, collating for the first time their life-history traits, distribution, conservation status, habitat preferences, and main threats in order to suggest future management actions. In northern, central, and eastern Europe, a relatively homogeneous species composition is found in most basins. In southern Europe, despite the lower species richness, spatially restricted species make these basins a high conservation priority. Information on freshwater mussels in Europe is unevenly distributed with considerable differences in data quality and quantity among countries and species. To make conservation more effective in the future, we suggest greater international cooperation using standardized protocols and methods to monitor and manage European freshwater mussel diversity. Such an approach will not only help conserve this vulnerable group but also, through the protection of these important organisms, will offer wider benefits to freshwater ecosystems.

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Environmental DNA (eDNA) can be used to detect the presence and abundance of aquatic organisms from water samples. Before implementing this methodology as a tool for monitoring, more knowledge is needed on variation in eDNA concentrations in relation to species abundance and potential confounding factors. Shedding and decay of eDNA may vary extensively over the season and are dependent on environmental factors such as water temperature and on biological processes such as activity level and reproduction. In lotic systems, eDNA concentrations are also affected by downstream transport of eDNA. Sessile freshwater mussels provide an ideal study system for investigating the relationship between species spatial distribution and eDNA concentrations in lotic systems. We quantified freshwater pearl mussel (Margaritifera margaritifera) eDNA concentrations at four localities in a natural river with detailed knowledge of mussel spatial distribution: (a) upstream of the known species distribution, just downstream (b) a small and (c) a large aggregation and (d) 1,700 m downstream of the large aggregation. To study seasonal variation, we quantified eDNA concentrations during three periods: (a) in late spring, with cold water and relatively inactive mussels; (b) in mid-summer, with higher water temperature and active mussel filtration; and (c) in late summer, during the release of larvae.Species detection was highly reliable, with no detection of eDNA upstream of the species distribution and complete detection downstream of the large aggregation.Detection success of the small aggregation was low, with 13% of the samples testing positive. Downstream transport was efficient, with no significant decrease in eDNA concentrations over 1,700 m river distance. Seasonal variation was strong, with a 20-fold increase in eDNA concentrations from late spring to late summer, during reproduction. Our results highlight both the potential and challenges of eDNA monitoring in lotic systems. K E Y W O R D Sconservation genetics, environmental DNA, freshwater mussels, species detection, unionidae | 65 WACKER Et Al.

ABSTRACT1. The highly threatened unionid mussels are obligate parasites on fish. This study investigated larval encapsulation of the freshwater pearl mussel (Margaritifera margaritifera) on its sympatric and three allopatric brown trout (Salmo trutta) strains.2. Encystment abundance differed between the brown trout strains shortly after encapsulation. Encystment abundance then decreased at different rates and resulted in a changed relationship in encystment abundance between the brown trout strains when the experiment was terminated. One of the allopatric brown trout strains had higher encystment abundance than the other brown trout strains.3. The larvae grew at different rates, and the allopatric brown trout strain with the highest encystment abundance had the largest larvae at the end of the experiment. There was a significant positive relationship between the mean condition factor and shell length of the brown trout strains.4. The experiment showed that the potential numbers of juvenile mussels may be restricted at the parasitic life-stage on sympatric brown trout strains. Innate differences in energy resources and immune defence between brown trout strains may be important for parasitic growth, because the condition factor of brown trout strains may be positively related to energy resources for the larvae, and negatively related to host fish immune defence.5. The present experiment showed that it may be important to investigate and manage unionid mussels' parasitic larval stage on host fish. Infestation experiments, like the one presented here, may inform managers if the parasitic stage is functioning properly. They can also evaluate host fish strains used for introductions in streams where natural fish strains have disappeared, but mussels persist. Such experiments have applications in breeding programmes for mussels, as this is an increasing management measure in threatened mussel populations.

Acidification has exterminated or seriously reduced Atlantic salmon ( Salmo salar ) populations in 40–45 Norwegian rivers. As this problem still exists, liming to restore salmon has been necessary, which now involves 21 of these rivers. Thirteen rivers were electrofished 1 year before liming and again 1–12 years later. There was a significant effect both of time after liming and status (e.g., formerly lost and reduced stocks) on the densities of both fry (age 0+) and parr (age ≥ 1+). However, the rate of increase in densities of young salmon in these two status categories was not significantly different in either age group. The development in parr densities suggests that more than 20 years of liming is required to restore salmon in rivers with lost native populations. Stocked rivers and rivers unaltered by hydropower developments generally had higher fry densities and faster increase in parr densities. Annual rod catches of adult salmon increased significantly after liming started, reaching about 45 t after 10 years of treatment. This is 11%–12% of the current total catch of Atlantic salmon in all Norwegian rivers. Liming thus makes an important contribution to the restoration of salmon in formerly acidified rivers.

A total of 21 acidified Norwegian rivers are now being limed to re‐establish or restore Atlantic salmon, Salmo salar L., stocks. Natural reproduction of Atlantic salmon was evident 1 year after the first year of liming in all rivers that had lost their native stocks (n = 9) except for one river. The density of fry (age 0+) developed significantly more rapidly in rivers that supported remnant stocks than in rivers that had lost their stocks, based on data 5 years after treatment. Nine of the study rivers were supplied with hatchery‐reared salmon, mainly unfed fry. Of the rivers with lost stocks, those which were supplied with fish had significantly higher densities than those that were not enhanced. On the other hand, rivers with remnant stocks that were supplied with fish had significantly lower densities of salmon fry than those that did not undergo such mitigation measures. In 2001, all limed rivers yielded 41.9 t of salmon.

Parasites often depend on their hosts for long distance transport, and genetic population structure can be strongly affected by host specificity and dispersal. Freshwater pearl mussel ( Margaritifera margaritifera ) populations have previously been found to naturally infest either primarily Atlantic salmon (‘salmon-mussel’) or exclusively brown trout (‘trout-mussel’) across a wide geographic range. Here, we experimentally test whether this intraspecific variation in natural infestation can be explained by host specificity in freshwater pearl mussel. Our experiments show that when both host species were exposed to larvae from salmon- and trout-mussel respectively, salmon-mussel larvae almost never infested brown trout and vice versa. This suggests that host specificity can explain variation in natural infestation among the studied freshwater pearl mussel populations. Host specificity provides a link to the species’ variable population genetic structure, as mussel populations limited to Atlantic salmon, the host with stronger dispersal, show higher genetic diversity and weaker differentiation than populations limited to brown trout as host.

Multiple paternity is an important characteristic of the genetic mating system and common across a wide range of taxa. Multiple paternity can increase within‐population genotypic diversity, allowing selection to act on a wider spectre of genotypes, and potentially increasing effective population size. While the genetic mating system has been studied in many species with active mating behavior, little is known about multiple paternity in sessile species releasing gametes into the water. In freshwater mussels, males release sperm into the water, while eggs are retained and fertilized inside the female (spermcast mating). Mature parasitic glochidia are released into the water and attach to the gills of fish where they are encapsulated until settling in the bottom substrate. We used 15 microsatellite markers to detect multiple paternity in a wild population of the freshwater pearl mussel (Margaritifera margaritifera). We found multiple paternity in all clutches for which more than two offspring were genotyped, and numbers of sires were extremely high. Thirty‐two sires had contributed to the largest clutch (43 offspring sampled). This study provides the first evidence of multiple paternity in the freshwater pearl mussel, a species that has experienced dramatic declines across Europe. Previous studies on other species of freshwater mussels have detected much lower numbers of sires. Multiple paternity in freshwater pearl mussels may be central for maintaining genetic variability in small and fragmented populations and for their potential to recover after habitat restoration and may also be important in the evolutionary arms race with their fish host with a much shorter generation time.