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Bivalves are ubiquitous members of freshwater ecosystems and responsible for important functions and services. The present paper revises freshwater bivalve diversity, conservation status and threats at the global scale and discusses future research needs and management actions. The diversity patterns are uneven across the globe with hotspots in the interior basin in the United States of America (USA), Central America, Indian subcontinent and Southeast Asia. Freshwater bivalves are affected by multiple threats that vary across the globe; however, pollution and natural system (habitat) modifications being consistently found as the most impacting. Freshwater bivalves are among the most threatened groups in the world with 40% of the species being near threatened, threatened or extinct, and among them the order Unionida is the most endangered. We suggest that global cooperation between scientists, managers, politicians and general public, and application of new technologies (new generation sequencing and remote sensing, among others) will strengthen the quality of studies on the natural history and conservation of freshwater bivalves. Finally, we introduce the articles published in this special issue of Hydrobiologia under the scope of the Second International Meeting on Biology and Conservation of Freshwater Bivalves held in 2015 in Buffalo, New York, USA.

Freshwater mussels are important in aquatic ecosystems but are among the most imperiled species globally. Freshwater pearl mussel (FPM) is a critically threatened species that probably relies on terrestrial organic matter (OM), which suggests that FPM is susceptible to disruptions in terrestrial–aquatic resource connectivity. We performed a regional‐scale field study covering 29 FPM populations across Finland to investigate whether terrestrial ecosystem modifications indirectly affect FPMs by disrupting OM inputs and impairing ecosystem connectivity. We also examined how the resource connectivity covaries along environmental gradients of water quality and catchment characteristics. We used Bayesian mixing models of hydrogen stable isotope ratios from terrestrial and aquatic OM sources to estimate the proportion of terrestrial inputs in FPM as a proxy for resource connectivity. We tested how environmental factors and terrestrial ecosystem alterations influenced connectivity and examined how connectivity and environmental conditions affected eight metrics of individual and population‐level condition in FPM. We found that FPM substantially relied on terrestrial OM across the region, implying strong connectivity. However, environments with higher anthropogenic pressures were associated with lower degrees of connectivity. Reduced connectivity and environmental change translated into lower relative growth and reduced muscle strength in FPM, implying poorer condition status in more human‐impacted areas. These results suggest that human‐induced environmental changes may alter the degree of resource connectivity, which in turn negatively impacts the condition of FPM. These findings highlight the importance of not only conserving ecosystems but also their connections with adjacent ecosystems to maintain the quality and quantity of energy flows which species may depend upon. Our findings also highlight the importance of catchment‐level conservation measures in preserving mussel habitats. Given the status of riverine mussels worldwide, our results are of importance not only to FPM but may inform conservation and management of threatened mussel species more broadly.

Recent research efforts have significantly advanced our knowledge on Asian freshwater mussel (Bivalvia: Unionidae) diversity and distribution. Here we provide a modern consensus of the diversity, biogeography and conservation of Unionida in the region comprising East and Southeast Asia (excluding Wallacea) and Asian Russia. A data review confirmed the presence of 228 native and 3 non-native Unionida (98% Unionidae, 2% Margaritiferidae), rendering the region a global hotspot of freshwater mussel diversity. Species richness was highest in China (particularly Yangtze basin) in absolute numbers and Cambodia when correcting for country area, and decreased gradually towards the south and steeply towards the north and east. Six of the seven unionid subfamilies are native to the region, with species richness peaking in Southeast Asia for Rectidentinae, Gonideinae, Parreysiinae and Modellnaiinae, China for Anodontinae and Unioninae, and Asian Russia for Margaritiferidae. Conservation status and data collected after 1980 were not available for 61 and 24% of species, respectively. Dams, deforestation and pollution are likely the major threats to mussels in the region, though data in this respect are scarce. The Philippines, Laos, Indonesia, Myanmar and Malaysia are among the countries with the poorest data availability and urgently require research.

Invasive freshwater bivalves harm native species, ecosystems and biodiversity, and incur economic costs. The enemy release hypothesis posits that invasive species are released from enemies during the invasion process, giving them a competitive advantage in the new environment. We compared parasitism in two invasive freshwater bivalves, Sinanodonta woodiana and Corbicula fluminea between their original range (China) and invaded range (Europe). For S. woodiana , the average sample‐size‐standardized population‐specific parasite taxon richness was 2.1 times as high, and sum of prevalence was 3.0 times as high in the native range (3 populations, 81 individuals studied) as in the invaded range (6 populations, 210 individuals studied). For C. fluminea , the average standardized population‐specific parasite taxon richness was 1.3 and sum of prevalences was 27.5 in the native range (4 populations, 749 individuals studied), whereas all European C. fluminea were free of parasites (7 populations, 418 individuals studied). The results demonstrate loss of parasites as a result of invasion. Previous studies have shown that parasite pressure on S. woodiana and C. fluminea in the invaded range in Europe is, on average, lower than on sympatric native freshwater mussel populations. Together, these results support one aspect of the enemy release hypothesis: invasive bivalves experience reduced parasite loads as a result of invasion, which may contribute to their success, given the costs typically imposed by parasitism.

Freshwater bivalves have been highly threatened by human activities, and recently their global decline has been causing conservational and social concern. In this paper, we review the most important research events in freshwater bivalve biology calling attention to the main scientific achievements. A great bias exists in the research effort, with much more information available for bivalve species belonging to the Unionida in comparison to other groups. The same is true for the origin of these studies, since the publishing pattern does not always correspond to the hotspots of biodiversity but is concentrated in the northern hemisphere mainly in North America, Europe and Russia, with regions such as Africa and Southeast Asia being quite understudied. We also summarize information about past, present and future perspectives concerning the most important research topics that include taxonomy, systematics, anatomy, physiology, ecology and conservation of freshwater bivalves. Finally, we introduce the articles published in this Hydrobiologia special issue related with the International Meeting on Biology and Conservation of Freshwater Bivalves held in 2012 in Bragança, Portugal.

Freshwater ecosystems are the most vulnerable worldwide and freshwater bivalves rank amongst the most threatened animals in the world. Surveying and monitoring freshwater bivalves are difficult tasks: they are difficult to find, hard to identify (taxonomic expertise is needed), and working underwater is technically challenging. It is therefore crucial to find more efficient methods to survey and monitor these species. Here, we present the first metabarcoding approach for freshwater bivalves and compare environmental DNA (eDNA) and traditional surveys. We describe two sets of primers (for Unionida and Venerida) developed for freshwater bivalves eDNA metabarcoding. These primers have been tested in the field, with about 300 studied sites. Results were compared to freshwater bivalves’ surveys using traditional methods, with eDNA always detecting more species than traditional surveys, especially when Sphaerids were taken into account. While our study initially focused on Western Palearctic freshwater bivalve species, our primers were confronted in silico with available sequences and have proven to be effective at a global scale. The results show that eDNA metabarcoding, with our developed primers, is a remarkable tool allowing for non-invasive surveys, detection of rare and inconspicuous species, absence data and overall freshwater bivalves routine monitoring.

An increasing threat to local, native freshwater mussels (Unionida)—an ecologically important but globally alarmingly declining group—is the invasion by exotic bivalves. The Enemy Release Hypothesis predicts that introduced species should benefit from enemy-mediated competition because they are less likely to be harmed by natural enemies, such as parasites, than their native competitors. We investigated within-site differences in parasitism between sympatric native (tot. five spp.) and invasive (tot. three spp.) bivalves in eight northern European waterbodies, which harboured totally 15 parasite taxa. In paired comparisons using within-site averages, the mean number of parasite species in the native bivalves was 2.3 times higher, and the sum of parasite prevalences 2.4 times higher, than in the invasive bivalves. This may lead to enemy-mediated competitive release of invaders and contribute to the success of invasive freshwater bivalves, in general. However, while the invasive clam Corbicula fluminea was completely free from parasites, parasite parameters of the other invader, Sinanodonta woodiana, were relatively high, indicating that the role of parasites can be invader-specific and urges further research. Understanding the factors affecting success of freshwater bivalve invasions, such as parasitism, can aid invasion control and conservation of local, native (endangered) bivalves.

Abstract Genetic elements encoded in nuclear DNA determine the sex of an individual in many animals. In certain bivalve lineages that possess doubly uniparental inheritance (DUI), mitochondrial DNA (mtDNA) has been hypothesized to contribute to sex determination. In these cases, females transmit a female mtDNA to all offspring, while male mtDNA (M mtDNA) is transmitted only from fathers to sons. Because M mtDNA is inherited in the same way as Y chromosomes, it has been hypothesized that mtDNA may be responsible for sex determination. However, the role of mitochondrial and nuclear genes in sex determination has yet to be validated in DUI bivalves. In this study, we used DNA, RNA, and mitochondrial short noncoding RNA (sncRNA) sequencing to explore the role of mitochondrial and nuclear elements in the sexual development pathway of the freshwater mussel Potamilus streckersoni (Bivalvia: Unionida). We found that the M mtDNA sheds a sncRNA partially within a male-specific mitochondrial gene that targets a pathway hypothesized to be involved in female development and mitophagy. RNA-seq confirmed the gene target was significantly upregulated in females, supporting a direct role of mitochondrial sncRNAs in gene silencing. These findings support the hypothesis that M mtDNA inhibits female development. Genome-wide patterns of genetic differentiation and heterozygosity did not support a nuclear sex-determining region, although we cannot reject that nuclear factors are involved with sex determination. Our results provide further evidence that mitochondrial loci contribute to diverse, nonrespiratory functions and additional insights into an unorthodox sex-determining system.

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 with 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.

This work was supported by Portuguese Foundation for Science and Technology (FCT) [grant number SFRH/BD/115728/2016 (MLL), grant number SFRH/BD/137935/2018 (AGS)]; Russian Foundation for Basic Research [grant number 18-34-20033 (IVV)]; Dawson Fellowship at St. Catharine's College, Cambridge (DCA); Life Margal Ulla [number LIFE09 NAT/ES/000514 (RA and PO)]; COMPETE 2020, Portugal 2020 and the European Union through the ERDF, and by Portuguese Foundation for Science and Technology (FCT) through national funds [UID/Multi/04423/2019] under project ConBiomics: the missing approach for the Conservation of freshwater Bivalves Project, and [project number NORTE-01-0145-FEDER-030286]; Federal Agency for Scientific Organizations under Grants [grant number 0409-2015-0143 (INB and IVV)]