a b s t r a c t Freshwater mussels of the order Unionida are key elements of freshwater habitats and are responsible for important ecological functions and services. Unfortunately, these bivalves are among the most threatened freshwater taxa in the world. However, conservation planning and management are hindered by taxonomic problems and a lack of detailed ecological data. This highlights the urgent need for advances in the areas of systematics and evolutionary relationships within the Unionida. This study presents the most comprehensive phylogeny to date of the larger Unionida family, i.e., the Unionidae. The phylogeny is based on a combined dataset of 1032 bp (COI + 28S) of 70 species in 46 genera, with 7 of this genera being sequenced for the first time. The resulting phylogeny divided the Unionidae into 6 supported subfamilies and 18 tribes, three of which are here named for the first time (i.e., Chamberlainiini nomen novum, Cristariini nomen novum and Lanceolariini nomen novum). Molecular analyses were complemented by investigations of selected morphological, anatomical and behavioral characters used in traditional phylogenetic studies. No single morphological, anatomical or behavioral character was diagnostic at the subfamily level and few were useful at the tribe level. However, within subfamilies, many tribes can be recognized based on a subset of these characters. The geographical distribution of each of the subfamilies and tribes is also presented. The present study provides important advances in the systematics of these extraordinary taxa with implications for future ecological and conservation studies.

Many doubts still exist about which freshwater mussel Unio species inhabit Northwest Africa. While some authors refer to the presence of Unio delphinus in the Atlantic North African basins of Morocco, a recent International Union for Conservation of Nature (IUCN) assessment performed on Moroccan Unio species, recognised the existence of a distinct species, Unio foucauldianus, with a critically endangered conservation status. The present study delivered new genetic, morphological, and geographical distribution data on two Unio species (i.e. U. delphinus and U. foucauldianus) greatly increasing the almost non-existent data on these taxa. Bayesian phylogenetic analysis revealed two highly supported geographically concordant clades, which diverged by 3.2 ± 0.6 % (uncorrected p distance): the first distributed across Iberia and corresponding to U. delphinus, and the second distributed across Morocco, corresponding to U. foucauldianus. These results were corroborated by the analysis of ten newly developed microsatellite loci as well as shell morphometry. We suggest that the IUCN critically endangered conservation status of U. foucauldianus should be revised and probably down-listed since its actual distribution is much wider than previously described. Phylogenetic relationships with the other Unio species were resolved, showing that U. delphinus and U. foucauldianus fall inside the pictorum lineage. The estimated molecular rate reported herein (0.265 ± 0.06 % per million years) represents the first for the Unionida and could be used as a reference in future studies.Electronic supplementary material The online version of this article

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Margaritifera marocana is one of the rarest freshwater mussel species (listed as critically endangered), and is endemic to Morocco. Despite its constrained distribution and low abundance, to date there are no quantitative studies addressing the conservation status of this species. Surveys were conducted in 36 sites along the Oum Er Rbia river basin (Rivers Derna, Laabid and Oum Er Rbia) to assess the distribution, abundance, population structure and genetic diversity of M. marocana. Just one specimen was found on River Oum Er Rbia and none on River Derna; however, a high abundance was found in the lower section of River Laabid (e.g., site Laabid 6 reached a mean density of 11.0 ± 6.8 ind.m(-2)). Contrary to earlier information, which reported an overall population size fewer than 250 individuals in a restricted area and no juvenile presence, this study showed that a much higher abundance exists in River Laabid alone. In addition, the species is present in more than 50 km of this river and is still recruiting since small specimens were found. Regarding genetic diversity, six of nine loci previously used in Margaritifera margaritifera were polymorphic and suitable in M. marocana. The spatial range contraction of this species is likely to be very recent, since no strong signature was detected by the molecular diversity indices. Information gathered in this study can be used as a reference to the present conservation status of M. marocana, and guide future research and management initiatives to better conserve it. We conclude discussing the potential major threats for the future survival of M. marocana and suggest some management measures (and research needs) that should be urgently applied.

With the biodiversity crisis continuing unchecked, we need to establish levels and drivers of extinction risk to effectively allocate conservation resources and develop targeted actions. Given that threat appears particularly high in freshwaters, we assessed the extinction risk of 1,500 randomly selected freshwater molluscs using the IUCN Red List Categories and Criteria, as part of the Sampled Red List Index project. We show that close to one third of species in our sample are estimated to be threatened with extinction, with highest levels of threat in the Palearctic, Australasia and Nearctic and among gastropods. Threat levels were higher in lotic than lentic systems. Twenty-seven species were classified as Extinct (eight bivalves and 19 gastropods), mostly from the Nearctic realm and lotic systems. Pollution and the modification of natural systems (e.g. through damming and water abstraction) were the most frequently reported threats to freshwater molluscs, with some regional variation. Given that we found little spatial congruence between species richness patterns of freshwater molluscs and other freshwater taxa, new additional conservation priority areas emerge from our study. We discuss the implications of our findings for freshwater mollusc conservation and important next steps to estimate trends in freshwater mollusc extinction risk over time.