Declining freshwater mussel diversity in the middle and lower reaches of the Xin River Basin: Threat and conservation

Freshwater mussels provide important functions and services for aquatic ecosystems, but populations of many species have been extirpated. Information on biodiversity plays an important role in the conservation and management of freshwater mussels. The Xin River Basin is a biodiversity hotspot for freshwater mussels in China, with more than 43 species known, but populations of which are decreasing. Here, we quantify the diversity of freshwater mussels in the middle and lower reaches of the Xin River Basin and study the correlation of habitat characteristics and freshwater mussel diversity. Compared to the historical period, the number of species, density, and biomass of freshwater mussels decreased 33%, 83%, and 82% in the current period, respectively. Fifty two percent of recorded species were empty shells, and 14 native freshwater mussels were not found in the study area. Four species are currently listed as vulnerable species using IUCN criteria and their global status. The assemblage structure of freshwater mussels exhibits significant spatial differences, and there was a correlation with substrate and physicochemical parameters. The main tributary of the Xin River with higher freshwater mussel diversity should be established as one large protected area because the nestedness component was the main pattern of beta diversity. These results indicated freshwater mussel diversity was declining rapidly, which can help focus conservation effort for freshwater mussel biodiversity.     

Strategies for the conservation of endangered freshwater pearl mussels (<Emphasis Type="Italic">Margaritifera margaritifera</Emphasis> L.): a synthesis of Conservation Genetics and Ecology

Freshwater pearl mussels (Margartifera margaritifera L.) are among the most critically threatened freshwater bivalves worldwide. The pearl mussel simultaneously fulfils criteria of indicator, flagship, keystone and umbrella species and can thus be considered an ideal target species for the process conservation of aquatic ecosystem functioning. The development of conservation strategies for freshwater pearl mussels and for other bivalve species faces many challenges, including the selection of priority populations for conservation and strategic decisions on habitat restoration and/or captive breeding. This article summarises the current information about the species’ systematics and phylogeny, its distribution and status as well as about its life history strategy and genetic population structure. Based on this information, integrative conservation strategies for freshwater mollusc species which combine genetic and ecological information are discussed. Holistic conservation strategies for pearl mussels require the integration of Conservation Genetics and Conservation Ecology actions at various spatial scales, from the individual and population level to global biodiversity conservation strategies. The availability of high resolution genetic markers for the species and the knowledge of the critical stages in the life cycle, particularly of the most sensitive post-parasitic phase, are important prerequisites for conservation. Effective adaptive conservation management also requires an evaluation of previous actions and management decisions. As with other freshwater bivalves, an integrative conservation approach that identifies and sustains ecological processes and evolutionary lineages is urgently needed to protect and manage freshwater pearl mussel diversity. Such research is important for the conservation of free-living populations, as well as for artificial culturing and breeding techniques, which have recently been or which are currently being established for freshwater pearl mussels in several countries.     

Conserving freshwater biodiversity: The value,status and management of high quality ditch systems

Freshwater biodiversity is globally threatened and while most conservation efforts are focussed on natural and larger freshwater systems such as rivers and lakes, in many lowland agricultural landscapes artificial water bodies including ditches may be equally important as habitats for freshwater species. Ditches occur across the agricultural landscape but in particular, those associated with coastal and floodplain grazing marsh, have high conservation value. The importance of this habitat for rare and threatened species afforded priority status under the UK Biodiversity Action Plan is explored. The characteristics of ditches that have high conservation value are described and a set of targets against which such ditches can be assessed are presented. An analysis of the current condition of Sites of Special Scientific Interest (SSSI) for the wider coastal and floodplain grazing marsh habitat demonstrates the range of pressures affecting these sites and highlights that alongside generic freshwater issues such as eutrophication and non-native species, these sites have a unique set of pressures associated with their ongoing management and the vulnerable location of many sites at the coast. Wider conservation strategies for freshwater biodiversity in lowland landscapes across Europe need to factor in the different management requirements of artificial habitats such as ditches alongside more ambitious restoration projects for natural waterbodies. In low lying coastal areas the threat of coastal squeeze for many important grazing marshes will require a strategic approach to allow upstream migration of important biodiversity.     

Analysing freshwater fish biodiversity records and respective conservation areas in Spain

The number of threatened freshwater fish species in Spain is among the highest recorded in Europe and includes a high percentage of endemic taxa. Investigated were the distribution of Spanish freshwater fish to identify priority areas for conservation and assess the extent to which freshwater fish are included in the existing network of protected areas. Considered were those threatened species recorded in the Spanish National inventories. From these data, several biodiversity indices were calculated and analysed. The results reveal important discrepancies between the national and international assessments of conservation status, whereby the current Spanish national catalogue needs updating to reconcile these inconsistencies. Several important areas for the conservation of freshwater fish lie outside protected areas. The results encourage the establishment of protected areas specifically for freshwater environments. An extensive database of Spanish freshwater fish species is needed to redefine priority areas and to maintain freshwater biodiversity.     

Establishing conservation units to promote recovery of two threatened freshwater mussel species (Bivalvia: Unionida: Potamilus)

Population genomics has significantly increased our ability to make inferences about microevolutionary processes and demographic histories, which have the potential to improve protection and recovery of imperiled species. Freshwater mussels (Bivalvia: Unionida) represent one of the most imperiled groups of organisms globally. Despite systemic decline of mussel abundance and diversity, studies evaluating spatiotemporal changes in distribution, demographic histories, and ecological factors that threaten long‐term persistence of imperiled species remain lacking. In this study, we use genotype‐by‐sequencing (GBS) and mitochondrial sequence data (mtDNA) to define conservation units (CUs) for two highly imperiled freshwater mussel species, Potamilus amphichaenus and Potamilus streckersoni. We then synthesize our molecular findings with details from field collections spanning from 1901 to 2019 to further elucidate distributional trends, contemporary status, and other factors that may be contributing to population declines for our focal species. We collected GBS and mtDNA data for individuals of P. amphichaenus and P. streckersoni from freshwater mussel collections in the Brazos, Neches, Sabine, and Trinity drainages ranging from 2012 to 2019. Molecular analyses resolved disputing number of genetic clusters within P. amphichaenus and P. streckersoni; however, we find defensible support for four CUs, each corresponding to an independent river basin. Evaluations of historical and recent occurrence data illuminated a generally increasing trend of occurrence in each of the four CUs, which were correlated with recent increases in sampling effort. Taken together, these findings suggest that P. amphichaenus and P. streckersoni are likely rare throughout their respective ranges. Because of this, the establishment of CUs will facilitate evidence‐based recovery planning and ensure potential captive propagation and translocation efforts are beneficial. Our synthesis represents a case study for conservation genomic assessments in freshwater mussels and provides a model for future studies aimed at recovery planning for these highly imperiled organisms.     

Species traits and catchment‐scale habitat factors influence the occurrence of freshwater mussel populations and assemblages

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Lifting the curtain on the freshwater mussel diversity of the Italian Peninsula and Croatian Adriatic coast

Freshwater mussels of the order Unionida have been dramatically declining globally. Despite their ecological importance, conservation of these animals has been hindered by unresolved taxonomy and a lack of data on the distribution and status of populations, especially in southern Europe. Although the Italian Peninsula has been noted as a centre of endemism and one of the major refugia of the glacial ages for several taxa, few studies have been performed on the genetic diversity of Unionida. Most importantly, the taxonomic status of several freshwater mussel populations of the Italian Peninsula is still unresolved. Here we present the first comprehensive dataset for the Unionida of the region spanning Italy and the coastal Croatian region (west of the Dinaric Alps). In total, 191 specimens were collected (85 Anodonta, 64 Unio, 17 Microcondylaea bonellii and 25 Sinanodonta woodiana) from 34 sites across the Italian Peninsula and coastal Croatian river basins for molecular identification (COI, 16S and 28S). Genetic analyses were performed to understand major phylogenetic and phylogeographic patterns. Seven species were detected: three Anodonta species (A. anatina, A. cygnea and A. exulcerata), two Unio species (U. mancus and U. elongatulus), Microcondylaea bonellii, and the invasive Sinanodonta woodiana. The presence of three endemic species (A. exulcerata, U. elongatulus and M. bonellii) confirms the importance of the region as a centre of endemism for freshwater mussels. The Apennine Mountains act as an important biogeographic barrier.     

High levels of multiple paternity in a spermcast mating freshwater mussel

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.     

Population genetic structure of three freshwater mussel (Unionidae) species within a small stream system: significant variation at local spatial scales

1. Unionid mussels are highly threatened, but little is known about genetic structure in populations of these organisms. We used allozyme electrophoresis to examine partitioning of genetic variation in three locally abundant and widely distributed species of mussels from a catchment in Ohio. 2. Within‐population variation was similar to that previously reported for freshwater mussels, but genotype frequencies exhibited heterozygote deficiencies in many instances. All three species exhibited significant among‐population variation. Evidence of isolation‐by‐distance was found in Elliptio dilatata and Ptychobranchus fasciolaris, while Lampsilis siliquoidea showed no geographical pattern of among‐population variation. 3. Our results suggest that the isolating effects of genetic drift were greater in L. siliquoidea than in the other species. Differentiation of populations occurred at a much smaller spatial scale than has previously been found in freshwater mussels. Differences among species may reflect differences in the dispersal abilities of fishes that serve as hosts for the glochidia larvae of mussels. 4. Based on our results, we hypothesise that species of mussels that are common to large rivers exhibit relatively large amounts of within‐population genetic variation and little differentiation over large geographical distances. Conversely, species typical of small streams show lower within‐population genetic variation and populations will be more isolated. If this hypothesis can be supported, it may prove useful in the design of conservation strategies that maintain the genetic structure of target species.     

Three deeply divided lineages of the freshwater mussel genus <Emphasis Type="Italic">Anodonta</Emphasis> in western North America

The surprising diversity and recent dramatic decline of freshwater mussels in North America have been well documented, although inventory efforts to date have been concentrated in the eastern United States. Unlike their eastern counterparts, western freshwater mussels have received comparatively little attention. The accurate identity of western lineages is a necessary component for future inventory, monitoring, and ecological work involving these taxa. Here we initiate a study involving the most speciose genus (Anodonta) in western North America, incorporating information about type localities and type specimen morphology and describing the discovery of three highly divergent lineages among four western Anodonta species. In a limited phylogenetic analysis, we find (1) that A. californiensis/nuttalliana and A.oregonensis/kennerlyi are distinct, highly divergent clades, and (2) that A. beringiana is more closely allied with A. woodiana, an Asian species, than either of the other two western North American clades. We were largely unable to resolve the placement of these three clades with respect to other anodontines, and suggest the need for a broader phylogenetic framework. We recommend, however, that the existence of these three deeply divergent groups be considered in the development of regional monitoring, conservation and research plans despite the taxonomic uncertainty.     

Null model of biotic homogenization: a test with the European freshwater fish fauna

In recent years, there has been growing concern about how species invasions and extinctions could change the distinctiveness of formerly disparate fauna and flora, a process called biotic homogenization. In the present study, a null model of biotic of homogenization was developed and applied to the European freshwater fish fauna. We found that non-native fish species led to the greatest homogenization in south-western Europe and greatest differentiation in north-eastern Europe. Comparing these observed patterns to those expected by our null model empirically demonstrated that biotic homogenization is a non-random ecological pattern, providing evidence for previous assumptions. The place of origin of non-native species was also considered by distinguishing between exotic (originating from outside Europe) and translocated species (originating from within Europe). We showed that exotic and translocated species generated distinct geographical patterns of biotic homogenization across Europe because of their contrasting effects on the changes in community similarity among river basins. Translocated species promoted homogenization among basins, whereas exotic species tended to decrease their compositional similarity. Quantifying the individual effect of exotic and translocated species is therefore an absolute prerequisite to accurately assess the spatial dynamics of biotic homogenization.     

Freshwater mussel (Mollusca: Bivalvia: Unionoida) richness and endemism in the ecoregions of Africa and Madagascar based on comprehensive museum sampling

The objective of this study was to assess freshwater mussel (Mollusca: Bivalvia: Unionoida) species distributions among the freshwater ecoregions of Africa and Madagascar to discover areas of high richness and endemism. These are among the top criteria for identifying biodiversity hotspots and establishing conservation priorities. Distributions were determined from museum specimens in 17 collections. In total, 5,612 records for 87 unionoid species could each be assigned to one of 90 freshwater ecoregions. The majority of species (55%) are known from only one (34 spp.) or two (14) ecoregions. Only three are known from more than 20 ecoregions: Etheria elliptica (38 ecoregions), Chambardia wahlbergi (25), and Mutela rostrata (21). The most species-rich ecoregions are Lake Victoria Basin (17 spp.), Upper Nile (16), Upper Congo (14), Senegal–Gambia (13), and Sudanic Congo–Oubangi (13). Those with the most endemic species are Lake Tanganyika (8 spp.), Lake Victoria Basin (6), Bangweulu–Mweru (4), and Lake Malawi (3). Twenty-five ecoregions have no known freshwater mussels. These patterns are significantly correlated with fish and general freshwater mollusk richness. Unionoid richness also varies significantly among major habitat types. These patterns are relevant to biogeography and conservation and indicate areas in need of further research. We argue that freshwater mussels are valuable as focal species for conservation assessments, and they themselves merit management consideration for their ecosystem functions and distributions in imperiled habitats. It is recommended that field surveys be conducted to determine the current status of species in all areas of Africa and Madagascar.     

Freshwater megafauna diversity: Patterns,status and threats

Aim

Freshwater megafauna remain underrepresented in research and conservation, despite a disproportionately high risk of extinction due to multiple human threats. Therefore, our aims are threefold; (i) identify global patterns of freshwater megafauna richness and endemism, (ii) assess the conservation status of freshwater megafauna and (iii) demonstrate spatial and temporal patterns of human pressure throughout their distribution ranges.

Location

Global.

Methods

We identified 207 extant freshwater megafauna species, based on a 30 kg weight threshold, and mapped their distributions using HydroBASINS subcatchments (level 8). Information on conservation status and population trends for each species was extracted from the IUCN Red List website. We investigated human impacts on freshwater megafauna in space and time by examining spatial congruence between their distributions and human pressures, described by the Incident Biodiversity Threat Index and Temporal Human Pressure Index.

Results

Freshwater megafauna occur in 76% of the world’s main river basins (level 3 HydroBASINS), with species richness peaking in the Amazon, Congo, Orinoco, Mekong and Ganges‐Brahmaputra basins. Freshwater megafauna are more threatened than their smaller counterparts within the specific taxonomic groups (i.e., fishes, mammals, reptiles and amphibians). Out of the 93 freshwater megafauna species with known population trends, 71% are in decline. Meanwhile, IUCN Red List assessments reported insufficient or outdated data for 43% of all freshwater megafauna species. Since the early 1990s, human pressure has increased throughout 63% of their distribution ranges, with particularly intense impacts occurring in the Mekong and Ganges‐Brahmaputra basins.

Main conclusions

Freshwater megafauna species are threatened globally, with intense and increasing human pressures occurring in many of their biodiversity hotspots. We call for research and conservation actions for freshwater megafauna, as they are highly sensitive to present and future pressures including a massive boom in hydropower dam construction in their biodiversity hotspots.

     

Using a multi‐model ensemble approach to determine biodiversity hotspots with limited occurrence data in understudied areas: An example using freshwater mussels in México

Species distribution models (SDMs) are an increasingly important tool for conservation particularly for difficult‐to‐study locations and with understudied fauna. Our aims were to (1) use SDMs and ensemble SDMs to predict the distribution of freshwater mussels in the Pánuco River Basin in Central México; (2) determine habitat factors shaping freshwater mussel occurrence; and (3) use predicted occupancy across a range of taxa to identify freshwater mussel biodiversity hotspots to guide conservation and management. In the Pánuco River Basin, we modeled the distributions of 11 freshwater mussel species using an ensemble approach, wherein multiple SDM methodologies were combined to create a single ensemble map of predicted occupancy. A total of 621 species‐specific observations at 87 sites were used to create species‐specific ensembles. These predictive species ensembles were then combined to create local diversity hotspot maps. Precipitation during the warmest quarter, elevation, and mean temperature were consistently the most important discriminatory environmental variables among species, whereas land use had limited influence across all taxa. To the best of our knowledge, our study is the first freshwater mussel‐focused research to use an ensemble approach to determine species distribution and predict biodiversity hotspots. Our study can be used to guide not only current conservation efforts but also prioritize areas for future conservation and study.     

Conservation threats and future prospects for the freshwater fishes of Ecuador: A hotspot of Neotropical fish diversity

Freshwater fish communities in Ecuador exhibit some of the highest levels of diversity and endemism in the Neotropics. Unfortunately, aquatic ecosystems in the country are under serious threat and conditions are deteriorating. In 2018–19, the government of Ecuador sponsored a series of workshops to examine the conservation status of Ecuador's freshwater fishes. Concerns were identified for 35 species, most of which are native to the Amazon region, and overfishing of Amazonian pimelodid catfishes emerged as a major issue. However, much of the information needed to make decisions across fish groups and regions was not available, hindering the process and highlighting the need for a review of the conservation threats to Ecuador's freshwater fishes. Here, we review how the physical alteration of rivers, deforestation, wetland and floodplain degradation, agricultural and urban water pollution, mining, oil extraction, dams, overfishing, introduced species and climate change are affecting freshwater fishes in Ecuador. Although many of these factors affect fishes throughout the Neotropics, the lack of data on Ecuadorian fish communities is staggering and highlights the urgent need for more research. We also make recommendations, including the need for proper enforcement of existing environmental laws, restoration of degraded aquatic ecosystems, establishment of a national monitoring system for freshwater ecosystems, investment in research to fill gaps in knowledge, and encouragement of public engagement in citizen science and conservation efforts. Freshwater fishes are an important component of the cultural and biological legacy of the Ecuadorian people. Conserving them for future generations is critical.     

Genetic structuring in the freshwater mussel Anodonta corresponds with major hydrologic basins in the western United States

Freshwater mussels (unionids) are increasingly recognized as important providers of ecosystem services, yet are among the most endangered fauna in the world. Because unionids are generally sessile and require specific fish hosts for development and dispersal, they are particularly vulnerable to habitat degradation. Surprisingly, little is known about the distribution of genetic diversity in freshwater mussels and this gap has a negative impact on taxonomy, monitoring, conservation and ecological research in these species. Here, we focus on western North American Anodonta, one of only three genera known to exist in this broad landscape and which contains three highly divergent lineages. We describe phylogeographical subdivision in the most widespread and diverse of these lineages, which includes Anodonta californiensis and Anodonta nuttalliana and occurs from Canada to Mexico. Using mitochondrial and nuclear data, we found that genetic structuring within this clade is inconsistent with morphologically based species designations, but instead follows patterns of vicariance among major hydrogeologic basins. Furthermore, there was a strong tendency for population diversity within drainage systems to increase downstream, implying greater habitat or host fish availability in this direction. Microsatellite results indicated that sampling locations were all genetically distinct, even at short distances. Many of our sample populations showed evidence of a recent demographic bottleneck, although this effect seemed to be very local and not drainage or basin‐specific. This study provides a foundation for the establishment of appropriate management units and future research on adaptive differentiation and host fish relationships.     

Dispersal limitation of unionid mussels and implications for their conservation

1. Freshwater unionid mussels are a highly imperilled group. Their dispersal abilities depend on the availability and the movement of host fish on which their parasitic mussel larvae develop. 2. We examined the relationship between the dispersal abilities of unionid mussels and their conservation status on a regional (SW Ontario) scale and their distribution and abundance on a catchment scale (Sydenham River, SW Ontario) by determining host specificity and estimating the dispersal abilities of mussels on fish from a review of the literature. 3. On the regional scale, we found that mussels with the most precarious conservation status relied on host fish with short movement distances, whereas vulnerable and more secure mussel species had host fish with 2–3 orders of magnitude larger movement distances. We were not able to detect a clear pattern on the catchment scale. 4. Our results suggest that limited dispersal by host fish affects the abundance and distribution of unionid mussels and ultimately their conservation status on a regional scale. Information on dispersal limitations because of differences in host fish communities should be included in conservation and management decisions to ensure connectivity and maintain functioning mussel metacommunities.     

Bivalves in a bottleneck: taxonomy,phylogeography and conservation of freshwater mussels (Bivalvia: Unionoida) in Australasia

The conservation biology of Australasian freshwater mussels is hindered by lack of a taxonomic framework that employs molecular data as a complement to shell characters, larval forms and internal anatomy. The fauna includes more than 32 known species (30+ Hyriidae, 2 Unionidae), but has not been revised for 55 years, despite minor amendments. The hyriids are relics of Gondwana, represented in Australia and New Guinea by the ancestral Velesunioninae and in Australia and New Zealand by the Hyriinae (Tribe Hyridellini). Many taxonomic and phylogeographic issues await resolution, including the relationships between Australasian and South American species, and between Australian and New Zealand species, and the status of species in New Guinea (including uncertain reports of Unionidae) and the Solomon Islands. Once these are clarified, it will be easier to identify threatened species and evaluate the conservation status of the fauna. At present, only seven taxa are named in the IUCN Red List or under national/state legislation, and these are not representative. Threatening processes include altered flow regimes, catchment disturbances, salinisation, pollution and invasive species. While the need for a taxonomic revision is paramount, progress in conservation may depend also upon involving the wider community.     

The freshwater pearl mussel <Emphasis Type="Italic">Margaritifera margaritifera</Emphasis> in Bavaria,Germany—Population status,conservation efforts and challenges

The freshwater pearl mussel Margaritifera margaritifera is a highly specialized, sensitive and critically endangered freshwater bivalve with a complex life cycle. An overview on the population status in Bavaria, Germany, one of the species’ main areas of distribution in Central Europe, is presented. Using the example of the freshwater pearl mussel and a Bavarian conservation project for this species, a scheme for the development of site-specific conservation concepts is discussed.     

The role of local adaptation in shaping fish‐mussel coevolution

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