Drought‐induced changes in flow regimes lead to long‐term losses in mussel‐provided ecosystem services

Extreme hydro‐meteorological events such as droughts are becoming more frequent, intense, and persistent. This is particularly true in the south central USA, where rapidly growing urban areas are running out of water and human‐engineered water storage and management are leading to broad‐scale changes in flow regimes. The Kiamichi River in southeastern Oklahoma, USA, has high fish and freshwater mussel biodiversity. However, water from this rural river is desired by multiple urban areas and other entities. Freshwater mussels are large, long‐lived filter feeders that provide important ecosystem services. We ask how observed changes in mussel biomass and community composition resulting from drought‐induced changes in flow regimes might lead to changes in river ecosystem services. We sampled mussel communities in this river over a 20‐year period that included two severe droughts. We then used laboratory‐derived physiological rates and river‐wide estimates of species‐specific mussel biomass to estimate three aggregate ecosystem services provided by mussels over this time period: biofiltration, nutrient recycling (nitrogen and phosphorus), and nutrient storage (nitrogen, phosphorus, and carbon). Mussel populations declined over 60%, and declines were directly linked to drought‐induced changes in flow regimes. All ecosystem services declined over time and mirrored biomass losses. Mussel declines were exacerbated by human water management, which has increased the magnitude and frequency of hydrologic drought in downstream reaches of the river. Freshwater mussels are globally imperiled and declining around the world. Summed across multiple streams and rivers, mussel losses similar to those we document here could have considerable consequences for downstream water quality although lost biofiltration and nutrient retention. While we cannot control the frequency and severity of climatological droughts, water releases from reservoirs could be used to augment stream flows and prevent compounded anthropogenic stressors.     

Coextirpation of host–affiliate relationships in rivers: the role of climate change,water withdrawal,and host‐specificity

The role of climate‐related disturbances on complex host–affiliate relationships remains understudied, largely because affiliate species vary in host use and are often differentially susceptible to disturbance relative to their hosts. Here we report the first set of host–affiliate species–discharge relationships (SDR) in freshwater and examine how anticipated shifts in water availability (flow) will impact coextirpations. We used SDR for freshwater mussels and fish across 11 regions (over 350 rivers) in the continental United States that we coupled to future water availability (2070) to model mussel and fish coextirpations. We also used river‐specific host–affiliate matrices (presence–absence) to evaluate how host‐specificity (mean number of hosts used by an affiliate) and host‐overlap (extent to which affiliates share hosts) relate to extirpation vulnerability. We found that the strength and predictability of SDR models vary geographically and that mussels were more susceptible to flow alterations than fish. These patterns of extirpations were strongest in the southeast where: (1) flow reductions are expected to be greatest; (2) more species are lost per unit flow; (3) and more mussels are expected to be lost per unit of fish. We also found that overall mussel losses associated with reduction in habitat (water availability) were greater than those associated with loss of fish hosts which we assumed to be a function of host redundancy. These findings highlight the utility of SDR as a tool for conservation efforts but they also demonstrate the potential severity of reductions in mussel and fish richness as consequence of climate change and water use. Mussels provide key ecosystem services but face multiple pronged attacks from reductions in flow, habitat, and fish hosts. These losses in biodiversity and ecosystem functions can translate into major effects on food webs and nutrient recycling.     

A proximate cue for oviposition site choice in the bitterling (Rhodeus sericeus)

1. We investigated two possible proximate cues used for oviposition site choice by females of the bitterling ( Rhodeus sericeus ), a freshwater fish that spawns on the gills of live unionid mussels. The two cues were the flow velocity and/or oxygen content of water emerging from the exhalant siphon of a mussel.
2. Field observations showed that female bitterling always inspected the exhalant siphons of mussels before they spawned in them. Siphon inspection was not always a prelude to spawning and it may serve as a means of assessing mussel quality. Female skimming behaviour, swimming over a mussel without spawning, may also be used to assess mussel quality, although the mechanism for this is unclear.
3. Measurements of the flow velocity of water emerging from the exhalant siphons of four mussel species ( Anodonta anatina, A. cygnea, Unio pictorum and U. tumidus ) showed a significant difference among species, with U. tumidus having the highest mean flow velocity and U. pictorum the lowest.
4. Measurements of the change in oxygen concentration of water entering a mussel inhalant siphon and leaving its exhalant siphon in field and laboratory studies showed a significant difference among the four mussel species, with A. cygnea exhibiting a significantly higher change in oxygen concentration than the other species.
5. The presence of bitterling embryos in the gills of a mussel significantly increased its oxygen consumption whereas larval glochidia had no significant effect. We discuss oxygen availability as a possible proximate cue for oviposition site choice in bitterling.     

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.     

Population genetic structure of a rare unionid (<Emphasis Type="Italic">Lampsilis cariosa</Emphasis>) in a recently glaciated landscape

The yellow lampmussel (Lampsilis cariosa) is a rare unionid species in need of conservation, as it is declining throughout most of its Atlantic slope range in North America. Because freshwater mussels rely on a fish host for dispersal of their larvae, barriers to the movement of hosts, such as habitat fragmentation by dams, may indirectly affect population genetic structure. We used microsatellite loci to assess genetic variation for L. cariosa within and among three river drainages in the northern part of its range, which emerged from glaciation only ∼ ∼8–10 kya. Despite this relatively recent emergence, significant differences were observed among populations both within and among drainages, possibly because low effective population sizes meant that populations of these mussels achieved drift-migration equilibrium rapidly following glaciation. L. cariosa individuals could be assigned to their own drainages with 89.3% accuracy. Among-population differences were modest, however, in comparison to differences observed in another study of rare mussels south of the recently glaciated region. L. cariosa populations exhibited significant isolation by distance, but there was no additional variation explained by the number, size, or age of intervening dams. An understanding of mussel population genetic structure provides information, useful for conservation planning, on patterns of isolation and connectivity among populations.     

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.     

Dynamic measurements of black oystercatcher (Haematopus bachmani) predation on mussels (Mytilus californianus)

Intertidal zone mussels can face threats from a variety of predatory species during high and low tides, and they must balance the threat of predation against other needs such as feeding and aerobic respiration. Black oystercatchers (Haematopus bachmani) on the Pacific coast of North America can depend on the mussel Mytilus californianus for a substantial portion of their diet. Observations suggest that oystercatchers tend to focus on mussels beginning to gape their valves during rising tides, following periods of aerial emersion. We present detailed, autonomous field measurements of the dynamics of three such predation events in the rocky intertidal zone. We measured accelerations of up to 4 g imposed on mussels, with handling times of 115–290 s required to open the shell and remove the majority of tissue. In each case a single oystercatcher attacked a mussel that had gaped the shell valves slightly wider than its neighbors as the rising tide began to splash the mussel bed, but no other obvious characteristic of the mussels, such as body temperature or orientation, could be linked to the oystercatcher's individual prey choice.     

Microsatellite loci for dreissenid mussels (Mollusca: Bivalvia: Dreissenidae) and relatives: markers for assessing exotic and native populations

We developed and tested 14 new polymorphic microsatellite loci for dreissenid mussels, including the two species that have invaded many freshwater habitats in Eurasia and North America, where they cause serious industrial fouling damage and ecological alterations. These new loci will aid our understanding of their genetic patterns in invasive populations as well as throughout their native Ponto-Caspian distributions. Eight new loci for the zebra mussel Dreissena polymorpha polymorpha and six for the quagga mussel D. rostriformis bugensis were compared with new results from six previously published loci to generate a robust molecular toolkit for dreissenid mussels and their relatives. Taxa tested include D. p. polymorpha, D. r. bugensis, D. r. grimmi, D. presbensis, the 'living fossil'Congeria kusceri, and the dark false mussel Mytilopsis leucophaeata (the latter also is invasive). Overall, most of the 24 zebra mussel (N = 583) and 13 quagga mussel (N = 269) population samples conformed to Hardy-Weinberg equilibrium expectations for the new loci following sequential Bonferroni correction. The 11 loci (eight new, three previously published) evaluated for D. p. polymorpha averaged 35.1 alleles and 0.72 mean observed heterozygosity per locus, and 25.3 and 0.75 for the nine loci (six new, three previously published) developed for D. r. bugensis. All but three of these loci successfully amplified the other species of Dreissena, and all but one also amplified Congeria and Mytilopsis. All species and populations tested were significantly divergent using the microsatellite data, with neighbour-joining trees reflecting their evolutionary relationships; our results reveal broad utility for resolving their biogeographic, evolutionary, population and ecological patterns.     

Spatio-temporal patterns of the decline of freshwater mussels in the Little South Fork Cumberland River,USA

The Little South Fork Cumberland River, Kentucky and Tennessee, USA, was a globally important conservation refugium for freshwater mussels (Mollusca: Unionidae) because it supported an intact example (26 species) of the unique Cumberland River mussel fauna including imperiled species. We used previous surveys and our 1997–1998 survey to reconstruct the historical fauna, to describe spatio-temporal patterns of density and number of species, and to evaluate the probable sequence and cause of observed mussel declines. We were specifically interested in better understanding how mussel assemblages respond to chronic disturbances, and how these changes manifest in persistence patterns. Density and numbers of species declined steadily from 1981 to 1998, but declines occurred first in the lower river (early 1980s), followed by declines in the upper river (late 1980s to early 1990s). Of the total species recorded from the Little South Fork, 17 (65%) are seemingly extirpated and five others appear near extirpation. Declines are associated with at least two, temporally distinct major insults. Lower river declines are associated with surface mining, whereas, oil extraction activities are implicated in upper river declines. Regardless of causal factors, species persistence was primarily a function of predecline population size with only the most numerous and widespread species surviving. At this time, the river appears lost as a conservation refugium for mussels despite its remoteness, predominantly forested watershed, and several layers of existing statutory and regulatory environmental safeguards. We suggest that the river could be restored and mussels reintroduced if an interagency task force is formed to identify and mitigate specific stressors now affecting most mussel species in the river.Nomenclature: Turgeon et al. (1998).     

Conservation status of freshwater mussels in Europe: state of the art and future challenges

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.     

The effect of seasonal variation in selective feeding by zebra mussels (Dreissena polymorpha) on phytoplankton community composition

1. To investigate the impact of zebra mussels ( Dreissena polymorpha ) on phytoplankton community composition, temporal variability in selective feeding by the mussels was determined from April to November 2005 in a natural lake using Delayed Fluorescence (DF) excitation spectroscopy.
2. Selective grazing by zebra mussels varied in relation to seasonal phytoplankton dynamics; mussels showed a consistent preference for cryptophytes and avoidance of chlorophytes and cyanobacteria. Diatoms, chrysophytes and dinoflagellates responded differentially to zebra mussel grazing depending on their size. Analysis of excreted products of the zebra mussels revealed that in addition to chlorophytes and cyanobacteria, phytoplankton >50  μ m and very small phytoplankton (≤7  μ m) were largely expelled in pseudofaeces.
3. The zebra mussel is a selective filter-feeder that alters its feeding behaviour in relation to phytoplankton composition to capture and ingest high quality phytoplankton, especially when phytoplankton occur in preferred size ranges. Flexibility of zebra mussel feeding behaviour and variation in susceptibility among phytoplankton groups to mussel ingestion indicate that invading zebra mussels could alter phytoplankton community composition of lakes and have important ecosystem consequences.     

Predator‐induced morphological defences in two invasive dreissenid mussels: implications for species replacement

相似文献   

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.     

Ecosystem Processes Performed by Unionid Mussels in Stream Mesocosms: Species Roles and Effects of Abundance

Unionid mussels are a guild of freshwater, sedentary filter-feeders experiencing a global decline in both species richness and abundance. To predict how these losses may impact stream ecosystems we need to quantify the effects of both overall mussel biomass and individual species on ecosystem processes. In this study we begin addressing these fundamental questions by comparing rates of ecosystem processes for two common mussel species, Amblema plicata and Actinonaias ligamentina, across a range of abundance levels and at two trophic states (low and high productivity) in stream mesocosms. At both low and high productivity, community respiration, water column ammonia, nitrate, and phosphorus concentrations, and algal clearance rates were all linearly related to overall mussel biomass. After removing the effects of biomass with ANCOVA, we found few differences between species. In a separate series of experiments, nutrient excretion (phosphorus, ammonia, and molar N:P) and biodeposition rates were only marginally different between species. For the species studied here, functional effects of unionids in streams were similar between species and linearly related to biomass, indicating the potential for strong effects when overall mussel biomass is high and hydrologic residence times are long.     

Life history traits and abundance can predict local colonisation and extinction rates of freshwater mussels

1. A critical need in conservation biology is to determine which species are most vulnerable to extinction. Freshwater mussels (Bivalvia: Unionacea) are one of the most imperilled faunal groups globally. Freshwater mussel larvae are ectoparasites on fish and depend on the movement of their hosts to maintain connectivity among local populations in a metapopulation. 2. I calculated local colonisation and extinction rates for 16 mussel species from 14 local populations in the Red River drainage of Oklahoma and Texas, U.S. I used general linear models and AIC comparisons to determine which mussel life history traits best predicted local colonisation and extinction rates. 3. Traits related to larval dispersal ability (host infection mode, whether a mussel species was a host generalist or specialist) were the best predictors of local colonisation. 4. Traits related to local population size (regional abundance, time spent brooding) were the best predictors of local extinction. The group of fish species used as hosts by mussels also predicted local extinction and was probably related to habitat fragmentation and host dispersal abilities. 5. Overall, local extinction rates exceeded local colonisation rates, indicating that local populations are becoming increasingly isolated and suffering an ‘extinction debt’. This study demonstrates that analysis of species traits can be used to predict local colonisation and extinction patterns and provide insight into the long‐term persistence of populations.     

Density effects on the clearance rate of the zebra mussel <Emphasis Type="Italic">Dreissena polymorpha</Emphasis>: flume study results

Zebra mussel filtration rates and regulating factors have been addressed earlier in a number of studies. Still, only a few of them have taken into consideration the refiltration phenomenon, and therefore the direct extrapolation of experimental results may only give the potential filtering capacity, and hence, over- or underestimate the actual amount of seston being removed by zebra mussels in an ecosystem. The current experimental study aimed to gain insight into the refiltration effect on the clearance rate of the zebra mussels at relatively high seston concentrations, and its potential role in controlling the filtration efficiency of the zebra mussel population. The experiment was conducted in a laboratory flume following the Latin squares design with one fixed (mussel density) and three random factors (initial total particulate matter (TPM) concentration, flume “wall effect” and distance from the flume inflow area) considered. The results showed the significant effects of mussel density and the TPM concentration on the effective clearance rate (ECR) of zebra mussels. The higher ECR values were obtained at denser mussel clumps and lower TPM concentrations. The flume “wall effect” had no significant effect on the ECR, whereas the distance from the flume inflow area appeared to have a significant impact. A positive relationship between ECR and the zebra mussel density was most evident in the proximity of the TPM source. Based on the results, we assume that at high TPM concentration, refiltration may assert itself by the elevated net clearance rate of mussels within dense clumps compared to that of mussels at relatively low individual densities. This should be taken into consideration while modelling and assessing the role of the zebra mussel in energy flow and redistribution of organic matter in an ecosystem.     

Indigenous microflora and opportunistic pathogens of the freshwater zebra mussel, Dreissena polymorpha

Freshwater fouling invertebrate zebra mussels (Dreissena polymorpha) harbor a diverse population of microorganisms in the Great Lakes of North America. Among the indigenous microorganisms, selective species are opportunistic pathogens to zebra mussels. Pathogenicity to zebra mussels by opportunistic bacteria isolated from the mussels was investigated in this study. Among the more than 30 bacteria isolated from temperature-stressed mussels, Aeromonas media, A. veronii, A. salmonicida subsp. salmonicida, and Shewanella putrefaciens are virulent pathogens to juvenile zebra mussels. Inoculation of a bacterial concentration of A. media, A. salmonicida subsp. salmonicida and S. putrefaciens at 10⁷ cells per zebra mussel resulted in 100% mortality within 5 days, and only 64.9% for A. veronii. In contrast, mortality was less than 12.3% following inoculation of a sterile phosphate buffer solution as a control. In addition, mortality was dependent on the size of the pathogen population used in inoculation and the incubation temperature, indicating the close relationship between the bacterial population and subsequent death. On the mussel tissue, a dense microbial population was evident from the moribund mussels viewed with Scanning Electron Microscope (SEM). Opportunistic bacteria invaded and destroyed the D. polymorpha tissue after 7 days of incubation when the bacterial inoculation was larger than 10⁵ per zebra mussel. Our results suggest that mussels are reservoirs of opportunistic pathogenic microorganisms to aquatic organisms and humans and a better understanding of the microbial ecology of the mussels will provide insights to the possible health hazards from these microorganisms.     

Intertidal assemblages associated with natural<Emphasis Type="Italic">corallina</Emphasis> turf and invasive mussel beds

There is considerable concern about conservation of biodiversity in highly disturbed and urbanized environments, although a very large proportion of biodiversity (i.e. the small and cryptic invertebrates) have been little studied in this regard. Many biogenic structures (e.g. coral reefs, mussel beds, foliose algae) provide habitat for a large number of small invertebrates. The features of these habitats to which these animals respond are complex and poorly documented. Invasive species are increasing in abundance and diversity in many disturbed estuaries, but most previous studies have concentrated on effects of invasive species on surrounding macroscopic assemblages. This study examines the assemblages of small invertebrates and algae living in natural patches of coralline turf and in patches of the invasive mussel, Mytilus galloprovincialis, on seawalls in Sydney Harbour. Although most taxa identified were common to both habitats, they were generally more abundant in turf than in the mussels, especially the more widespread and numerous taxa. Few taxa were unique to either habitat and those were generally sparse and patchy. In addition, there were relatively more smaller animals in the algal turf than in the mussels, although it is not known whether these were juveniles of adults present in both habitats, or different species. These data show that coralline turf and mussel beds do not provide similar intertidal habitat for associated assemblages and that overgrowth of natural biota by mussels may have strong indirect effects on associated assemblages. These warrant further experimental investigation, so that the effects of invasive species on local biodiversity can be better understood and managed.     

The impact of land use on the mussel Margaritifera margaritifera and its host fish Salmo trutta

Today, land use impacts a major proportion of all streams. Here, landscape features in corridors along streams and water chemical factors were analyzed in relation to recruitment of the threatened freshwater pearl mussel (Margaritifera margaritifera) and its host fish the brown trout (Salmo trutta). Mussel recruitment and trout density were negatively related to forest clear-cuts. Mussel recruitment was negatively related to water color and turbidity. Therefore, the threats to the mussel may be severe, as low mussel recruitment may be caused by direct effects on the juvenile mussels and indirect effects on the host fish. High proportions of lakes and ponds were found to be positive for recruitment and for trout, and deciduous forest was positively related to trout. The combination of investigations at different scales at the landscape level and at in-stream levels may be applicable to find threats to other threatened species. The results indicate that forestry activities may negatively affect recruitment of freshwater pearl mussels and its host fish. Reductions of forestry activities and the retaining of intact quantity and quality of the riparian zones next to streams, both for the mussel and its host fish may be important conservation measures to restore freshwater pearl mussel populations.