Stream channel geomorphology influences mussel abundance in southern Appalachian streams, U.S.A.

1. Studies of North American streams have shown that hydraulic parameters and stream geomorphology can explain unionid mussel abundance at both the reach and catchment scale. However, few studies have examined applicability of hydrogeomorphic variables across broader spatial scales, such as across whole catchments, or have elucidated conditions under which spates can affect mussel populations in streams. 2. We quantified freshwater mussel abundance and species richness and their physical habitat at 24 sites in eight streams in southern Appalachian catchments in 2000 and 2001. In addition, we modelled site‐specific hydraulic parameters during summer baseflow and bankfull stages to estimate high‐ and low‐discharge conditions, respectively. 3. Mussel abundance was related to stream geomorphology, whereas richness was related to stream size. Baseflow habitat parameters explained only minor variation in abundance or richness, and both measures were highly correlated with mean current velocity or stream size. Bankfull shear stress composed a relatively low proportion of overall mussel habitat variability, but it accounted for significant variation in abundance and richness. 4. Mussel abundance was highly variable at sites subject to low‐shear stress during spates, whereas abundance always was low at sites subject to high‐shear stress. These data suggest that habitat conditions during floods, rather than those at summer baseflow, limit the abundance of mussels in Appalachian streams. These data also suggest that mussel abundance and assemblage structure may be sensitive to any changes in channel geomorphology and hydraulic conditions that might result from land use in the catchment.     

Reach- and catchment-scale determinants of the distribution of freshwater mussels (Bivalvia: Unionidae) in south-eastern Michigan, U.S.A.

1. We investigated the diversity and distribution of freshwater mussels at 40 sites in an agricultural catchment, the River Raisin in south‐eastern Michigan, to relate mussel assemblages and individual taxa to reach and catchment‐scale variables. Unionids were surveyed by timed searches in 100‐m reaches, and in‐stream and riparian habitat were quantified as well as flow, water chemistry and channel morphology. Land use/cover and surficial geology were determined for site subcatchments and riparian buffers. 2. Some 21 mussel species were found overall; richness ranged from 0 to 12 living species per site. From the upper to middle to lower catchment, the number of individuals, number of species, Shannon–Weaver diversity and relative abundance of intolerant unionids all declined significantly. 3. Four groupings based on overall mussel diversity and abundance were significantly related to reach‐scale habitat variables. The richest mussel assemblages were associated with sites with higher overall habitat quality, greater flow stability, less fine substratum, and lower specific conductance. 4. Stepwise multiple regressions revealed that the distribution and abundance of the total mussel assemblage, as well as the most common species, could be predicted from a combination of reach‐ and catchment‐scale variables (R² = 0.63 for total mussels, R² = 0.51–0.86 for individual species). 5. Flow stability, substratum composition and overall reach habitat quality were the most commonly identified reach‐scale variables, and measures of surficial geology were the most effective catchment‐scale variables. The spatial pattern of geology is likely to be responsible for the diversity gradient from the upper to the lower catchment. 6. Prior studies, attempting to explain mussel distributions from local habitat features alone, have found relatively weak relationships. By employing a combination of reach‐ and catchment‐scale habitat variables, this study was able to account for a substantial amount of the spatial variability in mussel distributions.     

Distribution and microhabitats of freshwater mussels in waterbodies in the terrestrialized floodplains of a lowland river

Even when anthropogenically altered, river floodplains continue to contribute to biodiversity. This study examined the distribution of freshwater mussels in relation to environmental factors in waterbodies in the terrestrialized floodplain of a lowland river. Mussels were captured, and environmental measurements were taken in November of 2013 and 2014 in quadrats established in three floodplain waterbodies (FWBs), which were isolated from the main river channel. Among the three FWBs, mussel abundance was highest in a shallow FWB (depth range 18–45 cm) that had intermediate conditions of mud depth and fine sediment rate. Mussel abundance showed a hump-shaped relationship with water depth (the peak 45–50 cm) and mud depth (the peak 8–12 cm). Mussel abundance was also negatively related to the abundance of benthic litter. Litter abundance was positively related to branch abundance and the presence of tree cover, and negatively related to the distance to tree cover, indicating that benthic litter was derived from riparian trees. Our results indicate that relatively shallow (≤ 50 cm) FWBs with moderately accumulated mud, which are not scoured even during flooding, appear to be suitable habitats for mussels. Moreover, it is possible that riparian trees negatively impact mussel distribution in FWBs. Possible short-term measures for improving mussel habitat in FWBs may include the elimination of riparian trees and benthic litter.     

Modelling and mapping the distribution,diversity and abundance of freshwater mussels (Family Unionidae) in wadeable streams of Illinois,U.S.A.

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Novel insights into habitat suitability for Amazonian freshwater mussels linked with hydraulic and landscape drivers

Novel insights into habitat suitability for two Unionida freshwater mussels, Castalia ambigua Lamarck, 1819 (Hyriidae) and Anodontites elongatus (Swainson, 1823) (Mycetopodidae), are presented on the basis of hydraulic variables linked with the riverbed in six 500‐m reaches in an eastern Amazonian river basin. Within the reaches, there was strong habitat heterogeneity in hydrodynamics and substrate composition. In addition, we investigated stressors based on landscape modification that are associated with declines in mussel density. We measured hydraulic variables for each 500‐m reach, and landscape stressors at two spatial scales (subcatchment and riparian buffer forest). We used the Random Forest algorithm, a tree‐based model, to predict the hydraulic variables linked with habitat suitability for mussels, and to predict which landscape stressors were most associated with mussel density declines. Both mussel species were linked with low substrate heterogeneity and greater riverbed stability (low Froude and Reynolds numbers), especially at high flow (low stream power). Different sediment grain size preferences were observed between mussel species: Castalia ambigua was associated with medium sand and Anodontites elongatus with medium and fine sand. Declines in mussel density were associated with modifications linked to urbanization at small scales (riparian buffer forest), especially with percent of and distance from rural settlements, distance to the nearest street, and road density. In summary, the high variance explained in both hydraulic and landscape models indicated high predictive power, suggesting that our findings may be extrapolated and used as a baseline to test hypotheses of habitat suitability in other Amazonian rivers for Castalia ambigua and Anodontites elongatus and also for other freshwater mussel species. Our results highlight the urgent need for aquatic habitat conservation to maintain sheltered habitats during high flow as well as mitigate the effects of landscape modifications at the riparian buffer scale, both of which are important for maintaining dense mussel populations and habitat quality.     

Retention and supply of zebra mussel larvae in a large river system: importance of an upstream lake

1. Persistence of zebra mussel populations in river systems probably depends upon the presence of upriver sites capable of hosting self‐recruiting adult populations that act as sources of larvae. In this paper we examine the importance of Lake Pepin, a natural riverine lake in the Upper Mississippi River, as an upriver source of larvae to the downstream populations of zebra mussels. 2. Field studies and modelling suggest that Lake Pepin plays a major role in maintaining zebra mussel populations in the Upper Mississippi River. Long water residence times in Lake Pepin allow for self‐recruitment under the right hydraulic conditions. Larval abundance was low to absent upstream of the lake but increased dramatically downriver in all 3 years of the study. Travel time estimates in the Upper Mississippi River show that newly fertilised larvae drifting out of Lake Pepin can contribute substantially to the major downstream peak in larval abundance. In contrast, backwater and other off‐channel sites are unlikely to drive main‐channel abundance patterns. Larval abundances in off‐channel sites were less than or equal to those in the main channel. 3. A key factor in assessing the importance of Lake Pepin as a source population was the abundance of early stage, unshelled larvae. Studies that consider only abundances of older shelled stages (visible by cross polarised lighting) may yield misleading results. Results of this study suggest that efforts to control zebra mussels in the Upper Mississippi River should focus on controlling adult populations within Lake Pepin and reducing or eliminating larvae exiting the lake.     

Context-dependent effects of freshwater mussels on stream benthic communities

1. We asked whether unionid mussels influence the distribution and abundance of co‐occurring benthic algae and invertebrates. In a yearlong field enclosure experiment in a south‐central U.S. river, we examined the effects of living mussels versus sham mussels (shells filled with sand) on periphyton and invertebrates in both the surrounding sediment and on mussel shells. We also examined differences between two common unionid species, Actinonaias ligamentina (Lamarck 1819) and Amblema plicata (Say 1817). 2. Organic matter concentrations and invertebrate densities in the sediment surrounding mussels were significantly higher in treatments with live mussels than treatments with sham mussels or sediment alone. Organic matter was significantly higher in the sediment surrounding Actinonaias than that surrounding Amblema. Actinonaias was more active than Amblema and may have increased benthic organic matter through bioturbation. 3. Living mussels increased the abundance of periphyton on shells and the abundance and richness of invertebrates on shells, whereas effects of sham mussels were similar to sediment alone. Differences in the amount of periphyton growing on the shells of the two mussel species reflected differences in mussel activity and shell morphology. 4. Differences between living and sham mussel treatments indicate that biological activities of mussels provide ecosystem services to the benthic community beyond the physical habitat provided by shells alone. In treatments containing live mussels we found significant correlations between organic matter and chlorophyll a concentrations in the sediment, organic matter concentrations and invertebrate abundance in the sediment and the amount of chlorophyll a on the sediment and invertebrate abundance. There were no significant correlations among these response variables in control treatments. Thus, in addition to providing biogenic structure as habitat, mussels likely facilitate benthic invertebrates by altering the availability of resources (algae and organic matter) through nutrient excretion and biodeposition. 5. Effects of mussels on sediment and shell periphyton concentrations, organic matter concentrations and invertebrate abundance, varied seasonally, and were strongest in late summer during periods of low water volume, low flow, and high water temperature. 6. Our study demonstrates that freshwater mussels can strongly influence the co‐occurring benthic community, but that effects of mussels are context‐dependent and may vary among species.     

Predicting zebra mussel fouling on native mussels from physicochemical variables

1. Predictive models of impact are needed for the risk assessment of invasive species. One such species is the Eurasian zebra mussel ( Dreissena polymorpha ), a fouling bivalve that overgrows and kills native mussels (Unionidae) in many North American lakes and rivers. The level of mortality in a native mussel population increases with the fouling intensity – i.e. the mean number of zebra mussels attached to each native mussel.
2. We conducted a multi-site survey within the St Lawrence and Richelieu rivers (Quebec, Canada) to determine whether zebra mussel fouling intensity can be predicted from environmental variables. We found fouling intensity to be positively correlated with calcium concentration [Ca²⁺] and negatively correlated with sediment size, but not affected by local macrophyte cover.
3. A multiple regression model that includes calcium concentration and sediment size explains 86% of variation in fouling intensity across all sites.
4. Analysis of data from invaded sites in North America and Europe revealed a nonlinear relationship in which fouling intensity increases with calcium concentration up to an asymptotic threshold of 24 mg L⁻¹ Ca²⁺.
5. Our results suggest that the community-level impacts of zebra mussels are mediated by abiotic environmental variables and gradients in these variables may provide local habitat refugia for native mussels.     

Macroecology of a host-parasite relationship

The larvae of freshwater mussels are obligate ectoparasites on fishes while adults are sedentary and benthic. Dispersal of mussels is dependent on the movement of fish hosts, a regional process, but growth and reproduction should be governed by local processes. Thus, mussel assemblage attributes should be predictable from the regional distribution and abundance of fishes. At a broad spatial scale in the Red River drainage, USA, mussel species richness and fish species richness were positively associated; maximum mussel richness was limited by fish richness, but was variable beneath that constraint. Measured environmental variables and the associated local fish assemblages each significantly accounted for the regional variation in mussel assemblages. Furthermore, mussel assemblages showed strong spatial autocorrelation. Variation partitioning revealed that pure fish effects accounted for 15.4% of the variation in mussel assemblages; pure spatial and environmental effects accounted for 16.1% and 7.8%, respectively. Shared variation among fish, space and environmental variables totaled 40%. Of this shared variation, 36.8% was associated with the fish matrix. Thus, the variation in mussel assemblages that was associated with the distribution and abundance of fishes was substantial (> 50%), indicating that fish community structure is an important determinant of mussel community structure. Although animals commonly disperse plants and, thus, influence the structure of plant communities, our results show a strong macroecological association between two disparate animal groups with one strongly affecting the assemblage structure of the other.     

Scale‐dependent longitudinal patterns in mussel communities

1. Species richness and assemblage patterns of organisms are dictated by numerous factors, probably operating at multiple scales. Freshwater mussels (Unionidae) are an endangered, speciose faunal group, making them an interesting model system to study the influence of landscape features on organisms. In addition, landscape features that influence species distributions and the scale at which the factors have the greatest impact are important issues that need to be answered to conserve freshwater mussels. 2. In this study, we quantified freshwater mussel communities at 16 sites along three mid‐sized rivers in the south‐central United States. We addressed the following questions: (i) Are there predictable longitudinal changes in mussel community composition? (ii) What landscape variables best explain shifts in community composition? and (iii) At what scale do landscape variables best predict mussel community composition? 3. After controlling for the influence of longitudinal position along the stream, we compared mussel distributions to a suite of hypothesised explanatory landscape variables across multiple scales – catchment scale (entire drainage area), buffer scale (100‐m riparian buffer of the entire catchment) and reach scale (100‐m riparian buffer extending 1 km upstream from the sampling site). 4. We found a significant and consistent longitudinal shift in dominant mussel species across all three rivers, with community composition strongly related to distance from the headwaters, which is highly correlated with stream size. After accounting for stream size, variables at the buffer scale were the best predictors of mussel community composition. After accounting for catchment position, mean channel slope was the best explanatory variable of community composition and appeared in all top candidate models at the catchment and buffer scales. Coverage of wetland and urban area were also correlated with community composition at the catchment and buffer scales. 5. Our results suggest that landscape‐scale habitat factors influence mussel community composition. Landscape features at the buffer scale performed best at determining community composition after accounting for position in the catchment; thus, further protection of riparian buffers will help to conserve mussel communities.     

Parasitic mite and trematode abundance are associated with reduced reproductive output and physiological condition of freshwater mussels

Although historically understudied, parasites may play an important role in freshwater invertebrate population ecology and evolution. We quantified abundance of parasitic mites and trematodes in the freshwater mussel Pyganodon grandis, in a southeastern Alabama stream (USA), to assess parasite impact on adult mussel physiological condition and reproductive output. We used stepwise multiple regression analyses to assess the effects of mussel size and parasites on reproduction and condition. Multiple regression analysis found no multivariate models that predicted reproductive output or physiological condition. However, univariate models revealed that increased parasite densities predict reduced mussel reproductive output and physiological condition. These findings suggest that parasites may have important negative consequences for freshwater mussels. We hypothesize that elevated parasite loads may reduce mussel fitness in impounded or nutrient-enriched streams with high densities of intermediate hosts (chironomid midges).     

Effects of zebra mussels on phytoplankton and ciliates: a field mesocosm experiment

Many observational studies in North American lakes have documenteddecreases in phytoplankton abundance after the invasion of thezebra mussel (Dreissena polymorpha). However, few field experimentshave examined in detail the effect of zebra mussels on phytoplanktonabundance and species composition over an extended period. Replicatedin situ mesocosms were used to evaluate the impact of naturaldensities of zebra mussels on phytoplankton and ciliate biovolume,and algal species composition over a 5-week period in a habitatthat lacked extant mussel populations. Mussel biomass used inthe experiment was determined using a regression model basedon a data analysis that predicts zebra mussel biomass from totalphosphorus concentration. Within 1 week, zebra mussels decreasedphytoplankton biovolume by 53% and ciliate biovolume by 71%.The effect of zebra mussels on ciliate biovolume was sustainedthroughout the study. However, the effect of zebra mussels onphytoplankton abundance gradually waned over the remaining 4weeks of the experiment, such that the declining effect of zebramussels could not be explained by a shift towards less edibleand/or faster growing algal species. The mussels’ decliningcondition could help to explain the effect observed over thecourse of the experiment.     

Effects of a small dam on freshwater mussel growth in an Alabama (U.S.A.) stream

1. Dams, ubiquitous features in many lotic ecosystems, are believed to have many broad‐ranging and predominantly negative effects on stream biota. Whereas the impacts of larger dams are well studied, few studies have quantified effects of small dams on streams. 2. Recent surveys found numerous locations where mussels were abundant and larger in reaches immediately downstream from small dams. We examined mussel shell growth and resource conditions in Sandy Creek, a small (third‐order) tributary of the Tallapoosa River in east‐central Alabama (U.S.A.), to determine whether larger populations and individuals result from more rapid growth or longer lifespans of mussels downstream from the dam. 3. Growth rates for populations occurring immediately downstream from the dam (mill reach), c. 5 km downstream from the dam (downstream reach) and upstream from the impoundment (upstream reach) were compared with environmental conditions (seasonal measures of nutrient concentrations and water chemistry) and food availability [total suspended solids (TSS)]. Water temperature was continuously monitored using data loggers. 4. Analysis of length‐at‐age data using multiple growth models found that mill reach mussels grew faster than both up‐ and downstream populations. This dam appears to substantially increase water temperatures and may extend the shell growth period in the mill reach. TSS quantity varied seasonally between sites but was generally highest in the impoundment and mill reach during spring and autumn. TSS quality was highest in the upstream reach from spring through autumn but was highest in the impoundment and mill reach during winter. 5. Our data suggest that some small impoundments enhance conditions for freshwater mussel growth in downstream reaches. However, we do not know how far downstream this subsidy extends or how different species respond to mill dam augmentation. Regardless, mounting evidence suggests that this phenomenon is geographically and taxonomically widespread in eastern North America. Heretofore, undocumented positive effects of small dams suggest that some older dams may warrant protection or restoration if downstream reaches support imperilled mussel populations. Further, some small dams may prove useful conservation tools for natural resource managers attempting to identify sites for mussel culture facilities or translocation refugia.     

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.     

Sediment accumulation predicts the distribution of a unionid mussel (Elliptio complanata) in nearshore areas of a Canadian Shield lake

1. The importance of native freshwater mussels for ecosystem processes depends on their density, size distribution and activity. In lakes, many of the factors that affect mussels (fish hosts, habitat, food) could be directly or indirectly related to wind‐driven physical processes. 2. We tested whether the abundance and size of Elliptio complanata in the shallow, nearshore areas of a medium‐sized lake were related to site exposure, substratum type and fish distribution. To disentangle some of the correlated variables known to affect mussel distribution, we used paired exposed and sheltered sampling sites along the 7‐km fetch of the lake basin. 3. The distribution of sediment characteristics in nearshore areas was highly predictable. The mean depth of accumulated soft sediments decreased with increasing fetch at wind‐exposed sites, but increased with increasing fetch at sheltered sites. Sediments were deeper along the main shoreline than around islands. Deeper sediments tended to be finer and higher in silt content and organic fraction. 4. The density and proportion of juvenile mussels along the main shoreline varied in a unimodal way with sediment depth. These results suggest that wind‐driven physical forces affect the transport of young juveniles to sediment depositional areas and create sediment conditions that influence their growth and survival. In contrast, the proportion of juvenile mussels around islands was not related to sediment characteristics, but decreased with remoteness of the island, suggesting that the distribution of juvenile mussels may be limited by fish movements. These results are tentative since they do not include buried juvenile mussels. 5. We also found a unimodal relationship between total mussel density (juveniles and adults) and sediment depth but, in contrast to the relationship for juveniles only, it applied to all sites with soft sediments, including islands. We conclude that factors related to sediment depth affect the growth and survival of adult mussels around islands and that these factors are strong enough to modify the pattern of distribution established via dispersal during earlier life stages. 6. The mean shell length of adults at different sites within the lake basin ranged from 60 to 85 mm. Mussel shell length decreased with increasing fetch at sites exposed to the prevalent winds, but was relatively constant on the sheltered side of peninsulas and islands. The size of unionid mussels in different parts of the lake seems to be determined both by their exposure to physical forces and by sediments. 7. The local distribution of E. complanata is determined, directly and indirectly, by wind‐driven forces. These processes are likely to be important for other benthic organisms affected by similar habitat conditions (e.g. sediment characteristics, physical disturbance).     

Mussel responses to flood pulse frequency: the importance of local habitat

1. Understanding mechanisms behind the distribution of organisms along a gradient of hydrological connectivity is crucial for sustainable management of river–floodplain systems. We tested the hypothesis that frequency of flood pulses exerts a direct influence on the distribution of freshwater mussels (Unionoida) by creating a local environment that limits their fitness. 2. Multiscale habitat analyses combined with transplant‐rearing experiments were carried out with a focus on abundance, presence/absence, survival rates and growth rates of mussels. Sixty‐nine floodplain waterbodies (FWBs) were surveyed within a 15‐km lowland segment of the Kiso River in Japan. 3. The abundance of mussels significantly increased with increased frequency of inundation associated with flood pulses at the among‐FWB scale, while the probability of occurrence of mussels was negatively predicted by the amount of benthic organic matter at the within‐FWB scale. 4. Field‐rearing experiments showed that survival rates were low and growth rates nearly zero in infrequently inundated FWBs (these FWBs had no naturally occurring resident mussels). In such FWBs, hypoxia (DO < 2 mg L^?1) was frequently observed near the bottom when temperature was optimal for mussel growth (>15 °C). 5. These findings demonstrated that flood pulse frequency was the most important factor in determining mussel distribution in FWBs because it directly limits mussels’ fitness by mediating local environmental factors, possibly dissolved oxygen (DO) levels. Successful restoration efforts for mussel habitat conservation should focus on processes that lead to improved local conditions.     

Estimating the effective clearance rate and refiltration by zebra mussels, Dreissena polymorpha, in a stratified reservoir

1. The grazing impact of zebra mussels, Dreissena polymorpha Pallas, is often evaluated by applying the individual filtration rate measured in the laboratory to the field abundance and then by comparing the total volume of water filtered with the whole lake volume. Since this approach overlooks refiltration, it overestimates the grazing impact of zebra mussels. To deal with this problem, the present authors developed an in situ method for collecting faeces and pseudofaeces to measure the actual volume of water that is cleared of suspended particles by Dreissena in a unit time under a given set of environmental conditions. This is termed the effective clearance rate (ECR). 2. The experiment was conducted in Hargus Lake, OH, U.S.A., a small thermally stratified reservoir, to test the effects of spatial aggregation, mussel density and the concentration of particulate inorganic matter (PIM) on the effective clearance rate of Dreissena. 3. Over 40 measurements, the ECR values ranged from 15.3 to 68.6 mL ind^??1 h^??1. Much of the variation can be explained by colony form, mussel density and seston concentration. The effects of these variables were all statistically significant. The average ECR for isolated individuals was higher than that for those in clumps (40.4 versus 32.8 mL ind^??1 h^??1), which is attributed to increased refiltration in the cores of the clumps. The ECR decreased with increased zebra mussel density because of intensified competition for food particles within the group. The ECR increased with increased PIM concentration in the lake water, which may be interpreted as a result of enhanced water mixing which ultimately caused increases in both sediment resuspension and particle delivery to the mussels. 4. Taking the filtration rates for a 20-mm mussel to be between 116 and 234 mL ind^??1 h^??1, based on data from the literature, the clumped mussels under the present experimental conditions would have a refiltration ratio between 3.4 and 6.9. 5. The present authors developed an areal clearance model which predicts that seston removal by the Dreissena population is limited by the particle delivery from the ambient water to the mussel bed and will reach a maximum value beyond which no further increase will occur with increased population density. 6. It is concluded that the direct grazing impact of zebra mussel on phytoplankton in thermally stratified lakes is much less effective than predicted from simple filtration rate estimation.     

Physical habitat of zebra mussel (Dreissena polymorpha) in the lower Ebro River (Northeastern Spain): influence of hydraulic parameters in their distribution

Dreissena polymorpha (zebra mussel) is a freshwater bivalve mollusc and has been present for more than one decade in Spain. The zebra mussel causes serious ecological and socioeconomic impacts in areas where they settle. Our research aims to analyze its hydraulic habitat, developing physical models for this species, which indicates its preference and optimal microhabitat requirements. To get it, a survey was completed in a lotic reach in the lower Ebro River (Tarragona, Spain). Habitat suitability curves for the variables include water velocity, depth, Froude number, velocity/depth ratio, shear stress and shear stress ratio (RSS). In addition, interactive effects between hydraulic parameters on habitat selection and its use were studied and bivariate habitat models were developed. A close relationship was observed between the D. polymorpha presence and mean flow velocity combined with depth, Froude number and RSS. Suitable habitat requirements for Dreissena were detected in areas with stable river beds under high flows, and velocity below 1.2 m/s with a depth of less than 5 m under regular flows. Information about zebra mussel preferences may enhance the design of monitoring programmes and the integrated control management of this invasive mussel.     

THE BITTERLING–MUSSEL COEVOLUTIONARY RELATIONSHIP IN AREAS OF RECENT AND ANCIENT SYMPATRY

Host–parasite relationships are often characterized by the rapid evolution of parasite adaptations to exploit their host, and counteradaptations in the host to avoid the costs imposed by parasitism. Hence, the current coevolutionary state between a parasite and its hosts is predicted to vary according to the history of sympatry and local abundance of interacting species. We compared a unique reciprocal coevolutionary relationship of a fish, the European bitterling (Rhodeus amarus) and freshwater mussels (Unionidae) between areas of recent (Central Europe) and ancient (Turkey) sympatry. Bitterling parasitize freshwater mussels by laying their eggs in the gills of mussel and, in turn, mussel larvae (glochidia) parasitize the fish. We found that all bitterling from both regions avoided one mussel species. Preferences among other mussel species tended to be related to local mussel abundance rather than duration of sympatry. Individual fish were not consistent in their oviposition choices, precluding the evolution of host‐specific lineages. Mussels were demonstrated to have evolved strong defenses to bitterling parasitism in the area of ancient sympatry, but have no such defenses in the large areas of Europe where bitterling are currently invasive. Bitterling avoided glochidia infection irrespective of the duration of sympatry.