Comparing latitudinal and upstream–downstream gradients: life history traits of invasive mosquitofish

Aim Variation of life history traits along spatial gradients is poorly understood in invasive species and particularly in freshwater fish. We aimed to examine life history variation in a highly invasive fish (Gambusia holbrooki) along latitudinal and upstream–downstream river gradients and to assess the effects of age on this variation. We hypothesized similar responses in populations inhabiting environments more favourable to this species (lower latitudes and lower reaches of rivers). Location European rivers from southern Spain to southern France. Methods We sampled mosquitofish from the lowest reaches of ten river basins along 6° of latitude in the Mediterranean region and seven sites along the upstream–downstream gradient in three of the rivers. We examined abundance, population structure, size‐at‐age and other life history traits along these gradients. Results As hypothesized, lower reaches and lower latitudes both resulted in higher reproductive effort and lower body condition of mosquitofish. However, these patterns explained low per cent variation, were nonlinear and strongly depended on fish age. Independently of fish size, age groups differed in reproductive effort, in the gonadal weight–size relationship and its variation along spatial gradients. Mean size‐at‐age (or overall body size) did not vary with latitude (so the intra‐specific version of Bergmann’s rule or its converse does not apply) and in contrast increased upstream in rivers. Main conclusions Our findings suggest that for life history traits of freshwater organisms, river longitudinal variation plays a role as important as climate, with often differential effects. Our results also illustrate the poor knowledge of spatial variation of many life history traits, which precludes the understanding and prediction of biological invasions in a rapidly changing world.     

Byssal hairs in the invasive Asian freshwater bivalve Limnoperna fortunei (Mytilidae) in the Paraná River system with comments on this species in South America

We recorded, for the first time, byssal hairs in the Asian freshwater bivalve Limnoperna fortunei from the Paraná River system. We analysed the presence of hairs and their distribution on the shell in relation to habitat and shell size in 12 sites. Hairs were present in lentic habitats associated with macrophytes or organic matter, but were absent in lotic environments. The proportion of mussels with hairs was negatively correlated with current velocity. Hairs were more frequent and abundant in larger mussels. In general, the hairs are a similar length over the bivalve and almost entirely cover the shells in≥60% in lentic habitats. The projections allow L. fortunei to be camouflaged among the roots of macrophytes or coarse organic matter, assisting in avoiding visual predators. The clear-cut separation of L. fortunei populations into two different groups could be associated with phenotypic plasticity in this species.     

Morphological clines in dendritic landscapes

1. In complex landscapes such as river networks, organisms usually face spatio‐temporal heterogeneity and gradients in geomorphological, water, ecological or landscape characteristics are often observed at the catchment scale. These environmental variables determine developmental conditions for larval stages of freshwater insects and influence adult phenotypic characteristics. Environmental clines are therefore expected to generate morphological clines. Such a process has the potential to drive gradual geographical change in morphology‐dependent life history traits, such as dispersal. 2. We studied the influence of aquatic and terrestrial environmental factors on morphological variations in Calopteryx splendens across the Loire drainage. To investigate these effects we took explicitly into account the hierarchical structure of the river network. 3. We analysed eight morphological traits. Results showed significant body size variation between tributaries and the presence of a morphological cline at the drainage scale. We observed an effect of pH and water temperature on body size. Individuals in downstream sites were larger than individuals in upstream sites, and adults whose larval stages were exposed to alkaline pH and high temperatures during summer were larger. 4. Body size affects flight abilities in insects. Thus, our results suggest that morphological clines may generate an asymmetric dispersal pattern along the downstream–upstream axis, downstream populations dispersing farther than upstream ones. Such a process is expected to influence population genetic structure at the drainage scale if larval drift and floods do not balance an asymmetrical dispersal pattern of adults along the downstream–upstream gradient. To assess the influence of environmental gradients on the variation of life history traits it is important to understand the population biology of freshwater insects, and more generally of riverine organisms. It is also essential to integrate such data in conservation or restoration programmes.     

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 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.     

Speciation in ninespine stickleback: reproductive isolation and phenotypic divergence among cryptic species of Japanese ninespine stickleback

Although similar patterns of phenotypic diversification are often observed in phylogenetically independent lineages, differences in the magnitude and direction of phenotypic divergence have been also observed among independent lineages, even when exposed to the same ecological gradients. The stickleback family is a good model with which to explore the ecological and genetic basis of parallel and nonparallel patterns of phenotypic evolution, because there are a variety of populations and species that are locally adapted to divergent environments. Although the patterns of phenotypic divergence as well as the genetic and ecological mechanisms have been well characterized in threespine sticklebacks, Gasterosteus aculeatus, we know little about the patterns of phenotypic diversification in other stickleback lineages. In eastern Hokkaido, Japan, there are three species of ninespine sticklebacks, Pungitius tymensis and the freshwater type and the brackish‐water type of the P. pungitius–P. sinensis species complex. They utilize divergent habitats along coast–stream gradients of rivers. Here, we investigated genetic, ecological and phenotypic divergence among three species of Japanese ninespine sticklebacks. Divergence in trophic morphology and salinity tolerance occurred in the direction predicted by the patterns observed in threespine sticklebacks. However, the patterns of divergence in armour plate were different from those previously found in threespine sticklebacks. Furthermore, the genetic basis of plate variation may differ from that in threespine sticklebacks. Because threespine sticklebacks are well‐established model for evolutionary research, the sympatric trio of ninespine sticklebacks will be an invaluable resource for ecological and genetic studies on both common and lineage‐specific patterns of phenotypic diversification.     

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.     

Phytoplankton growth rates in the freshwater tidal reaches of the Schelde estuary (Belgium) estimated using a simple light-limited primary production model

During the course of 1996, phytoplankton was monitored in the turbid, freshwater tidal reaches of the Schelde estuary. Using a simple light-limited primary production model, phytoplankton growth rates were estimated to evaluate whether phytoplankton could attain net positive growth rates and whether growth rates were high enough for a bloom to develop. Two phytoplankton blooms were observed in the freshwater tidal reaches. The first bloom occurred in March and was mainly situated in the most upstream reaches of the freshwater tidal zone, suggesting that it was imported from the tributary river Schelde. The second bloom occurred in July and August. This summer bloom was situated more downstream in the freshwater tidal reaches and appeared to have developed within the estuary. A comparison between phytoplankton growth rates estimated using a simple primary production model and flushing rate of the water indicated that no net increase in phytoplankton biomass was possible in March while phytoplankton could theoretically increase its biomass by 20% per day during summer. Chlorophyllaconcentrations at all times decreased strongly at salinities between 5–10 psu. This decline was ascribed to a combination of salinity stress and light limitation. Phytoplankton biomass and estimated annual net production were much higher in the freshwater tidal zone compared to the brackish reaches of the estuary (salinity > 10 psu) despite mixing depth to euphotic depth ratios being similar. Possible reasons for this high production include high nutrient concentrations, low zooplankton grazing pressure and import of phytoplankton blooms from the tributary rivers.     

Population structure,shell morphology,age and condition of the freshwater mussel Hyridella menziesi (Unionacea: Hyriidae) from seven lake and river sites in the Waikato River system

Populations of the freshwater mussel Hyridella menziesi were compared from six lakes and one river site along the Waikato River system, North Island, New Zealand. Estimated densities ranged from several hundred per square metre in Lake Taupo to less than one per square metre in Lake Karapiro. Length-frequencies were similar at all sites, with unimodal distributions and a notable absence of any individuals less than 20 mm long. The extent of shell erosion varied from sites where most mussels had less than 1% of the right valve affected (i.e., Ohakuri & Karapiro), to Lake Aratiatia where most shells had erosion over 1–20% of their surface. Taupo and Ohakuri mussels showed an unusually high incidence (over 50%) of internal shell abnormalities (including dulling, deformities and protuberances). Canonical discrimination based on shell length, height, width and weight clearly separated Taupo and Ohakuri mussels from all other sites, and ANCOVA on morphological characteristics confirmed significant differences between sites. Physical condition indices (based on dry flesh weight per shell weight and shell volume) showed an increasing downstream trend, but this was not significantly correlated with food availability (i.e., chlorophyll a concentration) because of unexpectedly low condition in lakes Ohakuri and Karapiro. Length v. age relationships contained considerable scatter and did not show between-site differences. Measured differences in external erosion, internal shell abnormalities and morphology were attributed to physical, chemical and biological factors. Use of mussels for environmental monitoring must take these background differences into account.     

The site of settlement indicates commensalism between bluemussel and its epibiont

Summary The site of settlement of barnacles (Balanus improvisus) attached on shells of bluemussels (Mytilus edulis) was mapped from a sample of mussels collected in the Baltic Sea. Most barnacles had settled near the siphonal apertures of the mussel. An experiment was made to measure the disadvantages and advantages that living in close association brings to barnacles and mussels. The barnacles on shells of living mussels were shown to grow significantly faster than those on empty mussel shells. Presence of barnacles had no effects on growth of mussels. The two-species association under study was demonstrated to be a case of commensalism.     

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.     

Evidence of phenotypic plasticity in the response of unionid mussels to turbidity

1. Increases in total suspended solids (TSS) reduce feeding and reproductive success of unionid mussels, but mussels in turbid rivers are less affected than those in clear rivers, probably due to differences in gill and palp morphology. This study was designed to determine whether the differences observed between adult mussels in populations from turbid versus clear rivers are due to phenotypic plasticity.
2. Parasitic larvae (glochidia) of Lampsilis siliquoidea (Fatmucket) obtained from a low turbidity river (TSS < 5 mg/L) were transformed on Ambloplites rupestris (Rock Bass) in the laboratory to obtain juvenile mussels for rearing under clear and turbid conditions in the laboratory (i.e. nominally 0 versus 50 mg/L suspended river sediment). Juveniles obtained were reared under these contrasting conditions until age 3–4 weeks, when they were examined in a feeding experiment under a range of TSS concentrations (0, 5, 10, 15, 20, 25 mg/L).
3. The clearance rate (volume cleared of particles per unit time) of algae by juveniles from both rearing conditions was similar for the no-TSS control and declined with increased TSS. The rate of decline was, however, lower in the group reared under turbid conditions, which is consistent with reports for adult mussels.
4. Our results indicate that differences in the clearance rate response to high TSS observed in mussels in clear and turbid rivers are probably driven in part by phenotypic plasticity. This finding provides a mechanism to explain how freshwater mussels thrive under turbid conditions as well as informs conservation efforts involving reintroduction of mussels in this highly imperilled taxon.

     

Detection of DNA damage in haemocytes of zebra mussel using comet assay

The aim of the study was to use the comet assay on haemocytes of freshwater mussel, Dreissena polymorpha Pallas, for detection of possible DNA damage after exposure to pentachlorophenol (PCP) and to evaluate the potential application of the comet assay on mussel haemocytes for genotoxicity monitoring of freshwater environment. Zebra mussels were exposed for seven days to different concentrations (10, 80, 100, 150 μg/l) of PCP and in the river Sava downstream from Zagreb municipal wastewater outlet. Significant increase in DNA damage was observed after exposure to PCP at doses of 80 μg/l and higher and after in situ exposure in the river Sava as well. This study confirmed that the comet assay applied on zebra mussel haemocytes may be a useful tool in determining the potential genotoxicity of water pollutants.     

Linking genotype to phenotype in a changing ocean: inferring the genomic architecture of a blue mussel stress response with genome‐wide association

A key component to understanding the evolutionary response to a changing climate is linking underlying genetic variation to phenotypic variation in stress response. Here, we use a genome‐wide association approach (GWAS) to understand the genetic architecture of calcification rates under simulated climate stress. We take advantage of the genomic gradient across the blue mussel hybrid zone (Mytilus edulis and Mytilus trossulus) in the Gulf of Maine (GOM) to link genetic variation with variance in calcification rates in response to simulated climate change. Falling calcium carbonate saturation states are predicted to negatively impact many marine organisms that build calcium carbonate shells – like blue mussels. We sampled wild mussels and measured net calcification phenotypes after exposing mussels to a ‘climate change’ common garden, where we raised temperature by 3°C, decreased pH by 0.2 units and limited food supply by filtering out planktonic particles >5 μm, compared to ambient GOM conditions in the summer. This climate change exposure greatly increased phenotypic variation in net calcification rates compared to ambient conditions. We then used regression models to link the phenotypic variation with over 170 000 single nucleotide polymorphism loci (SNPs) generated by genotype by sequencing to identify genomic locations associated with calcification phenotype, and estimate heritability and architecture of the trait. We identified at least one of potentially 2–10 genomic regions responsible for 30% of the phenotypic variation in calcification rates that are potential targets of natural selection by climate change. Our simulations suggest a power of 13.7% with our study's average effective sample size of 118 individuals and rare alleles, but a power of >90% when effective sample size is 900.     

Muskrat predation on a diverse unionid mussel community: impacts of prey species composition,size and shape

1. The muskrat, Ondatra zibethicus, is a predator known to affect species composition, population size and age structure of freshwater unionid mussel communities. Muskrats leave large piles of dead shells (middens) on the edges of rivers, lakes and streams. We compared the species composition and size structure of shells collected from muskrat middens to the nearby live unionid community in the lower Licking River (Kentucky, USA). 2. Like previous studies, our results suggest that muskrats are both size‐selective and species‐specific predators; however, our results indicate that mussel shape is also an important factor. 3. We generated a shape metric (‘cubocity’) sensitive to the overall shape of the mussel. Species with relatively lower cubocity values (around 0.85) are plate‐like or spike‐like, while mussels with more cuboidal shells have higher cubocity values (near 1.0). 4. Our results suggest muskrats prefer cuboidal mussels and generally avoid spike‐shaped mussels. The endangered fanshell, Cyprogenia stegaria, was the most favoured prey; the fanshell’s relative size and shape appear to make it particularly vulnerable to muskrats. 5. We believe the predictive capabilities of this shape metric will be of benefit to those who monitor and manage threatened mussel populations.     

Predation on exotic zebra mussels by native fishes: effects on predator and prey

SUMMARY 1. Exotic zebra mussels, Dreissena polymorpha, occur in southern U.S. waterways in high densities, but little is known about the interaction between native fish predators and zebra mussels. Previous studies have suggested that exotic zebra mussels are low profitability prey items and native vertebrate predators are unlikely to reduce zebra mussel densities. We tested these hypotheses by observing prey use of fishes, determining energy content of primary prey species of fishes, and conducting predator exclusion experiments in Lake Dardanelle, Arkansas. 2. Zebra mussels were the primary prey eaten by 52.9% of blue catfish, Ictalurus furcatus; 48.2% of freshwater drum, Aplodinotus grunniens; and 100% of adult redear sunfish, Lepomis microlophus. Blue catfish showed distinct seasonal prey shifts, feeding on zebra mussels in summer and shad, Dorosoma spp., during winter. Energy content (joules g⁻¹) of blue catfish prey (threadfin shad, Dorosoma petenense; gizzard shad, D. cepedianum; zebra mussels; and asiatic clams, Corbicula fluminea) showed a significant species by season interaction, but shad were always significantly greater in energy content than bivalves examined as either ash-free dry mass or whole organism dry mass. Fish predators significantly reduced densities of large zebra mussels (>5 mm length) colonising clay tiles in the summers of 1997 and 1998, but predation effects on small zebra mussels (≤5 mm length) were less clear. 3. Freshwater drum and redear sunfish process bivalve prey by crushing shells and obtain low amounts of higher-energy food (only the flesh), whereas blue catfish lack a shell-crushing apparatus and ingest large amounts of low-energy food per unit time (bivalves with their shells). Blue catfish appeared to select the abundant zebra mussel over the more energetically rich shad during summer, then shifted to shad during winter when shad experienced temperature-dependent stress and mortality. Native fish predators can suppress adult zebra mussel colonisation, but are ultimately unlikely to limit population density because of zebra mussel reproductive potential.     

Horizontal variation of biomass and C:N:P ratios of benthic algae in lakes

The horizontal variation of chlorophyll a (chl a) and C:N:P (carbon:nitrogen:phosphorus) ratio was estimated for benthic algal communities attached to living substrates (mussels and macrophytes) and to rocks and stones in three lakes of different trophy. Samples were taken in a nested hierarchical design with replicates separated by several cm, dm, 10 m, and km. The observed horizontal variation of chl a, C:N, and C:P ratios did not differ for horizontal scales, substrates, or lakes. Although the investigated lakes were quite unlike regarding nutrient status, light regime, or morphology, the patchiness was similar in all lakes. Moreover, patchiness was also similar on stones, macrophytes and mussels, although those substrates differed in longevity and surface structure. Similar patchiness regardless of scale, substrate, or sampled lake, implies the possibility of using an optimal sampling design calculated for one lake and substrate also in other lakes and on other substrates.     

Ecological correlates of the distribution limits of two poeciliid species along a salinity gradient

Identifying the environmental factors responsible for the formation of a species' distribution limit is challenging because organisms interact in complex ways with their environments. However, the use of statistical niche models in combination with the analysis of phenotypic variation along environmental gradients can help to reduce such complexity and identify a subset of candidate factors. In the present study, we used such approaches to describe and identify factors responsible for the parapatric distribution of two closely‐related livebearer fish species along a salinity gradient in the lowlands of Trinidad, West Indies. The downstream distribution limits of Poecilia reticulata were strongly correlated with the brackish–freshwater interface. We did not observe significant phenotypic variation in life‐history traits for this species when comparing marginal with more central populations, suggesting that abrupt changes in conditions at the brackish–freshwater interface limit its distribution. By contrast, Poecilia picta was present across a wide range of salinities, although it gradually disappeared from upstream freshwater localities. In addition, P. picta populations exhibited an increase in offspring size in localities where they coexist with P. reticulata, suggesting a role for interspecific competition. The parapatric distribution of these two species, suggests that P. reticulata distributions are limited by an abiotic factor (salinity), whereas P. picta is limited by a biotic factor (interspecific competition). Similar parapatric patterns have been previously described in other systems, suggesting they might be a common pattern in nature. © 2013 The Linnean Society of London     

Indicator bacteria in freshwater and marine molluscs

The freshwater mussel Anodonta cygnea and four marine shellfish (mussels, Mytilus edulis; cockles, Cerastoderma edule; clams, Mya arenaria; Scrobicularia plana) from a total of six sites were surveyed for Escherichia coli, Clostridium perfringens, faecal streptococci, 25 and 37 °C coliforms, 25 °C and 37 °C total viable numbers and fluorescent pseudomonads. The A. cygnea from an urban lake contained greater numbers of the faecal indicator bacteria than animals from a rural lake. There were also differences in the other bacterial counts and these were discussed with respect to bacterial parameter and animal characteristics. When freshwater mussels were transferred from the city site to the rural site for 24 h the load of faecal indicator bacteria was eliminated or significantly reduced. Other bacterial types took longer to become stabilised. Loss of indicator bacteria from Anodonta was also demonstrated using cleansing in the laboratory. Very high bacterial numbers were found in some marine molluscs notably Scrobicularia plana and most shellfish contained significant numbers of the three faecal indicator bacteria at every sampling occasion. The relationship between bacterial types was discussed and it was concluded that in both freshwater and marine animals the bacterial numbers were determined more by sampling site than by species of shellfish.     

Sexual,habitat‐constrained and parasite‐induced dimorphism in the shell of a freshwater mussel (Anodonta anatina,Unionidae)

Intraspecific trends in freshwater mussel (unionoid) shells that are consistently associated with differences in the mussels' sex and/or parasitic infestation can potentially be used to reconstruct sex ratios or parasitic levels of modern and ancient unionoid populations. In contrast to morphological patterns within mammal species, such dimorphic trends within unionoid species are, however, poorly understood. This study investigates, for the first time, to what extent sex, trematode infection and indirect habitat effects determine shell morphology in the freshwater mussel Anodonta anatina. Three of the five study populations displayed significant sexual shell width dimorphism. Here, shells of females were significantly wider than males, probably as a result of altered shell growth to accommodate marsupial gills. In two of these populations, female shells were additionally significantly thinner than those of males, which could be a result of resource depletion by offspring production. Two other A. anatina populations showed no significant dimorphic patterns, and our results indicate that this interpopulational difference in the degree of sexual dimorphism may reflect the overarching effect of habitat on morphology. Thus, populations in the most favourable habitats exhibit faster growth rates, attain larger maximum sizes and produce more offspring, which results in more swollen gills and consequently more inflated shells of gravid females compared to less fecund populations. None of the populations showed any evidence for sexual dimorphism in overall size, growth rate, sagittal shape and density of shells. In addition to sexual dimorphisms, infestation by bucephalid trematode parasites (Rhipidocotyle sp.) significantly altered sagittal and lateral shell shape of A. anatina in one of the populations, with infected specimens growing wider and more elongated. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.