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.     

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.     

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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Relative influence of variables at multiple spatial scales on stream macroinvertebrates in the Northern Lakes and Forest ecoregion, U.S.A.

• 1 We used 94 sites within the Northern Lakes and Forests ecoregion spanning Minnesota, Wisconsin and Michigan to identify environmental variables at the catchment, reach and riparian scales that influence stream macroinvertebrates. Redundancy analyses (RDA) found significantly influential variables within each scale and compared their relative importance in structuring macroinvertebrate assemblages.
• 2 Environmental variables included landcover, geology and groundwater delivery estimates at the catchment scale, water chemistry, channel morphology and stream habitat at the reach scale, and landcover influences at three distances perpendicular to the stream at the riparian scale. Macroinvertebrate responses were characterised with 22 assemblage attributes, and the relative abundance and presence/absence of 66 taxa.
• 3 Each scale defined macroinvertebrates along an erosional to depositional gradient. Wisconsin's macroinvertebrate index of biotic integrity, Ephemeroptera–Plecoptera–Trichoptera taxa and erosional taxa corresponded with forest streams, whereas organic pollution tolerant, Chironomidae and depositional taxa corresponded with wetland streams. Reach scale analyses defined the gradient similarly as dissolved oxygen and wide, shallow channels (erosional) opposed instream macrophytes and pool habitats (depositional). Riparian forests within 30 m of the stream coincided with an erosional assemblage and biotic integrity.
• 4 Next, we combined all significant environmental variables across scales to compare the relative influence of each spatial scale on macroinvertebrates. Partial RDA procedures described how much of the explained variance was attributable to each spatial scale and each interrelated scale combination.
• 5 Our results appeared consistent with the concept of hierarchical functioning of scale in which large‐scale variables restrict the potential for macroinvertebrate traits or taxa at smaller spatial scales. Catchment and reach variables were equally influential in defining assemblage attributes, whereas the reach scale was more influential in determining relative abundance and presence/absence.
• 6 Ultimately, comprehending the relative influence of catchment and reach scale properties in structuring stream biota will assist prioritising the scale at which to rehabilitate, manage and derive policies for stream ecosystem integrity.

     

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.     

Importance of scale,land‐use,and stream network properties for riparian plant communities along an urban gradient

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Greater macroinvertebrate diversity and freshwater mussel density in meander margins of an Amazon river

Meanders are complex aquatic environments exhibiting different flow and sediment characteristics that influence macroinvertebrate distribution. Differences in macroinvertebrate composition, freshwater mussel density (mainly Castalia ambigua Lamarck, 1819) and habitat variables were investigated, using uni‐ and multivariate analyses, in two zones, the margin and thalweg, of the meanders of a 7‐km stretch of a morphologically unaltered tropical alluvial lowland river. Clear differences were found between meander zones, with greater taxonomic and functional diversity of macroinvertebrates and greater freshwater mussel densities in the meander margin. A total of 12 families of macroinvertebrates were exclusive to the meander margin, and macroinvertebrate indicators of zones were Philopotamidae and Leptoceridae (meander margin) and Chironomidae and Elmidae (meander thalweg). In the meander margin, there were no differences in macroinvertebrate abundance, and taxonomic and functional group diversity among areas with low, medium and high mussel density. Macroinvertebrate abundance did not vary between zones, but in both, abundance was associated with lower pH and higher electrical conductivity. Low shear velocity, which stabilises the river bed, high organic matter content and percentage silt in the meander margin were associated with higher taxonomic and functional macroinvertebrate diversity, as well as the presence of freshwater mussels. Natural unaltered meanders are spatially heterogeneous in both habitat and biodiversity and, similar to restored meanders, support greater macroinvertebrate taxonomic and functional diversity, as well as maintain freshwater mussel beds, mainly by stabilising the river bed. The meander margin may also be important for attracting mussel host fish that feed on macroinvertebrates and aid mussel dispersal. Thus, the meander margins, and other similar riparian habitats with low shear velocity, for example, side channels and pools, have potentially high conservation value and should be afforded due protection.     

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.     

Relationships between land use, spatial scale and stream macroinvertebrate communities

1. The structure of lotic macroinvertebrate communities may be strongly influenced by land‐use practices within catchments. However, the relative magnitude of influence on the benthos may depend upon the spatial arrangement of different land uses in the catchment. 2. We examined the influence of land‐cover patterns on in‐stream physico‐chemical features and macroinvertebrate assemblages in nine southern Appalachian headwater basins characterized by a mixture of land‐use practices. Using a geographical information system (GIS)/remote sensing approach, we quantified land‐cover at five spatial scales; the entire catchment, the riparian corridor, and three riparian ‘sub‐corridors’ extending 200, 1000 and 2000 m upstream of sampling reaches. 3. Stream water chemistry was generally related to features at the catchment scale. Conversely, stream temperature and substratum characteristics were strongly influenced by land‐cover patterns at the riparian corridor and sub‐corridor scales. 4. Macroinvertebrate assemblage structure was quantified using the slope of rank‐abundance plots, and further described using diversity and evenness indices. Taxon richness ranged from 24 to 54 among sites, and the analysis of rank‐abundance curves defined three distinct groups with high, medium and low diversity. In general, other macroinvertebrate indices were in accord with rank‐abundance groups, with richness and evenness decreasing among sites with maximum stream temperature. 5. Macroinvertebrate indices were most closely related to land‐cover patterns evaluated at the 200 m sub‐corridor scale, suggesting that local, streamside development effectively alters assemblage structure. 6. Results suggest that differences in macroinvertebrate assemblage structure can be explained by land‐cover patterns when appropriate spatial scales are employed. In addition, the influence of riparian forest patches on in‐stream habitat features (e.g. the thermal regime) may be critical to the distribution of many taxa in headwater streams draining catchments with mixed land‐use practices.     

Hard rock versus soft bottom: the fauna associated with intertidal mussel beds on hard bottoms along the coast of Chile,and considerations on the functional role of mussel beds

The fauna associated with hard bottom mussel beds along the exposed Pacific coast of Chile was examined. The abundance of adult (>10 mm body length) purple mussels Perumytilus purpuratus varied between 32 and 75 individuals per 50 cm², and their biomass between 4.8 and 8.6 g AFDW per 50 cm² at eight sampling sites between Arica (18°S) and Chiloé (42°S). At all sampling sites, the associated fauna was dominated by suspension-feeding organisms (cirripeds, spionid and sabellid polychaetes, a small bivalve) followed by grazing peracarids and gastropods. Predators and scavengers also reached high abundances while deposit- and detritus-feeding organisms were of minor importance. The majority of organisms associated with these hard bottom mussel beds feed on resources obtained from the water column or growing on the mussels rather than on materials deposited by the mussels. This is in contrast to the fauna associated with mussel beds on soft bottoms, which comprises many species feeding on material accumulated by mussels (faeces and pseudofaeces) and deposited within the mussel bed. Many of the organisms dwelling between mussels both on hard bottoms and on soft bottoms have direct development, but organisms with pelagic development also occur abundantly within mussel beds. We propose that species with direct development are disproportionately favoured by the structurally complex habitat with diverse interstitial spaces between the mussels, which provides ample shelter for small organisms. We conclude that mussels on hard-bottoms primarily provide substratum for associated fauna while mussels on soft bottoms provide both substratum and food resources. Electronic Publication     

Limited Effects of Large‐Scale Riparian Restoration on Seed Banks in Agriculture

I examined the effect of riparian forest restoration on plant abundance and diversity, including weed species, on agricultural lands along the Sacramento River in California (United States). Riparian forest restoration on the Sacramento River is occurring on a large‐scale, with a goal of restoring approximately 80,000 ha over 160 km of the river. In multiuse habitats, such as the Sacramento River, effects of adjoining habitat types and movement of species across these habitats can have important management implications in terms of landscape‐scale patterns of species distributions. Increased numbers of pest animals and weeds on agricultural lands associated with restored habitats could have negative economic impacts, and in turn affect support for restoration of natural areas. In order to determine the distribution and abundance of weeds associated with large‐scale restoration, I collected seed bank soil samples on orchards between 0 and 5.6 km from adjacent restored riparian, remnant riparian, and agricultural habitats. I determined the abundance, species richness, and dispersal mode of plant species in the seed bank and analyzed these variables in terms of adjacent habitat type and age of restored habitat. I found that agricultural weed species had higher densities at the edge of restored riparian habitat and that native plants had higher densities adjacent to remnant riparian habitat. Weed seed abundance increased significantly on walnut farms adjacent to restored habitat with time since restored. I supply strong empirical evidence that large areas of natural and restored habitats do not lead to a greater penetration of weed species into agricultural areas, but rather that weed penetration is both temporally and spatially limited.     

Habitat persistence for sedentary organisms in managed rivers: the case for the federally endangered dwarf wedgemussel (Alasmidonta heterodon) in the Delaware River

1. To manage the environmental flow requirements of sedentary taxa, such as mussels and aquatic insects with fixed retreats, we need a measure of habitat availability over a variety of flows (i.e. a measure of persistent habitat). Habitat suitability measures in current environmental flow assessments are measured on a ‘flow by flow’ basis and thus are not appropriate for these taxa. Here, we present a novel measure of persistent habitat suitability for the dwarf wedgemussel (Alasmidonta heterodon), listed as federally endangered in the U.S.A., in three reaches of the Delaware River. 2. We used a two‐dimensional hydrodynamic model to quantify suitable habitat over a range of flows based on modelled depth, velocity, Froude number, shear velocity and shear stress at three scales (individual mussel, mussel bed and reach). Baseline potentially persistent habitat was quantified as the sum of pixels that met all thresholds identified for these variables for flows ≥40 m³ s^?1, and we calculated the loss of persistently suitable habitat by sequentially summing suitable habitat estimates at lower flows. We estimated the proportion of mussel beds exposed at each flow and the amount of change in the size of the mussel bed for one reach. 3. For two reaches, mussel beds occupied areas with lower velocity, shear velocity, shear stress and Froude number than the reach average at all flows. In the third reach, this was true only at higher flows. Together, these results indicate that beds were possible refuge areas from the effects of these hydrological parameters. Two reaches showed an increase in the amount of exposed mussel beds with decreasing flow. 4. Baseline potentially persistent habitat was less than half the areal extent of potentially suitable habitat, and it decreased with decreasing flow. Actually identified beds and modelled persistent habitat showed good spatial overlap, but identified beds occupied only a portion of the total modelled persistent habitat, indicating either that additional suitable habitat is available or the need to improve habitat criteria. At one site, persistent beds (beds where mussels were routinely collected) were located at sites with stable substratum, whereas marginal beds (beds where mussels were infrequently collected or that were lost following a large flood event) were located in scoured areas. 5. Taken together, these model results support a multifaceted approach, which incorporates the effects of low and high flow stressors, to quantify habitat suitability for mussels and other sedentary taxa. Models of persistent habitat can provide a more holistic environmental flow assessment of rivers.     

Effects of substratum stability on diversity of stream invertebrates during baseflow at two spatial scales

1. The effects of substratum stability on the diversity of stream invertebrates were assessed at two spatial scales in a Japanese stream during baseflow, from May to June 1998. Deposition and erosion were examined separately as distinct elements of substratum stability by a newly developed method using small steel pins. Stream invertebrates were sampled after 28 days of measurement of substratum stability. We also measured physical environmental variables, current velocity and depth, and food resource parameters including periphyton and particulate organic matter (POM) standing crops.
2. At the scale of the habitat patch, the effects of substratum stability on invertebrates were overwhelmed by those of POM standing crop. Moreover, higher taxon richness at high abundance may simply result from a higher likelihood of more taxa being included in samples. Therefore, this small scale revealed no role for substratum stability in explaining spatial pattern of community diversity.
3. At the reach scale ( n =10), taxon richness and evenness peaked at an intermediate level of deposition, whereas invertebrate abundance did not show any significant relationship. This result, and the pattern of relative abundance of common taxa, implies that the diversity of stream invertebrates was determined by subtle substratum movements and by the habitat preference of each taxon.
4. The difference in the determinant of community parameters between the habitat patch and the reach affirm the importance of a cross-scale analysis to choose an appropriate spatial scale for investigating the community structure of stream invertebrates. Prominent effects of substratum stability, particularly the deposition of substratum particles, during baseflow suggest that subtle and constant movement of small substratum particles can contribute to determine the diversity of stream invertebrates.     

Impacts of land use at the catchment scale constrain the habitat benefits of stream riparian buffers

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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.     

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.     

Stream macroinvertebrate assemblage uniformity and drivers in a tropical bioregion

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Impacts of an invasive willow (Salix × rubens) on riparian bird assemblages in south‐eastern Australia

We explored how a woody plant invader affected riparian bird assemblages. We surveyed 15 200‐m‐long transects in riparian zones in a much‐changed landscape of eastern Victoria, Australia. Abundance, species‐richness, foraging‐guild richness and composition of birds were compared in transects in three habitat types: (i) riparian zones dominated by the invasive willow Salix × rubens; (ii) riparian zones lined with native woody species; and (iii) riparian zones cleared of almost all woody vegetation. We also measured abundance and richness of arthropods and habitat structure to explore further the effects of food resources and habitat on the avifauna. We observed 67 bird species from 14 foraging guilds. Native riparian transects had more birds, bird species and foraging guilds than willow‐invaded or cleared transects. Habitat complexity increased from cleared to willow‐invaded to native riparian transects, as did abundance of native and woodland‐dependent birds. Native shrub and trees species had more foliage and branch‐associated arthropods than did willows, consistent with a greater abundance and variety of foraging guilds of birds dependent on this resource. Willow spread into cleared areas is unlikely to facilitate greatly native bird abundance and diversity even though habitat complexity is increased. Willow invasion into the native riparian zone, by decreasing food resources and altering habitat, is likely to reduce native bird biodiversity and further disrupt connectivity of the riparian zone.     

Species traits and environmental conditions govern the relationship between biodiversity effects across trophic levels

Changing environments can have divergent effects on biodiversity–ecosystem function relationships at alternating trophic levels. Freshwater mussels fertilize stream foodwebs through nutrient excretion, and mussel species-specific excretion rates depend on environmental conditions. We asked how differences in mussel diversity in varying environments influence the dynamics between primary producers and consumers. We conducted field experiments manipulating mussel richness under summer (low flow, high temperature) and fall (moderate flow and temperature) conditions, measured nutrient limitation, algal biomass and grazing chironomid abundance, and analyzed the data with non-transgressive overyielding and tripartite biodiversity partitioning analyses. Algal biomass and chironomid abundance were best explained by trait-independent complementarity among mussel species, but the relationship between biodiversity effects across trophic levels (algae and grazers) depended on seasonal differences in mussel species’ trait expression (nutrient excretion and activity level). Both species identity and overall diversity effects were related to the magnitude of nutrient limitation. Our results demonstrate that biodiversity of a resource-provisioning (nutrients and habitat) group of species influences foodweb dynamics and that understanding species traits and environmental context are important for interpreting biodiversity experiments.     

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.