Scale-dependent associations between native freshwater mussels and invasive Corbicula

In North America there is conflicting evidence concerning whether the invasive Asian clam, Corbicula fluminea, and native mussels (Unionidae), can successfully co-exist. One reason underlying disparate conclusions may be the different spatial scales at which data have been collected. We compared the distribution and abundance of native unionid mussels and Corbicula at two spatial scales, stream reaches and 0.25 m² patches, within one biogeographic region, the Ouachita Highlands, of the south central U.S. We found that Corbicula abundance was negatively related to native mussel abundance at small spatial scales. While Corbicula densities varied widely in patches without native mussels, and in patches where mussels occurred at low abundance, Corbicula density was never high in patches where mussels were dense. We hypothesize that the likelihood of successful Corbicula invasion decreases with increasing abundance of adult native mussels. Several mechanisms may potentially drive this pattern including lack of space for Corbicula to colonize, physical displacement by actively burrowing mussels, and locally reduced food resources in patches where native mussels are feeding. In addition, Corbicula may be unable to withstand environmental bottlenecks as readily as unionids. When patch-scale density and biomass information were pooled to represent entire stream reaches, the negative relationship between native mussels and Corbicula was no longer as apparent, and there was not a significant relationship between native mussels and Corbicula. These results point to the importance of appropriate sample scale in examining potential associations between species.     

High abundance of an invasive species gives it an outsized ecological role

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Population-specific responses to an invasive species

Predicting the impacts of non-native species remains a challenge. As populations of a species are genetically and phenotypically variable, the impact of non-native species on local taxa could crucially depend on population-specific traits and adaptations of both native and non-native species. Bitterling fishes are brood parasites of unionid mussels and unionid mussels produce larvae that parasitize fishes. We used common garden experiments to measure three key elements in the bitterling–mussel association among two populations of an invasive mussel (Anodonta woodiana) and four populations of European bitterling (Rhodeus amarus). The impact of the invasive mussel varied between geographically distinct R. amarus lineages and between local populations within lineages. The capacity of parasitic larvae of the invasive mussel to exploit R. amarus was higher in a Danubian than in a Baltic R. amarus lineage and in allopatric than in sympatric R. amarus populations. Maladaptive oviposition by R. amarus into A. woodiana varied among populations, with significant population-specific consequences for R. amarus recruitment. We suggest that variation in coevolutionary states may predispose different populations to divergent responses. Given that coevolutionary relationships are ubiquitous, population-specific attributes of invasive and native populations may play a critical role in the outcome of invasion. We argue for a shift from a species-centred to population-centred perspective of the impacts of invasions.     

Simulating selection and evolution at the community level using common garden data

A key issue in evolutionary biology is whether selection acting at levels higher than the individual can cause evolutionary change. If it can, then conceptual and empirical studies must consider how selection operates at multiple levels of biological organization. Here, we test the hypothesis that estimates of broad‐sense community heritability, HC2, can be used to predict the evolutionary response by community‐level phenotypes when community‐level selection is imposed. Using an approach informed by classic quantitative genetics, we made three predictions. First, when we imposed community‐level selection, we expected a significant change in the average phenotype of arthropod communities associated with individual tree genotypes [we imposed selection by favoring high and low NMDS (nonmetric multidimensional scaling) scores that reflected differences in arthropod species richness, abundance and composition]. Second, we expected HC2 to predict the magnitude of the community‐level response. Third, we expected no significant change in average NMDS scores with community‐level selection imposed at random. We tested these hypotheses using three years of common garden data for 102 species comprising the arthropod communities, associated with nine clonally replicated Populus angustifolia genotypes. Each of our predictions were met. We conclude that estimates of HC2 account for the resemblance among communities sharing common ancestry, the persistence of community composition over time, and the outcome of selection when it occurs at the community level. Our results provide a means for exploring how this process leads to large‐scale community evolutionary change, and they identify the circumstances in which selection may routinely act at the community level.     

Patch utilization and flower visitations by wild bees in a honey bee‐dominated,grassland landscape

Understanding habitat needs and patch utilization of wild and managed bees has been identified as a national research priority in the United States. We used occupancy models to investigate patterns of bee use across 1030 transects spanning a gradient of floral resource abundance and richness and distance from apiaries in the Prairie Pothole Region (PPR) of the United States. Estimates of transect use by honey bees were nearly 1.0 during our 3.5‐month sampling period, suggesting honey bees were nearly ubiquitous across transects. Wild bees more frequently used transects with higher flower richness and more abundant flowers; however, the effect size of the native flower abundance covariate (β^native = 3.90 ± 0.65 [1SE]) was four times greater than the non‐native flower covariate (β^non‐native = 0.99 ± 0.17). We found some evidence that wild bee use was lower at transects near commercial apiaries, but the effect size was imprecise (β^distance = 1.4 ± 0.81). Honey bees were more frequently detected during sampling events with more non‐native flowers and higher species richness but showed an uncertain relationship with native flower abundance. Of the 4039 honey bee and flower interactions, 85% occurred on non‐native flowers, while only 43% of the 738 wild bee observations occurred on non‐native flowers. Our study suggests wild bees and honey bees routinely use the same resource patches in the PPR but often visit different flowering plants. The greatest potential for resource overlap between honey bees and wild bees appears to be for non‐native flowers in the PPR. Our results are valuable to natural resource managers tasked with supporting habitat for managed and wild pollinators in agroecosystems.     

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.     

Population structure of an introduced species (Dreissena polymorpha) along a wave-swept disturbance gradient

The zebra mussel Dreissena polymorpha was introduced to North America during the mid-1980s, and is now a dominant member of many benthic communities in the lower Great Lakes. In this study, I explored the abundance, biomass, size structure and settlement of Dreissena inhabiting rocks along a wave-swept disturbance gradient near Middle Sister Island in western Lake Erie. Ten rocks were collected from quadrats at six sites along each of three transect lines oriented perpendicular to shore. Occurrence, abundance and biomass of Dreissena on smaller, movable rocks were positively associated with rock distance from shore (lake depth) and with rock area; rocks at nearshore sites supported little, if any, Dreissena, whereas those at offshore sites were heavily colonized. Mussel size distributions also differed in relation to shore distance. Large mussels (19 mm) were underrepresented or absent on rocks collected at nearshore sites, but were overrepresented at offshore locations (37 m). Settlement of larval mussels on settling pads was positively correlated with distance offshore and with time of exposure, though settlement was substantial even at a nearshore (10 m) location. Area-adjusted mussel dry mass increased more rapidly with distance offshore on large than on small rocks. Large rocks also required more force to displace and were significantly less likely to be disturbed when transplanted at the study site. Results from this study indicate that occurrence, abundance and size structure of Dreissena in nearshore waters of Lake Erie correspond with the frequency of habitat disturbance, though other factors including food limitation and larval supply may also contribute to these patterns. These patterns complement studies that established the significance of physical disturbance in other aquatic systems.     

Filtration activity of invasive mussel species under wave disturbance conditions

Self-purification is one of the most important ecosystem functions of rivers. Multiple human activities regularly impact this ecosystem service, consequently altering river morphology, hydrology, and the composition of biotic assemblages that contribute to self-purification. However, little quantitative information is available about the importance of such impacts. Hence, we tested how invasive mussel species contribute to self-purification under disturbed riverine conditions. In laboratory experiments, invasive mussel species equipped with magnetic sensors that recorded filtration activity were exposed to artificial waves of varying intensity that simulated the hydraulic effects of inland navigation. Our results suggest that invasive mussel species are more resistant to wave disturbance compared to native species, as estimated threshold values for initiating shell closure are very high (Dreissena rostriformis bugensis) or the duration (Corbicula fluminea) and degree of shell closing (D. rostriformis bugensis, C. fluminea) very low. Also we demonstrated that the invasive species D. rostriformis bugensis and C. fluminea continued filtering during wave impact, whereas Dreissena polymorpha did not behave significantly differently than previously studied native mussel species, based on the studied susceptibility parameters. Thus, D. rostriformis bugensis and C. fluminea appear to be pre-adapted to hydraulic or morphological disturbance, and may compensate against other losses regarding this important ecosystem function in rivers that are intensively used for inland navigation. However, as the dominance of invasive species in river systems may disrupt natural food webs, this compensation of filter-feeding activity may be accompanied by the loss of other ecosystem functions.     

Climate controls the distribution of a widespread invasive species: implications for future range expansion

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Habitat engineering by the invasive zebra mussel <Emphasis Type="Italic">Dreissena polymorpha</Emphasis> (Pallas) in a boreal coastal lagoon: impact on biodiversity

Habitat engineering role of the invasive zebra mussel Dreissena polymorpha (Pallas) was studied in the Curonian lagoon, a shallow water body in the SE Baltic. Impacts of live zebra mussel clumps and its shell deposits on benthic biodiversity were differentiated and referred to unmodified (bare) sediments. Zebra mussel bed was distinguished from other habitat types by higher benthic invertebrate biomass, abundance, and species richness. The impact of live mussels on biodiversity was more pronounced than the effect of shell deposits. The structure of macrofaunal community in the habitats with >10³ g/m² of shell deposits devoid of live mussels was similar to that found within the zebra mussel bed. There was a continuous shift in species composition and abundance along the gradient ‘bare sediments—shell deposits—zebra mussel bed’. The engineering impact of zebra mussel on the benthic community became apparent both in individual patches and landscape-level analyses.     

Impact of Dreissena fouling on the physiological condition of native and invasive bivalves: interspecific and temporal variations

The impact of Dreissena fouling on unionids has hardly been studied in Europe, despite the fact that in some ecosystems (e.g. Lake Balaton, Hungary) infestations of several hundreds to a thousand individuals per unionid have been observed. At present, the zebra mussel Dreissena polymorpha is a dominant species in Lake Balaton and in the last decade three other invasive bivalves were introduced, potentially increasing the pressure on native unionid survival. We examined whether the fouling of dreissenids (zebra and quagga (D. rostriformis bugensis) mussels) has a negative impact on native (Anodonta anatina, Unio pictorum and U. tumidus) and invasive (Corbicula fluminea and Sinanodonta woodiana) bivalves and whether there are any interspecific and temporal variations in fouling intensity and physiological condition measured by standard condition index and glycogen content. A significant negative impact was detected on native unionids only in July and September (no impact was detected in May), when the fouling rate was high. For invasive species, a significant negative impact was detected on S. woodiana with a high level of dressenid infestation; whereas no significant impact was detected on C. fluminea. Overall, this study confirms that Dreissena may threaten unionid species including the invasive S. woodiana, although high interspecific and temporal variations were observed. This situation should be taken into account in future ecological and conservational assessments because species respond differently to Dreissena fouling and effects seem to be more pronounced in late summer/early autumn. In addition, this study provides the first evidence that the invasive C. fluminea appear to be less vulnerable to dressenid fouling.     

Linking terrestrial and benthic estuarine ecosystems: organic matter sources supporting the high secondary production of a non-indigenous bivalve

The Asian clam, Corbicula fluminea, is among the most pervasive invasive species in freshwater ecosystems worldwide. Our objective was to study C. fluminea’s functional response in terms of feeding behavior and food selectivity, using the natural variation in organic matter (OM) sources that occur in estuarine environments. Using C and N stable isotopes, we identified and quantified the contribution of different OM sources supporting the production of C. fluminea along the salinity gradient occupied in the Minho River estuary (NW-Iberian Peninsula, Europe), where this species presently dominates the benthic macrofauna biomass. We observed a pronounced shift in the quality of OM available for C. fluminea along the estuarine mixing zone. Stable isotope analysis, POM C/N, and phytoplankton contribution estimates based on C:Chl a revealed that POM was largely comprised of terrestrial-derived OM in tidal freshwater stations (TFW) and was increasingly comprised of phytoplankton, a more palatable food source, towards the polyhaline estuary. A similar shift in the isotopic composition along the estuarine mixing zone was observed in C. fluminea, suggesting a shift in food resources. Accordingly, based on a Bayesian stable isotope mixing model, there was an upstream–downstream counter gradient in the contribution to C. fluminea biomass from terrestrial-derived OM (41–64 % in TFW) and phytoplankton (29–55 % in the brackish estuary). Although the majority of the food sources identified were filtered from the water column (70–80 %), reliance on sediment OM and microphytobenthos provided evidence for deposit feeding by C. fluminea. We conclude that C. fluminea has the ability to adapt to environments with low food quality because it can consume terrestrial-derived OM. This can be a competitive adaptation in systems with perennial low food quality such as the Minho River estuary. Moreover, its ability to couple benthic and pelagic environments and terrestrial ecosystems demonstrates a strong potential to alter food web flows in aquatic ecosystems.     

Thermodynamic examination of 1- to 5-nt purine bulge loops in RNA and DNA constructs

Bulge loops are common features of RNA structures that are involved in the formation of RNA tertiary structures and are often sites for interactions with proteins and ions. Minimal thermodynamic data currently exist on the bulge size and sequence effects. Using thermal denaturation methods, thermodynamic properties of 1- to 5-nt adenine and guanine bulge loop constructs were examined in 10 mM MgCl₂ or 1 M KCl. The ΔG37∘ loop parameters for 1- to 5-nt purine bulge loops in RNA constructs were between 3.07 and 5.31 kcal/mol in 1 M KCl buffer. In 10 mM magnesium ions, the ΔΔG° values relative to 1 M KCl were 0.47–2.06 kcal/mol more favorable for the RNA bulge loops. The ΔG37∘ loop parameters for 1- to 5-nt purine bulge loops in DNA constructs were between 4.54 and 5.89 kcal/mol. Only 4- and 5-nt guanine constructs showed significant change in stability for the DNA constructs in magnesium ions. A linear correlation is seen between the size of the bulge loop and its stability. New prediction models are proposed for 1- to 5-nt purine bulge loops in RNA and DNA in 1 M KCl. We show that a significant stabilization is seen for small bulge loops in RNA in the presence of magnesium ions. A prediction model is also proposed for 1- to 5-nt purine bulge loop RNA constructs in 10 mM magnesium chloride.     

Inversion of a part of the numerator relationship matrix using pedigree information

Background

In recent theoretical developments, the information available (e.g. genotypes) divides the original population into two groups: animals with this information (selected animals) and animals without this information (excluded animals). These developments require inversion of the part of the pedigree-based numerator relationship matrix that describes the genetic covariance between selected animals (A₂₂). Our main objective was to propose and evaluate methodology that takes advantage of any potential sparsity in the inverse of A₂₂ in order to reduce the computing time required for its inversion. This potential sparsity is brought out by searching the pedigree for dependencies between the selected animals. Jointly, we expected distant ancestors to provide relationship ties that increase the density of matrix A₂₂ but that their effect on A22-1 might be minor. This hypothesis was also tested.

Methods

The inverse of A₂₂ can be computed from the inverse of the triangular factor (T^-1) obtained by Cholesky root-free decomposition of A₂₂. We propose an algorithm that sets up the sparsity pattern of T^-1 using pedigree information. This algorithm provides positions of the elements of T^-1 worth to be computed (i.e. different from zero). A recursive computation of A22-1 is then achieved with or without information on the sparsity pattern and time required for each computation was recorded. For three numbers of selected animals (4000; 8000 and 12 000), A₂₂ was computed using different pedigree extractions and the closeness of the resulting A22-1 to the inverse computed using the fully extracted pedigree was measured by an appropriate norm.

Results

The use of prior information on the sparsity of T^-1 decreased the computing time for inversion by a factor of 1.73 on average. Computational issues and practical uses of the different algorithms were discussed. Cases involving more than 12 000 selected animals were considered. Inclusion of 10 generations was determined to be sufficient when computing A₂₂.

Conclusions

Depending on the size and structure of the selected sub-population, gains in time to compute A22-1 are possible and these gains may increase as the number of selected animals increases. Given the sequential nature of most computational steps, the proposed algorithm can benefit from optimization and may be convenient for genomic evaluations.     

Predation on invasive zebra mussel, Dreissena polymorpha, by pumpkinseed sunfish, rusty crayfish, and round goby

The enemy release hypothesis states that invasive species are successful in their new environment because native species are not adapted to utilize the invasive. If true for predators, native predators should have lower feeding rates on the invasive species than a predator from the native range of the invasive species. We tested this hypothesis for zebra mussel (Dreissena polymorpha) by comparing handling time and predation rate on zebra mussels in the laboratory by two North American species (pumpkinseed, Lepomis gibbosus, and rusty crayfish, Orconectes rusticus) and one predator with a long evolutionary history with zebra mussels (round goby, Neogobius melanostomus). Handling time per mussel (7 mm shell length) ranged from 25 to >70 s for the three predator species. Feeding rates on attached zebra mussels were higher for round goby than the two native predators. Medium and large gobies consumed 50–67 zebra mussels attached to stones in 24 h, whereas pumpkinseed and rusty crayfish consumed <11. This supports the hypothesis that the rapid spread of zebra mussels in North America was facilitated by low predation rates from the existing native predators. At these predation rates and realistic goby abundance estimates, round goby could affect zebra mussel abundance in some lakes.