Lyubov E. Burlakova

Changes in global economies and trade: the potential spread of exotic freshwater bivalves.

The globalization of economies and trade have facilitated the spread of exotic species including the five most important freshwater suspension feeding invaders Dreissena polymorpha, D. bugensis, Corbicula fluminea, C. fluminalis, and Limnoperna fortunei. We suggest that the spread of these exotic species has not been a continuous process, but rather punctuated by periods of rapid long distance spread (jump), during which species greatly expanded their geographic ranges. Each jump has been associated with changes in the tempo of some human activity, such as the construction of shipping canals for trade, building of reservoirs for water storage and power production, political boundary changes or changes in political systems, which affected the position or permeability of national borders, human migration, changes in the mode and volume of international trade, or recent industrial practices and environmental laws. We hypothesize that the rate of spread of exotic species depends on the spatial scale of spread and may be accelerated or slowed by various human activities. In general, aquatic exotic species may quickly spread along connected waterways in a new continent they invade and soon reach their maximum range (continental scale). However, it will take much longer to colonize all isolated regions (regional scale) and longer still to spread to all isolated lakes and river systems (local scale). The difference in the rate of colonization across scales may be several orders of magnitude.

Impacts of Zebra Mussels on Aquatic Communities and their Role as Ecosystem Engineers

Zebra mussels (Dreissena polymorpha) are not only an extremely aggressive invasive species, often dominating water bodies they invade, they are also very effective ecosystem engineers, altering the environments they invade. They are effective engineers, altering both ecosystem structure and function. They change existing and provide new habitat for other organisms, affect trophic interactions and the availability of foods for both pelagic species and other benthic species, and they affect the rates of other ecosystem processes including mineralization of nutrients, oxygen availability and sedimentation rates. These physical impacts on the environment feedback directly to other species that interact with or are impacted by zebra mussels, or indirectly through food chains, disturbance, succession, or other longer-term community and ecosystem processes.

THE INVASIVE BIVALVES DREISSENA POLYMORPHA AND LIMNOPERNA FORTUNEI: PARALLELS, CONTRASTS, POTENTIAL SPREAD AND INVASION IMPACTS

Abstract We contrast ecological and life history traits of the well studied freshwater invader, the zebra mussel (Dreissena polymorpha), with the lesser known invasive golden mussel (Limnoperna fortunei) to compare salient biological traits and environmental limits, and to predict the potential spread and ecosystem impacts of L. fortunei in areas where it is introduced. Both species are sessile, byssate bivalves with a planktonic larval stage and extremely high reproductive capacity. For both species adults attain much higher biomass in waterbodies they invade than all of the native invertebrates combined, and they create substrate complexity otherwise not found in freshwater systems. Both are very active suspension feeders, greatly enhance benthic-pelagic coupling, and act as effective ecosystem engineers. Although taxonomically unrelated, their ecosystem impacts are surprisingly similar and follow from the novel ecological niche they share, rather than being species specific. The golden mussel has broader environmental tolerances and therefore may be a much more successful invader than D. polymorpha in regions dominated by acidic, soft and contaminated waters. In the near future L. fortunei may colonize the southern and central parts of North America, much farther north than has been previously predicted. Although to date the zebra mussel is considered the most aggressive freshwater invader, soon many waterbodies may receive another, even more aggressive invader.

Invaders are not a random selection of species

We assembled information on 119 species of freshwater macroinvertebrate invaders in North America and Europe, and compared them to all native freshwater species in North America and Europe. We tested whether the invaders were a random or selected group among taxa (phylum or class), water quality requirements, and feeding habit. We found that freshwater macroinvertebrate invaders are not a random selection of species, and are over-represented by molluscs and crustaceans, while taxa richness of native communities are dominated by insects. Over 35% of native species of aquatic invertebrates in North America are only able to live in areas with excellent or very good water quality, and are intolerant of organic pollution. In contrast, all invaders are tolerant of at least moderate amounts of organic pollution. There was a significant difference in the distribution of feeding habits between native species and invaders: collector-filterers (including suspension feeders) were 2.5–3 times more abundant, and predators were 3–4 times less abundant among invaders than among native invertebrates. The ongoing spread of exotic species affects the biodiversity of selected taxa, shifts communities toward greater tolerance of organic pollution and increases the numbers of suspension feeders, thereby enhancing benthic pelagic coupling in waterbodies with high densities of invaders. Because these processes are very similar in Europe and North America, we suggest that the observed patterns may have a common global effect.

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

Freshwater mussels of the Order Unionida provide important ecosystem functions and services, yet many of their populations are in decline. We comprehensively review the status of the 16 currently recognized species in Europe, collating for the first time their life‐history traits, distribution, conservation status, habitat preferences, and main threats in order to suggest future management actions. In northern, central, and eastern Europe, a relatively homogeneous species composition is found in most basins. In southern Europe, despite the lower species richness, spatially restricted species make these basins a high conservation priority. Information on freshwater mussels in Europe is unevenly distributed with considerable differences in data quality and quantity among countries and species. To make conservation more effective in the future, we suggest greater international cooperation using standardized protocols and methods to monitor and manage European freshwater mussel diversity. Such an approach will not only help conserve this vulnerable group but also, through the protection of these important organisms, will offer wider benefits to freshwater ecosystems.

The Impact of Dreissena polymorpha (Pallas) Invasion on Unionid Bivalves

Dreissena polymorpha, the zebra mussel, is one of the most aggressive and important invading aquatic species world wide. Its spread has followed the path of human activity, initially following human constructed canals connecting the Black Sea and Baltic Sea basins. One consequence of this invasion is the impact of zebra mussels on native bivalves. Overgrowth by Dreissena can cause a dramatic decrease in unionid density. The extent of this effect is determined by several factors including Dreissena density, time since invasion by Dreissena, biomass of attached Dreissena, and type of bottom sediments (sand versus silt). We found a correlation between overall Dreissena density and the number of zebra mussels per overgrown unionid, and between Dreissena density and the ratio of the mass of attached zebra mussels to the mass of the host unionid. The extensive overgrowth of unionids by Dreissena, resulting in mass mortality, is characteristic of periods of rapid population growth, when Dreissena invade a new waterbody.

Zebra versus quagga mussels: a review of their spread, population dynamics, and ecosystem impacts

Dreissena polymorpha (zebra mussel) and D. rostriformis bugensis (quagga mussel) continue to spread in Europe and in North America, and have large ecological and economic impacts where they invade. Today many more waterbodies are invaded by zebra mussels, and therefore the extent of their impact is greater than that of quagga mussels. Both species provide additional space and food for invertebrates in the littoral zone, increasing their diversity and density. In contrast, in the profundal zone, quagga mussels may compete for space and food resources with benthic invertebrates, decreasing their diversity and density. The system-wide effect of dreissenids depends on water mixing rates, lake morphology, and turnover rates. Because quagga mussels are found in all regions of a lake, and form larger populations, they may filter larger volumes of water and may have greater system-wide effects, especially in deep lakes, than zebra mussels, which are restricted to shallower portions of lakes. Shortly after initial invasion, as populations increase, both dreissenids will have their largest effects on communities, and most of them will be direct effects. After the initial stage of invasion, impacts are less predictable, and more likely to be caused by indirect effects through changes in the ecosystem.

Landscape patterns of an aquatic invader: Assessing dispersal extent from spatial distributions

Assessing the spatial distribution of organisms across landscapes is a key step toward determining processes that produce observed patterns. The spatial distribution of an invasive aquatic mollusk, the zebra mussel (Dreissena polymorpha), was examined in two lake-rich areas (Belarus and midwestern United States) with contrasting invasion histories. Spatial distribution patterns of invaded lakes were determined using Ripleys K. Aggregation of invaded lakes was found at similar spatial extents ( 120 km in Belarus. The observed spatial extent of aggregation likely reflected the scale of secondary geographic spread, whereas the scale of long-distance dispersal events was reflected by the spatial extent of segregation. Isolated Belarus lakes were less likely to be invaded than those connected by waterways. Although one-dimensional aggregation of invaded lakes along connected Belarus waterways was not observed, nearest neighbor analysis indicated that zebra mussel dispersal occurred at distances <15 km within these waterways. Based on observed spatial pattern, we concluded that zebra mussels have not yet saturated European and North American lake landscapes, including many suitable lakes. Similar distribution patterns of invaded lakes in Belarus and North America suggest that similar processes have influenced zebra mussel spread in both landscapes.

GROWTH RATE AND LONGEVITY OF DREISSENA POLYMORPHA (PALLAS): A REVIEW AND RECOMMENDATIONS FOR FUTURE STUDY

Abstract We review the variety of methods that have been used over the last 50 y in the Former Soviet Union, Eastern and Western Europe, and recently in North America to determine growth rate and longevity in zebra mussels (Dreissena polymorpha [Pallas]). These methods include: counting annual rings, analysis of size-frequency distributions, following growth under experimental conditions and monitoring marked mussels under natural conditions, without removing them from substrate. The last method provides the most reliable data, however this is the least common method used. Dreissena polymorpha growth rates depend on water temperature, season of the year, location in the water column, food availability, oxygen concentrations, water velocity and various other environmental factors. However, it is very difficult to separate the independent effects of each of these factors, especially in natural waterbodies. Several factors may overlap and have additive or synergistic effect that makes it difficult to determine the effects of a single factor. When comparing among studies that used the same methods, we found that zebra mussels grow faster in reservoirs than in lakes. The reported longevity of D. polymorpha varies from 2–19 y and it is not clear to what extent this variation is caused by biological variability and environmental conditions and what amount of the variation is caused by the methods used to assess age and longevity.

Endosymbionts of Dreissena polymorpha (Pallas) in Belarus

Dreissena polymorpha were dissected and examined for endosymbionts from 17 waterbodies in Belarus - the country through whose waterways zebra mussels invaded Western Europe nearly two centuries ago. Fourteen types of parasites and other symbionts were observed within the mantle cavity and/or associated with internal tissues, including ciliates (Conchophthirus acuminatus, Ancistrumina limnica, and Ophryoglena sp.), trematodes (Echinostomatidae, Phyllodistomum, Bucephalus polymorphus, and Aspidogaster), nematodes, oligochaetes, mites, chironomids, and leeches. Species composition of endosymbionts differed among river basins and lake systems. The most common endosymbiont was the ciliate C. acuminatus. Its mean infection intensity varied significantly among waterbodies from 67 ± 6 to 3,324 ± 556 ciliates/mussel.