Introduction: The Biology, Ecology, and Physiology ...

AMER. ZOOL., 36:239-243 (1996)

Introduction: The Biology, Ecology, and Physiology of Zebra Mussels1 JEFFREY L. RAM Department of Physiology, Wayne State University, Detroit, Michigan 48201 AND

ROBERT F. MCMAHON Center for Biological Macrofouling Research, Department of Biology, Box 19498, The University of Texas at Arlington, Arlington, Texas 76019

1 From the Symposium Biology, Ecology and Physiology of Zebra Mussels presented at the Annual Meeting of the American Society of Zoologists, 4—8 January 1995, at St. Louis, Missouri.

Arkansas Rivers, but not the Missouri River (Griffiths et al, 1991; Ram et al, 1992; McMahon, 1992; O'Neill and Dextrase, 1994) (Fig. 1). By the end of 1995, the zebra mussel had invaded waters in 20 of the 38 U.S. states east of the Rocky Mountains and the Canadian provinces of Ontario and Quebec (Fig. 1). By 1994, it was also beginning to disperse, apparently by human mediated vectors, into smaller, inland bodies of water. As of January 1996, sightings of zebra mussels have been confirmed in 54 isolated inland lakes in Illinois, Indiana, Michigan, New York, Ohio, Vermont and Wisconsin (United States National Biological Service, 1995a). A second species of freshwater, dreissenid mussel, the quagga mussel, Dreissena bugensis, has also been introduced into the Great Lakes. Initially recorded in Lake Ontario, it is spreading rapidly and now occupies the eastern basin and southern shore of Lake Erie, the southern shore of Lake Ontario and the freshwater portions of the St. Lawrence River (May and Marsden, 1992; Mills et al., 1993, 1996; Zebra Mussel Information Clearinghouse, 1995). The tendency of zebra mussels to form dense aggregates on hard surfaces causes them to have great economic impact. Its planktonic veliger larvae and translocating juveniles are entrained on intake water into man-made raw water systems where they settle and attach in large numbers. Newly settled mussels reach such high densities (up to 700,000 irr 2 , Griffiths et al., 1991) and have such rapid post-settlement growth rates (Nichols et al., 1990, 1996) that they rapidly form thick mats, occluding or blocking flow even in large diameter piping

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The papers in this issue of American Zoologist were originally presented in a symposium entitled, "The Biology, Ecology, and Physiology of Zebra Mussels," held 5 January 1995, at the American Society of Zoologists (ASZ) Annual Meeting in St. Louis, Missouri. The zebra mussel (Dreissena polymorpha), is an endemic, freshwater, European bivalve mollusc accidentally introduced into the North American Great Lakes. Introduction was most likely by release of larvae with ship ballast water into Lake St. Clair, near Detroit, Michigan in 1986 (Hebert et al., 1989). The zebra mussel's high fecundity, passively dispersed planktonic veliger larval stage, and ability to attach by proteinaceous byssal threads to boat and barge hulls, nets, buoys, and floating debris allowed it to spread rapidly throughout the lower Great Lakes and freshwater portions of the St. Lawrence River after its initial introduction (Fig. 1). It subsequently dispersed east through the Erie Canal, into the Hudson River, overland (by unintentional human mediated vectors) into the upper Susquehana River in New York, and south and west through the Illinois River into the Mississippi River near St. Louis. After entering the Mississippi River, the zebra mussel rapidly spread downstream to New Orleans, LA, and upstream to La Crosse, WI. It is now found in the lower portions of most major Mississippi River tributaries, including the Ohio, Tennessee, Cumberland, and

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(Mackie et al., 1989; McMahon, 1992; Kovalak et al., 1993; Jenner and JanssenMommen, 1993). Early North American experience with mussel infestations in the raw water systems of power stations, potable water treatment plants and industrial facilities on the Great Lakes suggested that their fouling developed more rapidly and was more severe than reported in Europe (Griffiths et al., 1989; LePage, 1993; Claudi and Mackie 1993). Thus, the zebra mussel invasion of North America will lead to increased macrofouling impacts, the eventual costs for repair, replacement and control estimated to be several billion dollars annually (Roberts, 1990; Office of Technology Assessment, United States Congress, 1993). The zebra mussel is also negatively impacting native North American freshwater biota. Extensive mortality or even complete extirpation of native North American unionid bivalve populations has been reported in habitats densely colonized by ze-

While Dreissena polymorpha is likely to have negative environmental and economical impacts on North American freshwaters, North American investigators have found that this species can also be a model organism for biological, ecological and physiological investigations. The mussels are easy to collect, maintain, and culture (Nichols, this volume) making them adaptable to a wide variety of laboratory investigations, and, because they are epifaunal and sessile, they also make excellent subjects for field ecological or environmental studies. For example, the information base on zebra mussel physiology has been greatly increased by recent North American studies of ion transport (Horohov et al., 1992; Dietz et al., 1996), ciliary function (Silverman et al., 1996), physiological resistance and capacity adaptations (McMahon, et al., 1993 and McMahon, 1996), and reproductive/ developmental mechanisms (Ram et al.,


FIG. 1. Map showing the distribution of the zebra mussel, Dreissena polymorpha in North America as of October, 1995. Dotted lines show the borders of U.S. states and Canadian provinces and solid lines indicate major rivers and lakes. Solid circles are confirmed reports of individual zebra mussel populations while darkened areas indicate regions were mussel populations have become contiguous (data from United States National Biological Service, 1995b).

bra mussels. The mussels settle on the posterior portions of unionid shells extending above the substratum surface. Massive infestations cause unionids to be dislodged from the substratum and/or experience starvation as zebra mussels filter seston food from the unionid's inhalant flow (Mackie, 1993; Schloesser et al., 1996). Suspensionfeeding by dense zebra mussel populations may also result in density reductions of phytoplankton and suspended inorganic matter (Maclsaac et al., 1992). The resulting increase in water clarity can lead to increases in the density and biomass of benthic macrophytes and animals (Hebert et al., 1991). Such diversion of energy flow by zebra mussels from pelagic into benthic food chains has been hypothesized to negatively impact pelagic fish populations while increasing demersal fish productivity (Karnaukhov and Karnaukhov, 1993). Zebra mussels also accumulate environmental contaminants; thus, predators feeding on zebra mussels can have notably elevated tissue contaminant loads and reduced reproductive success (see Maclsaac et al., 1996). Zebra mussels have already caused considerable ecological change in the lower Great Lakes and their ecological impacts are likely to extend into other drainage systems as they disperse throughout North America.

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consin Sea Grant Institute, 1994; Ontario Hydro, 1995), these volumes have focused primarily on this species' macrofouling impacts and control, have been largely reports of nonzoologists, have not attempted to analyze, review, or integrate information from multiple sources, and have not been available to the general zoological research community. A more biologically oriented treatise on zebra mussels is Zebra Mussels: Biology, Impacts and Control, edited by Nalepa and Schloesser (1993), which is a collection of papers on aspects of the mussel's biology and macrofouling control. However, much of the research reported in this volume was carried out prior to 1991, before zebra mussels had dispersed little beyond Lake Erie and when most North American investigators had studied them for less than one year. Intensive study of zebra mussel biology in North America has occurred over the four years subsequent to publication of this treatise (Nalepa and Schoesser, 1993). The symposium papers in this issue of American Zoologist update the biology, ecology, and physiology of zebra mussels in North America and provide a comparison with European findings. As zebra mussels become available to an increasing number of North American investigators, this publication will provide a new and comprehensive data base supporting further research on this ecologically and economically important species. ACKNOWLEDGMENTS

This symposium was sponsored by the Division of Invertebrate Zoology and cosponsored by the Division of Comparative Physiology and Biochemistry and the Division of Ecology of the American Society of Zoologists. We are grateful to the symposium participants who devoted considerable time and effort to preparing both their symposium presentations and papers. Many The chapters in this symposium provide of the participants allocated personal or a new synthesis and integration of recent grant funds to partially support their travel biological studies on North American vari- expenses to the symposium. We would also eties of dreissenid mussels. Although full like to acknowledge the experts who relength papers have appeared in the pub- viewed the papers presented in this volume. lished proceedings of several annual con- Cornelia Schlenk, Assistant Director, New ferences on zebra mussels and other exotic York Sea Grant Institute, and Charles R. species {e.g., Tsou and Mussalli, 1993; Wis- O'Neill, of the "Zebra Mussel Information


1993; Fong et al., 1993, 1994; Miller et al., 1994; Ram et al., 1996). Although much information has been accumulated through European studies, the zebra mussel's invasion of North America has triggered a rather extensive American/Canadian research response, sponsored by many agencies, including NSF, NOAA, the U.S. Army Corps of Engineers, the U.S. National Biological Service, power and water utilities, and Canadian and Ontario governmental research and environmental agencies. As North American data have accumulated, it has become apparent that North American and European zebra mussels are not equivalent in all aspects of their biology, ecology and physiology (Mackie and Schloesser, 1996; McMahon, 1996). Lack of congruence between European and North American data may partly result from the rapid dispersal of North American zebra mussels, their elevated growth rates and lack of natural predators, and the fact that the probable source of North American mussels was the Black Sea region of the Ukraine at the extreme southern portion of the zebra mussel's European range. 'Zebra. mussels from this area have been little studied. Instead, the majority of European studies have concentrated on northern European mussels drawn from much colder waters. Thus, North American zebra mussels appear to be more thermally tolerant (McMahon et al., 1993), to have greater growth rates (Griffiths et al., 1991) and to display greater reproductive variation (Garton and Haag, 1993) than reported for mussels from northern Europe. In addition, genetic and morphological studies on North American quagga mussels prove decisively that they are genetically distinct from zebra mussels and that their source populations are in the Dneiper River/Black Sea Region of Ukraine (Spidle et al, 1994; Rosenberg and Ludyanskiy, 1994; Marsden et al., 1996).

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Clearing House of the New York Sea Grant Institute provided invaluable assistance and advice regarding seeking of funding for presentation and publication of the symposium. Presentation and publication of the symposium was supported by grants from New York Sea Grant Institute, The Zebra Mussel Information Clearing House of the New York Sea Grant Institute and NSF Grant No. IBN-9416907. REFERENCES


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