ML042800052

Zebra Mussels: Biology, Impacts, and Control, Chapter 13

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ZEBRA MUSSELS BIOLOGY, IMPACTS, AND CONTROL Edited by Thomas F. Nalepa Donald W. ScWoesser

CHAPTER 13 Some Aspects of the Zebra Mussel, (Dreissena polymorpha) in the Former European USSR with Morphological Comparisons to Lake Erie NataJiya F. Smimova, G. 1. Biochino, and Germane A. Vinogradov INTRODUCTION The main objectives of this work are to discuss the origin of the zebra mussel, Dreissena po/ymorpha. and to present data on the physiological, morphological, and cytogenetic polymorphism of this species in the former European USSR.

HISTORICAL Dreissena was first found in the lower course of the Ural River in 1769 and later described as a zoologicaJ species in 1771 by the Russian zoologist 0-87371-696-Si93JSO.OO

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218 ZEBRA MUSSELS: BIOLOGY. IMPACTS. AND CONTROL Piter Pallas. Dreissena became of great interest during the 1820swhen it was found at the London docks and then in different places of western Europe.

In Germany it acquired the name of the wandering mussel ("Wundermus-chel") because of its ability to spread rapidly to different areas. Later in the same century. this molluskbegan to block water supply pipes in Paris, Arlee.

Berlin. and many other towns and cities throughout Europe (Zhadin. 1946).

The contemporary family of Dreissenidae is represented by only two genera. Dreissena and Congeria. The most ancient representatives of Dreis-sena belong to the genus Congeria. which appeared in the early Eocene.

Congeria was most widespread and abundant during the epoch of the first and second Pontic Pier. However. in the Pliocene period Congeria almost completely disappeared from Europe and was replaced by the genus Dreis-sena. The largest distribution of Dreissena occurred during the Khvalynsk epoch of the Quarternary period. During this period. Dreissena was found in the Volga River and its tributaries. in northern areas of Eastern Europe. in Western Europe. and in the Aral Sea (Andrusov. 1897).

During the Quarternary glacial epoch. the geographic range of Dreissena declined dramatically. This was probablya resultof coarse material suspended in glacial outwash and the negative impact this material had on the sensitive siphons of this species. While the effects of turbid glacial flows were wide-spread, there were areas in its former range that were not affected and Dreis-sena survived. These areas were the brackish waters of the Caspian and Aral Seas, in the freshwater portions of the Azov and Black Seas. and also in some water bodies of the Balkan peninsula.

PHYSIOLOGICAL, MORPHOLOGICAL,AND CYTOLOGICAL VARIABiliTY Dreissena. as is typical of an organism that is very adaptable and able to occur over a wide range of environmentalconditions, can form populations that are locally distinct. Along the Voiga River, populations of Dreissena have formeddistinct ecotypes or races that differ in their tolerances to various environmental parameters. most notably temperature and salinity. Studies have shown that these different tolerance limits are distinctive at both the organism and cell level. For example, populations from six different sites along the Voiga (Figure I) were subjected to thermal tolerance tests. Indi-viduals from the most southern site (Astrahan) and from a site subject to thermal discharges (Kostroma) were more tolerant of elevated temperatures than individualsfrom twoof the more northern sites (Rybinsk and Kuibyshev)

(Figure 2). In experiments to determine salinity tolerances. individuals from the site nearestto the Caspian Sea (Astrahan) had a lower mortality in relation to increasesin salinity than individualsfrom the site farthest from the Caspian (Rybinsk) (Figure 3). In addition. survival of ciliary epithelial cells under

Smirno\\'O. Biochillo. and Vinogradov 219 I

400km I VolgaRiver Figure 1. Locationof samplingstations along the VolgaRiver;1 = Rybinsk. 2 = Kos-troma.3 = Kuibyshev.4 = Samara,5 = Chapaevsk.6 = Astrahan. (Redrawn from Shkorbatov. G. L. System of Integration of Species as a System (Vilnus.

1986]).

increased salinity conditions (25%0)was much greater in individuals from near the Caspian than from a freshwater site (Kuibyshev) (Figure 4).

An examination of polymorphism in both color and pattern of Dreissena shells from different parts of the fonner European USSR revealed six main varieties or phenotypes (Figure 5). Differences in the frequencies and relative proponions of occurrence of these varieties indicate five main population

220 ZEBRA MUSSELS: BIOLOGY, IMPACTS. AND CONTROL 100 80 6

Figure 2. Relationship between mortality (%) and temperature in Dreissena from different water bodies in the former European USSR. Numbers correspond to the' water bodies as shown in Figure 1. (Redrawn from Shkorbatov, G. L. System of Integration of Species as a System

[Viinus. 1986)).

groups: Aral-Caspian, Ponto-Caspian, Middle-Russian,Baltic. and Northeast.

The maximum number of varieties and the greatest differences in the ratio of these phenes are found in the Aral-Caspian group. which is indicative of the unique position of this group in the system of intraspecificdifferentiation of Dreissena populations (Biochino and Slynko. 1988 and 1990; Biochino.

1990). These five groups are mostly confined to specific and separate geo-graphical regions that can be differentiated by the time period in which Dreissena first colonized that region (Morduhai-Boltovski, 1960). Further.

since these regions coincide with the separation of Eurasian mammal fauna (Starobogatov. 1970), it may be assumed that these Dreissena groups possess the status of distinct geographical races (Mayr, 1974).

Specimens of Dreissena were collected from Lake Erie near Monroe. MI in summer 1990 to compare shell color and pattern of North American in-dividuals to those from the former European USSR. Preliminary analysis indicates that the specimens from Lake Erie are most similar to specimens from the Ponto-Caspianregion; that is. only Lake Erie and the Ponto-Caspian group have individuals with the DD phenotype (Table I). To further assess the degree of similarity between populations from the different regions. the similarity index of Zhivotovsky (1982) was used. This index is calculated as:

r = I (Pmqm)O,S where p is the occurrence frequency (or proportion) of l

60 40

E 20 0

28 30 32 34 36 Temperature (C.)

Smirnova, Biochino. and Vinogradov 221 50 40

~

~ 30

.~~o

E20 10 o

6 2

2 3

4 5

6 7

Salinity

("100) 9 10 8

Figure 3.

Relationship between mortality

(%) and salinity

{%oj in Dreissena from two different water bodies in the former European USSR. The exposure period was 2 weeks.

1 = Rybinsk (freshwater).

2 = Astrahan (near Caspian

Sea, brackish waler). (Redrawn from Antonov, P. I., and Shozbatov.

G. L Species and Its Productivity Wifhin Distribution Area [Moscow.

1983)).

100 o.o 30 60 90 120 150 Time of Exposure (minutes) 180 210 Figure 4.

Relationship between survival (%) and time of exposure (min) for ciliated cells from the gill epithelium of Dreissena in water of 25%0 salinity. The Dreissena cells were taken from individuals from two different water bodies in the former European USSR.

1 =

Rybinsk (fresh water).

2

= Astrahan (near Caspian Sea, brackish water).

(Redrawn from Antonov.

P. I.. and Shozbatov, G. L Species and Its Productivity Within Distribution Area [Moscow.

1983]).

80 l

60 j.

J 40 CI) 20

222 ZEBRA MUSSELS: BIOLOGY, IMPACTS, AND CONTROL DO cc MM AA RR 00 Figure S. The six varieties (phenotypes) of Dreissena found in the former European USSR. Differencesbasedon color and pattem of shells. DO = light, stripeless; AA = arched stripes; CC = mixed, zigzag; RA = radial striped; MM =

spotted; 00

= dark, stripeless.

Table 1. Proportionof DifferentPhenotypesIn Populations of Drelssena from Different Regions in the Former European USSR and from Lake Erie Phenotype Region AA DD 00 RR MM CC Ponto-Caspian 0.094 0.022 0.056 0.008 0.006 0.814 Middle-Russian 0.170 0.000 0.110 0.002 0.010 0.708 Baltic 0.080 0.000 0.015 0.005 0.001 0.899 Lake Erie 0.120 0.030 0.170 0.110 0.190 0.380 Note: The different phenotypes are shown in Figure 5. Specimens from the various regionswere collected from the following water bodies: Ponto-Caspian Region -

lower Volga River, Tsymlyansk and Krenenchug reservoirs, and mouth of the Don river; Middle-Russian Region -

Kuibyshev, Cheboksarsk, Saratov, and Volgograd reservoirs; Baltic Region -

Aybinsk and Ivankovsk reservoirs and Kurshsky Bay.

Smirnova.

Biochino. and Vinogradov 223 Table 2. Index of Similarity Between Populations of Dreissena from Regions in the Former European USSR and Lake Erie Region Ponto-Caspian Middle-Russian 0.976 Region Ponto-Caspian Middle-Russian Baltic Lake Erie Note: Samples collected fromwater bodies as given in Table 1. See text for index derivation.

Baltic 0.980 0.961 Lake Erie 0.849 0.830 0.760 a phene of the first population and q is the occurrence frequency of the same phene in the second population. Values are calculated for each phene. and then the values are added to get the index r. The index ranges from 0 (no common phene) to I (all phenes similar). Based on this index. it is apparent that specimens from the POnlo-Caspian.Middle-Russian. and Baltic regions are more similar to each other than to specimens from Lake Erie (fable 2).

Thus. given the extent of these phenotypic differences. it may be proposed that the introduction of Dreissena into Nonh America occurred with individ-uals from outside these panicular regions of the former European USSR. It must be noted. however. that Lake Erie specimens were not compared to specimens from the Nonheast and Aral-Caspian regions.

In addition to examining the physiological and morphological variability in these populations. it is also useful to examine cytogenetic variability.

Peculiarities in the organization of chromosomes are directly linked to for-mation of reproductive isolation. differences in fecundity and viability of the organisms. formationof geographical races, etc. Investigationsof cells of the gill epithelium and gametes of Dreissena from the former European USSR have shown that the chromosome number varies from 21 to 32 and that the karyotypeof 32 chromosomesis modal(Grishanin. 1990).The modalcomplex included 24 meta, 4 submetacentric. 2 subtelocentric. and 6 acrocentric chro-mosomes. The percentage of modal chromosomes. however, is highly vari-able. For instance. in specimens from cooling waters of the Litovskayapower station. the modal number of chromosomes was found in 87% of the nuclei; but in specimens from the Rybinsk reservoir. the modal number of chro-mosomes was found in only 23% of the nuclei (Barshene, 1990).

YEAR-TO-YEAR FLUCTUATIONS Various long-term studies of Dreissena larvae in reservoirs of the former European USSR have shown natural year-to-year fluctuations in numbers. In the Dnepropetrovskreservoir, numbers declinedevery thirdyear (Dyga. 1966)

224 ZEBRA MUSSELS: BIOLOGY. IMPACTS. AND CONTROL 108 N

Number of Larvae 108 (m~/S)

I

Iga River DiScharge o

1 20 6.H Caspian Sea (em) 0 WaterLevel 1

1975 1

1980 1

1985 1

1990 Figure 6. Year-to-yearlIuctuations in the number of Dreissena in the Ivankovsk and Uglich Reservoir relative to fluctuations in the discharge in the Volga River and water levels of the Caspian Sea. N = Mean total number of larvae in the two reservoirs, a = discharge of the Volga River (r03/s), 6.H = changes in water levels of the Caspian Sea (em).

and in the Kyibyshevskreservoirnumbersdeclinedeveryfourth year (Kir-pichenko, 1964). We examined annual fluctuations in larvae number, water temper.1ture,and water transparency in the Ivankovsk and Uglich Reservoirs over the period of 1973-1988. In general, larvae numbers were greater in years when temperatures were higher than nonnal and watertransparencywas lower. Since these reservoirs are located on the Voiga River, many of their physical and chemical characteristics are influenced by discharge patterns of

Sm;rnova, Biochino, alld V;lIogradm' 225 this river. Therefore, we further examined changes in larval numbers relative to annual changes in water volume of the Volga. Water volume is primarily influencedby changes in the levelof the CaspianSea which, in turn, isrelated to changes in the earth poles (Smirnov, 1969, Sarukhanyan and Smirnov.

1971). The number of larvae in the reservoirs was clearly related to discharge volume of the river and changes in levels of the Caspian Sea (Figure 6).

ACKNOWLEDGMENT We would like to acknowledge the contribution of Olga Zhavoronkova who drew the various zebra mussel phenotypes.

REFERENCES.

Andrusov, N. I. "Extinct and Living Dreissenidae of Eurasia," Tr. SPB. Oba. Es.

restVoisp.,Otd. Geol. Miner. 27:683 (1897).

Antonov. P. I., and G. L. Shkorbatov. "Ecological and Morphological Variabilityof the Volga Populations of Dreissena polymorpha (Pallas):'

in Species and Its Productivity Within Distribution Area (Moscow, 1983), pp. 116-128.

Barshene. Y. V. "Cytogenetic Peculiarities of Dreissenas and Unionids," in Species in Its Area. Biology, Ecology, and Production of Aquatic Inverrebrates (Minsk.,

1990), pp. 126-130.

Biochino, G. I. "Polymorphism and Geographical Variabilityof Dreissena polymor-pha (Pallas)," in Microevolution of Freshwater Orgallisms (Rybinsk, 1990), pp.

143-158.

Biochino, G. I., and Yu. V.Slynko, "Interspecific Differentiationof DreissenaWithin Its Area," Ecology of Populations, Part I (Novosibirsk, 1988), pp. 87-89.

Biochino, G. I., and Yu. V. Slynko. "Population Structure of Dreissena polymorphD (Pallas) Within Its Area," inSpecies inIts Area. Biology, Ecology, and Production of Aquatic Invertebrates (Minsk, 1990), pp. 130-135.

Dyga. A. K. "Biological Foulingon HydrotechnicalStructures in the Dnepropetrovsk Reservoir and Their Control," Cand. Thesis (Dnepropetrovsk, 1966) 19p.

Grishanin, A. K. "Kariotype of the Bivalve Mollusc Dreissena," in Species in Its Area. Biology, Ecology, alld Production of Aquatic Invertebrares (Minsk. 1990).

pp. 121-123.

Kirpichenko, M. Y. "Phenology, Number, Dynamics, and Growth ofDreissena Larva in the Kuibyshevsk Reservoir," in Dreissena Biology and Control (Moscow and Leningrad. 1964), pp. 19-30.

Editors' Note: Citations are presented as provided by the authors. The anicles are published in Russian with titles translated into English.

For further infonnation about a specific

citation, contact the senior author:

Dr. Nataliya

Smimova, Institute Biology of Inland

Waters, Nekouzskiy Raion, Yaroslavslcaya,

Oblast, Borok, Russia.

226 ZEBRA MUSSELS: BIOLOGY. IMPACTS, AND CONTROL Mayr, E. Populations. Species. and Evolution (Moscow, 1974). 46 Op.

Sarukhanyan, E. I.. and N. P. Smimov. Many-Year FluctUlJtionsof the Volga Dis-cJw.rge(Leningrad, 1971) 164p.

Smimov, N. P. "Causes of Long-Period Stream Flow Fluctuations," Sov. Hydrol.

3:308-314 (1969).

Starobogatov, Y. 1. "The Mollusc Fauna and Zoogeographical Division into Districts of the Continental Waters," Nauka (970) 372 p.

Shkorbatov. G. L. "Interspecific Differentiationand Integrityof Speciesas aSystem."

in System of Inregrationof Species (Vilnus, 1986). pp. 118-137.

Zhadin, A. Y. "The Wandering Mussel Dreissena," Priroda 5:29-37 (1946).

Zhivotovsky, L. A. "Indices of Population Variabilityon Polymorphic Characters,"

in Population Phenetics (Moscow, 1982). pp. 38-45.