Establishment of the Baikalian Amphipod Gmelinoides Fasciatus (Amphipoda, Crustacea) in Lake Onega1 N

Establishment of the Baikalian Amphipod Gmelinoides fasciatus (Amphipoda, Crustacea) in Lake Onega1 N. A. Berezina and V. E. Panov Zoological Institute, Russian Academy of Sciences, St. Petersburg, 199034 Russia Received March 12, 2002

1 The number of biological invasions has recently taken. Water temperature in the littoral zone was 10Ð increased drastically all over the world (Carlton, 1991). 17°C in Onega Lake and 18.5Ð20°C in the Svir’ River. This problem is especially important for European Water in Onega Lake had very low mineralization, 33Ð aquatic ecosystems where invasive2 species often bring 40 mg/l (Sabylina, 1999). At most sites studied, the about considerable reconstruction of natural communi- content of the main cations was similar (Na+, 2.1; Ca2+, ties (Invasive Aquatic Species of Europe, 2002). The 7.6Ð8.2; K+, 0.7 mg/l), being somewhat increased near amphipod Gmelinoides fasciatus (Stebbing) of Baikal Petrozavodsk (Na+, 5.1; Ca2+, 12.1; K+, 1.1 mg/l), origin was introduced into some Russian water bodies which appears to be associated with an increased tech- to increase their fish productivity. In most these water nogenic load on the water body and a greater trophicity. bodies, G. fasciatus became a mass species, displacing Pondweed (Potamogeton gramineus) was the dominant from littoral communities other amphipod species, macrophyte in the water basins studied. including the previously dominant Gammarus lacustris Sars (Panov and Berezina, 2002). In the late 1980s, the Abundant data are available in the literature on the Baikalian amphipod was found in Ladoga, the largest distribution of amphipods and their role in zoobenthic European lake, and in 1994, in Beloe Lake (Panov, communities of Onega Lake over the last 100 years 1994; Shcherbina et al., 1997). Since these lakes are (Yarzhinskii, 1870; Segestrole, 1954; Gordeev, 1965; connected to Onega Lake by the Svir’ River and the Alimov et al., 1982; Polyakova, 1999; etc.). According VolgaÐBaltic Canal (Fig. 1), there is the possibility of to these authors, the following amphipods inhabit G. fasciatus becoming established in Onega Lake, with Onega Lake: Gammaracanthus lacustris (at a depth accompanying changes in communities. The dispersion 20Ð28 m), Monoporeia affinis, Pallasea quadrispinosa, speed of G. fasciatus is known to be considerable. For and Gammarus lacustris. Gmelinoides fasciatus was example, in the Volga basin, this amphipod successfully not recorded in Onega Lake before the mid-1990s. Our colonized in 20 years water bodies situated 300Ð400 km investigation in the summer of 2001 revealed this upstream from the introduction site. Tolerance of Baikalian amphipod throughout the west coast of G. fasciatus to various abiotic factors may also facili- Onega Lake. It must have established itself there during tate rapid dispersion (Berezina et al., 2001; Berezina the last several years. and Panov, 2002). High values of abundance and biomass of G. fascia- In August 2001, we analyzed the state of littoral bio- tus populations have been noted (Table 1), which cenoces in the western part of Onega Lake, from means that G. fasciatus naturalized successfully under Petrozavodsk to the Svir’ River estuary (Fig. 1). Six new conditions. Maximal abundance (8300 ind./m2) areas of stony littoral zone of Onega Lake and two areas and biomass (34 g/m2) were noted at Station 5. At all of the Svir’ River were investigated. The sampling was sites studied, the abundance G. fasciatus in pondweed performed on August 16Ð17, 2001 with a special tube growths was two to five times higher than in the open with a sampling area of 1/16 m2 at a depth of 0.2Ð0.5 m. littoral. At Station 2, where water temperature was the Altogether, 30 samples of bottom organisms were lowest (10°C in the middle of August), the biomass of G. fasciatus was much lower (4 g/m2) than at other sites 1 This article was submitted by the author in English. studied, and so was the fecundity (Table 2). A consider- 2 Species that form stable populations in a new environment out- able decrease in quantitative indices of the amphipod side the natural distribution area as a result of introduction (intended or accidental) or self-dispersion are called alien species was noted in the Svir’ River (Table 1), which is proba- (Shine et al., 2000). Alien species are called invasive if they exer- bly due to different hydrodynamic conditions and lack cise a negative impact on structure and functions of natural com- of suitable habitats. On the average, the abundance munities, which may result in a decrease of the role of aboriginal species, disruption of the trophic structure of the community, a G. fasciatus in the Svir’ River was at least two times decreased diversity of ecosystems, etc. (Convention…, 1992). lower than in Onega Lake.

S171

S172 BEREZINA, PANOV

32° 34° 36° 38°

62° 62°

2 1 Onega Lake

3 4 5 7 61° 61° 8 Svir’ River 6 Ladoga Lake

Beloe Lake

32° 34° 36° 38°

Fig. 1. Discovery sites of Gmelinoides fasciatus in Onega Lake (1Ð8, stations).

At the time of investigation, the populations of may be associated with scarcity of food. At other sites, G. fasciatus were represented by two generations, over- the fecundity was, on the average, 10Ð11 eggs per wintered individuals and those of the summer genera- female. At Stations 3 and 5, where G. fasciatus abun- tion. Females constituted the greatest share in popula- dance and biomass were maximal, the greatest share of tions (33Ð55%), 13Ð23% of them bore eggs or newly the young in a population was observed, 25Ð44%, hatched young in marsupial chambers (Fig. 2). Share of whereas at other stations, it was 15Ð16%. reproducing females and their fecundity (on the aver- At all the sites studied, the share of the Baikalian age, 5Ð6 eggs per female) were the lowest in Onega amphipod in bottom communities was considerable Lake at Station 2 and in the Svir’ River (Table 2), which (Table 2), constituting 50Ð83% of the total abundance and 46Ð79% of the total biomass. The main indices 2 were also high for chironomids, oligochaetes, mol- Abundance, ind./m lusks, stone flies, caddis flies, mayflies (Baetis rhodani, 6000 Heptagenia sulfurea), and isopods Asellus aquaticus. IIIIII IV The greatest species richness (more than 30) was noted 4500 in Onega Lake at Station 3, where, alongside with G. fasciatus, the aboriginal amphipod Gammarus 3000 lacustris was noted (50 ind./m2, 0.4 g/m2). In the Svir’ River communities, the share of G. fasciatus was lower (abundance, 37%; biomass, 42%) than in Onega Lake, 1500 a considerable share being constituted by chironomids (abundance, 38%), mollusks (biomass, 33%), and oli- 0 gochaetes (abundance and biomass, 20Ð21%). 1234567 Stations In summary, the Baikalian amphipod Gmelinoides fasciatus is established in Onega Lake. It may have Fig. 2. SexÐage population structure of Gmelinoides fascia- penetrated there in different ways: via the Svir’ River tus at the stations studied in Onega Lake and the Svir’ River (August 2001): I, males; II, females bearing eggs or young; from Ladoga Lake or via the VolgaÐBaltic Canal from III, other females; IV, young. Beloe Lake. Considerable values of abundance, biom-

ENTOMOLOGICAL REVIEW Vol. 83 Suppl. 2 2003

ESTABLISHMENT OF THE BAIKALIAN AMPHIPOD Gmelinoides fasciatus S173 ± Table 1. Quantitative characteristics of the amphipod Gmelinoides fasciatus (x Sx) at different sites in Onega Lake and the Svir’ River in August 2001 Station Coordinates Abundance, ind./m2 Biomass, g/m2 Onega Lake 1. South of Petrozavodsk 61°48′08″ 2680 ± 356 18.97 ± 2.62 34°26′30″ 2. Petrozavodsk, wharf 61°50′08″ 2280 ± 584 4.00 ± 0.06 34°18′06″ 3. Devichii Bay 61°31′37″ 3140 ± 2012 13.80 ± 6.41 30°00′03″ 4. Brusno Island 61°28′05″ 4128 ± 356 14.60 ± 0.87 35°17′45″ 5. Southwestern part 61°15′11″ 5208 ± 3048 27.40 ± 12.91 35°36′03″ 6. Svir’ Estuary 60°58′28″ 2684 ± 268 7.80 ± 0.22 35°29′59″ Svir’ River 61°04′44″ 1432 ± 536 5.37 ± 0.66 7. Voznesenskoe Village 35°03′34″

8. Lodeinoe Pole 60°44′47″ 10 ± 6 0.06 ± 0.02 33°36′57″

Table 2. The main indices of Gmelinoides fasciatus population in Onega Lake and the Svir’ River in August 2001 Onega Lake Svir’ River Indices 12345678 Maximal (average) body length (mm) 7.9 (6.7) 6.5 (5.7) 10.7 (6.0) 10.4 (7.2) 10.1 (7.1) 10.2 (7.0) 7.8 (6.4) – of males 7.4 (6.0) 7.0 (4.7) 9.8 (6.1) 7.9 (5.9) 7.8 (5.8) 8.1 5.9) 7.2 (5.2) – of females Maximal (average) fecundity 16 (11) 8 (6) 16 (10) 16 (11) 18 (10) 16 (10) 8 (5) 6 (eggs/female) Share of reproducing females, % 14 6 13 23152210– Relative abundance in community 50 83.5 56 81.5 85 64 37 50 ass, fecundity of G. fasciatus, and its leading role in the E.V. Borisovskaya, a research assistant of the Institute littoral biocenoses demonstrate that this amphipod has for Biology of Inland Waters, Borok, for analysis of the naturalized successfully in Onega Lake. Further moni- ionic composition of water. The research was finan- toring of the state of littoral communities of Onega cially supported by the Russian Foundation for Basic Lake is necessary to reveal inevitable transformations Research (grant no. 00-15-97825) and the Federal Spe- of their structural and functional organization after the cial Scientific–Technical Program “Investigations and establishment of a new species. Developments in Priority Directions of Science and Technology” (grant no. 43.073.1.1). ACKNOWLEDGMENTS The authors are deeply grateful to N.N. Filatov, REFERENCES Director of the Institute of Water Problems of the 1. Alimov, A.F., Finogenova, N.P., Balushkina, E.V., Bata- North, Karelian Scientific Centre, RAS, Petrozavodsk, lova, F.M., Kaufman, Z.S., and Polyakova, T.N., Produc- for providing the opportunity to take part in the voyage tivity of Benthos, Limnologicheskie issledovaniya na across Onega Lake on board the R/V Ekolog, and to zalive Onezhskogo ozera Bol’shoe Onego (Limnological

ENTOMOLOGICAL REVIEW Vol. 83 Suppl. 2 2003 S174 BEREZINA, PANOV

Investigations in the Bol’shoe Onego Bay of Onega logicheskie problemy (Lake Onega. Ecological Prob- Lake), Leningrad: Zool. Inst. Akad. Nauk SSSR, 1982, lems), Filatov, N.N., Ed., Petrozavodsk: Karel’sk. pp. 170Ð198. Nauch. Tsentr Ross. Akad. Nauk, 1999, pp. 211Ð227. 2. Berezina, N.A., Khlebovich, V.V., Panov, V.E., and 10. Sabylina, A.V., Modern Hydrochemical Regime of the Zaporozhets, N.V., Salinity Resistance in the Amphipod Lake, Onezhskoe ozero. Ekologicheskie problemy (Lake Gmelinoides fasciatus (Stebb.) Introduced into the Gulf Onega. Ecological Problems), Filatov, N.N., Ed., of Finland Basin (Baltic Sea), Dokl. Akad. Nauk, 2001, Petrozavodsk: Karel’sk. Nauch. Tsentr Ross. Akad. vol. 379, no. 3, pp. 414Ð416. Nauk, 1999, pp. 58Ð108. 3. Carlton, J.T., A Journal of Biological Invasions, Biol. Inv., 1991, no. 1, p. 1. 11. Segestrole, S., The Freshwater Amphipods Gammarus pulex and Gammarus lacustris in Denmark and Fennos- 4. Convention on Biological Diversity, 1992, http: candia—a Contribution to the Late and Postglacial //www.biodiv.org. Immigration History of the Aquatic Fauna in Northern 5. Gordeev, O.N., Higher Crustaceans of Karelian Lakes, Europe, Soc. Sci. Fenn. Comm. Biol., 1954, vol. 15, Fauna ozer Karelii. Bespozvonochnye (Fauna of Kare- no. 1. lian Lakes, Invertebrates), MoscowÐLeningrad: Nauka, 1965, pp. 153Ð171. 12. Shcherbina, G.H., Arkhipova, N.R., and Bakanov, A.I., 6. Invasive Aquatic Species of Europe, Leppäkoski, E., Changes in Biological Diversity of Zoobenthos in the Olenin, S., and Gollasch, S., Eds., Dordrecht: Kluwer, Upper Volga and the Rybinsk Reservoir, Problemy bio- 2002. logicheskogo raznoobraziya vodnykh organizmov basseina reki Volgi (Problems of Biological Diversity of 7. Panov, V.E., The Baikalian Endemic Amphipod Gmeli- Aquatic Organisms of the Volga River Basin), noides fasciatus Stebb. in Ladoga Lake, Dokl. Akad. Popchenko, V.I. and Bychek, E.A., Eds., Tol’yatti, 1997, Nauk, 1994, vol. 336, no. 2, pp. 279Ð282. pp. 108Ð114. 8. Panov, V.E. and Berezina, N.A., Invasion History, Biol- ogy and Impacts of the Baikalian Amphipod Gmeli- 13. Shine, C., Williams, N., and Gundling, L., A Guide to noides fasciatus, Invasive Aquatic Species of Europe, Designing Legal and Institutional Frameworks on Alien Leppäkoski, E., Olenin, S., and Gollasch, S., Eds., Dor- Invasive Species, Envir. Pol. Law Paper no. 40, IUCN, drecht: Kluwer, 2002, pp. 96Ð103. 2000. 9. Polyakova, T.N., Bottom Cenoses under Conditions of 14. Yarzhinskii, F.F., To the Fauna of Crustaceans in Lake Anthropogenic Eutrophication, Onezhskoe ozero. Eko- Onega, Tr. SPb. Obshch. Estestv., 1870, no. 1, pp. 61Ð63.

ENTOMOLOGICAL REVIEW Vol. 83 Suppl. 2 2003