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Key Role of the Amphipod Gmelinoides Fasciatus in Reed Beds of Lake Ladoga

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Key Role of the Amphipod Gmelinoides Fasciatus in Reed Beds of Lake Ladoga BOREAL ENVIRONMENT RESEARCH 14: 404–414 © 2009 ISSN 1239-6095 (print) ISSN 1797-2469 (online) Helsinki 30 June 2009 Key role of the amphipod Gmelinoides fasciatus in reed beds of Lake Ladoga Nadezhda A. Berezina1)*, Lyubov V. Zhakova1), Natalia V. Zaporozhets2) and Vadim E. Panov2) 1) Zoological Institute of Russian Academy of Sciences, St-Petersburg 199034, Russia (corresponding author’s e-mail: [email protected]) 2) St-Petersburg State University, St-Petersburg 199034, Russia Received 27 Feb. 2008, accepted 24 June 2008 (Editor in charge of this article: Johanna Mattila) Berezina, N. A., Zhakova, L. V., Zaporozhets, N. V. & Panov, V. E. 2009: Key role of the amphipod Gmelinoides fasciatus in reed beds of Lake Ladoga. Boreal Env. Res. 14: 404–414. The Baikalian amphipod Gmelinoides fasciatus, a successful invader in Eurasia, colonized the coastal zone of Lake Ladoga (northeastern Europe) in late 1990s. In the summers of 2000 and 2005 the density and biomass of benthic communities associated with macro- phyte beds (Phragmites australis) and role of the invader in fish diet were studied. Due to high production of macrophytes (600–800 g carbon m–2 y–1) the benthic biomass was high, averaging 70.6 g m–2 in 2005. Whilst the mean benthic biomass showed a 2-fold increase from 1990 to 2005, the biomass of native benthos 2-fold decreased, from 33.6 ± 6.6 to 16.5 ± 6.5 g m–2. Gmelinoides fasciatus reached 54 ± 13 g m–2 and constituted 77% in the zoo- benthic biomass in 2005 as compared with 6% in 1990. Some crustaceans such as Gamma- rus lacustris and Asellus aquaticus reduced notably their distribution area and decreased in density. G. fasciatus is an important food item in diet of ruffe, perch and young burbot. Introduction Sea basin including the Neva Estuary, large lakes (Lakes Ladoga, Onega and Peipsi) and most Invasive species can play important community of the small lakes in northwestern and central structuring roles in diverse aquatic ecosystems. Russia (Berezina 2007a). In Lake Ladoga, G. They may enter into direct competition with fasciatus was first found in the late 1980s (Panov native species through predation, resource com- 1996). Although it is known to be the most petition and modification of habitats, resulting in abundant species in all invaded European lakes replacement of vulnerable natives (Elton 1958, (Berezina 2007a, 2007b and literature herein), its Leppäkoski et al. 2002). Significant shifts in place in lake food webs is still not clear. native communities have been recorded as a con- The present study focuses on the state of sequence of amphipod invasions (Leppäkoski et macroinvertebrate communities associated with al. 2002, Kelly and Dick 2005, Berezina 2007a, macrophyte beds (mainly Phragmites australis) 2007b). The Baikalian amphipod, Gmelinoides in Lake Ladoga in 2000 and 2005 and an evalu- fasciatus (Stebbing), is one of the most success- ation of the role of G. fasciatus in these com- ful invaders in eastern Europe and Siberia. Since munities and fish diets. The density and biomass the 1970s, it has been established in the Baltic of macroinvertebrates was compared with earlier BOREAL ENV. RES. Vol. 14 • Gmelinoides fasciatus in reed beds of Lake Ladoga 405 records in 1990 (after Kurashov et al. 1996, Panov 1996) in order to find possible shifts in benthic communities caused by the establish- ment of a new species. Materials and methods Study sites Lake Ladoga, the largest in the northwestern part of Russia and Europe (18 135 km2 area, 46.9 m mean depth), has a mesotrophic status but is sub- ject to eutrophication in numerous shallow bays (Filatov and Pozdniakov 2000). The littoral zone, limited by a depth of 8 m, the lower boundary of macrophytes, constitutes 31% of the total lake area (Raspopov et al. 1996). Before 1970s Lake Ladoga had had oligotrophic status, then during Fig. 1. Sampling sites (1–16) in Lake Ladoga. two decades its ecological status deteriorated catastrophically as a result of anthropogenic pol- lution (Lozovik et al. 2000). Several thousands We chose 16 study sites in the shallow stone- of enterprises discharged waste waters into the sandy littoral zone of Lake Ladoga with devel- lake and its drainage area. The pulp, chemical oped macrophytes, mainly P. australis (Fig. 1 and metallurgical mills and agroindustrial com- and Table 1). Macrophyte and macroinvertebrate plexes are principal sources of pollution in the communities were studied during 20–31 July case of Lake Ladoga (Frumin et al. 2000). 2000 and 5–9 August 2005. All sampling sites In 1980–1990s phosphorus loading to this lake (and microhabitats) were the same during both varied from 2300 to 6800 tonnes per year (Kon- sampling periods, as confirmed by a satellite dratyev et al. 1998). During 1970–1980, the total navigation system (GPS). Geographical coordi- phosphorus concentration in water had been 21– nates, daytime temperatures and sampling depths 26 µg l–1 while in the 1990s it did not exceed 17 at the study sites are listed in Table 1. Bottom µg l–1 (Modern state … 1998, Filatov and Pozdn- sediments were of mixed substrates (stones, yakov 2000). By the year 2000, the state of many gravel and sand). The coverage of hard substrates polluted locations improved owing to liquidation reached a maximum of 90% at Site 10 and varied of some large pulp mills (Filatov and Pozdnyakov between 20% and 60% at all others. 2000, Frumin et al. 2000). However, eutrophi- During the study periods, the northern Sites 8, cation process in the lake is still in progress 9 and 11 were not subjected to significant anthro- especially in its coastal zone (Andronnikova and pogenic stress. The other sites were exposed to Raspopov 2007). “Hot spots” are southern and anthropogenic eutrophication and chemical pollu- western coasts of Lake Ladoga including mouths tion in different degree. The sites 1, 2, 15 and 16 in of Rivers Volkhov, Burnaya, Vladimirskij and southern part of the lake were testified as eutrophic Shchuchij Bays (near Priozersk city) and some areas. The Volkhov Bay (Site 3) was influenced by northern locations such as Pitkaranta, Sortavala, wastewater discharges from aluminum plant which Laskela and Lahdenpohja bays (Raspopov et al. is situated on the Volkhov River. These discharges 1996, Lozovik et al. 2000, Luotonen et al. 2004, are sources of nitrogen and phosphates resulting Petrova et al. 2005). The Valaam Archipelago, in eutrophication of the southern part of the lake eastern coast and northern skerry areas of the lake (Naumenko et al. 2000). Also, waste waters from are characterized by the most favorable ecological the Syas pulp mills released waste waters to this state (Raspopov et al. 1996, Petrova et al. 2005). part of the lake up to 1998 (Frumin et al. 2000). 406 Berezina et al. • BOREAL ENV. RES. Vol. 14 Pollution resulted in a nitrogen to phosphorus ratio ing to Katanskaya (1981). The projective area of 13–17 indicating hyper-eutrophic conditions coverage of species in beds was estimated visu- (Frumin et al. 2000). Besides, the Volkhov and ally. The sampling frames with areas of 0.125 m2 Svir Bays (Site 4) take humic waters from wet- (3 replicates) and 0.03 m2 (5 replicates) were lands and bogs. The amounts of humic compounds used for collecting quantitative samples of mac- contribute 65%–92% to the total organic carbon rophytes. To measure height and biomass of in waters of the Volkhov River (Korkishko et al. P. australis, 7–30 shoots of flowering plants 2000). Humic compounds can be an important (depending on the heterogeneity of the bed) were source of nutrients and microelements facilitat- randomly sampled. Prior to weighing, macro- ing further eutrophication of the lake (Korkishko phytes were rinsed in tap water and dried at et al. 2000). The Shchuchij Bay (Site 12) and 60 °C for 96 hours. Annual primary macrophyte tens of kilometers of lake area to the south from production (PP, g carbon (C) m–2) was estimated it were heavily polluted up to the 1980s because as a function of the maximal elevated biomass of –2 of intensive waste water discharges from a pulp plants (Bmax, g C m ): PP = 1.2Bmax (Raspopov mill (Slepukhina et al. 2000). During the study 1985). years, the Shchuchij Bay retained properties of an Quantitative samples of macroinvertebrates eutrophic area with intensive macrophyte beds. were obtained with a 0.03-m2 diameter cylindri- Moreover, high contents of technogenic materials cal metal frame (0.76 m height), with three rep- (to 70%) were recorded in sediments of the bay. licates taken at each site. The frame was forced Waters of the Vladimirovskij Bay (Site 14) were into the bottom and all the hard substrates and influenced by moderate oil pollution in both 2000 plants in the frame were transferred to a plastic and 2005. Earlier it had also been characterized container filled with fresh water. Animals were as eutrophied and polluted area (Raspopov et al. washed off the hard substrates or scraped off with 1996). a knife (for attached animals). A 3–7 cm layer of soft sediments was collected with a hand-held net (0.25 mm mesh size) for three minutes. All Sampling and laboratory procedures parts of the sample were placed in a sieve (0.25 mm), rinsed with water, transferred to a plas- Macrophyte communities were described accord- tic zip-bag and preserved in 4% formaldehyde. Table 1. Coordinates, sampling depths and daytime temperature of water at study sites. Site Lat. N Long. E Water depth (m) Temperature (°C) 2000 2005 2000 2005 01 60°01´04´´ 31°32´39´´ 0.65 0.7 20 20 02 60°13´17´´ 31°55´36´´ 0.6 0.6 21 20 03 60°07´52´´ 32°19´26´´ 0.65 0.75 21 19.5 04 60°31´26´´ 32°41´13´´ 0.4 0.7 21 20 05 60°58´45´´ 32°36´26´´ 0.6 0.7 19.5 20 06 61°20´30´´ 31°39´51´´ 0.45 0.5 19 20 07 61°24´03´´ 31°30´31´´ 0.6 0.6 21 19 08 61°37´16´´ 31°10´35´´ 0.6 0.5 20 19 09 61°38´07´´ 31°11´26´´ 0.5 0.7 21 19.5 10 61°21´35´´ 30°53´18´´ 0.2 0.5 18 18 11 61°31´07´´ 30°33´27´´ 0.5 0.6 17 17.5 12 61°04´59´´ 30°05´22´´ 0.3 0.6 17 18 13 60°50´10´´ 30°28´06´´ 0.5 0.7 17 19 14 60°37´05´´ 30°31´59´´ 0.3 0.6 17 18 15 60°22´46´´ 30°52´48´´ 0.5 0.7 18 17 16 60°06´39´´ 31°05´29´´ 0.4 0.7 17.5 19 BOREAL ENV.
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