We studied relationships between nutrient loading, local abiotic variables, richness and biomasses of macrophytes, and associated invertebrate species in the north-eastern Baltic Sea. The study showed that nutrient load strongly correlated to the richness and biomass of macroalgal and invertebrate species and functions and often interacted with local abiotic variables such as salinity and depth. Generally elevated nutrient loads increased the species richness of macrophytes and benthic invertebrates and the biomass of annual macroalgae and of the majority of invertebrate feeding groups, but reduced the biomass of perennial macroalgae. The study also showed that the effect of nutrient loading was scale-specific, i.e. different responses to nutrients were observed at water-body and gulf scales. The biomass of the majority of species and functions was a function of the gulf-scale nutrient loading whereas the variability of annual algae and chironomidae reflected changes in the water-body level nutrient loading.
Bendtsen, J., Gustafsson, K. E., Söderkvist, J., and Hansen, J. L. S. 2009. Ventilation of bottom water in the North Sea–Baltic Sea transition zone. Journal of Marine Systems, 75, 138–149.
http://dx.doi.org/10.1016/j.jmarsys.2008.08.006
Bergström, L. and Kautsky, L. 2006. Local adaptation of Ceramium tenuicorne (Ceramiales, Rhodophyta) within the Baltic Sea. Journal of Phycology, 42, 36–42.
http://dx.doi.org/10.1111/j.1529-8817.2006.00173.x
Bokn, T. L., Duarte, C. M., Pedersen, M. F., Marbà, N., Moy, F. E., Barrón, C., and Bjerkeng, B. 2003. The response of experimental rocky shore communities to nutrient additions. Ecosystems, 6, 577–594.
http://dx.doi.org/10.1007/s10021-002-0108-6
Bonsdorff, E. and Pearson, T. H. 1999. Variation in the sublittoral macrozoobenthos of the Baltic Sea along environmental gradients: a functional-group approach. Australian Journal of Ecology, 24, 312–326.
http://dx.doi.org/10.1046/j.1442-9993.1999.00986.x
Bonsdorff, E., Rönnberg, C., and Aarnio, K. 2002. Some ecological properties in relation to eutrophication in the Baltic Sea. Hydrobiologia, 475/476, 371–377.
http://dx.doi.org/10.1023/A:1020395526898
Clarke, K. R. 1993. Non-parametric multivariate analyses of changes in community structure. Australian Journal of Ecology, 18, 117–143.
http://dx.doi.org/10.1111/j.1442-9993.1993.tb00438.x
Clarke, K. R. and Gorley, R. N. 2006. Primer v6. User Manual ⁄ Tutorial. Primer-E, Plymouth.
Clarke, K. R. and Green, R. H. 1988. Statistical design and analysis for a biological effects study. Marine Ecology Progress Series, 46, 213–226.
http://dx.doi.org/10.3354/meps046213
Clarke, K. R., Somerfield, P. J., and Chapman, M. G. 2006. On resemblance measures for ecological studies, including taxonomic dissimilarities and a zero-adjusted Bray–Curtis coefficient for denuded assemblages. Journal of Experimental Marine Biology and Ecology, 330, 55–80.
http://dx.doi.org/10.1016/j.jembe.2005.12.017
Duarte, C. D. 1995. Submerged aquatic vegetation in relation to different nutrient regimes. Ophelia, 41, 87–112.
Elith, J., Leathwick, J. R., and Hastie, T. 2008. A working guide to boosted regression trees. Journal of Animal Ecology, 77, 802–813.
http://dx.doi.org/10.1111/j.1365-2656.2008.01390.x
Elmgren, R. 1989. Man’s impact on the ecosystem of the Baltic Sea: energy flows today and at the turn of the century. Ambio, 18, 326–332.
Elmgren, R. 2001. Understanding human impact on the Baltic ecosystem: changing views in recent decades. Ambio, 30, 222–231.
http://dx.doi.org/10.1639/0044-7447(2001)030[0222:UHIOTB]2.0.CO;2
Eriksson, B. K. and Bergström, L. 2005. Local distribution patterns of macroalgae in relation to environmental variables in the northern Baltic Proper. Estuarine, Coastal and Shelf Science, 62, 109–117.
http://dx.doi.org/10.1016/j.ecss.2004.08.009
European Union. 2000. Directive of the European Parliament and of the Council Establishing a Framework for Community Action in the Field of Water Policy. Legislative Acts and other instruments. ENV221 CODEC 513.
Field, C. B., Behrenfeld, M. J., Randerson, J. T., and Falkowski, P. 1998. Primary production of the biosphere: integrating terrestrial and oceanic components. Science, 281, 237–240.
http://dx.doi.org/10.1126/science.281.5374.237
Gee, J. M. and Warwick, R. M. 1994. Metazoan community structure on relation to the fractal dimensions of marine algae. Marine Ecology Progress Series, 103, 141–150.
http://dx.doi.org/10.3354/meps103141
Geertz-Hansen, O., Sand-Jensen, K., Hansen, D. F., and Christiansen, A. 1993. Growth and grazing control of abundance of the marine macroalga, Ulva lactuca L. in a eutrophic Danish estuary. Aquatic Botany, 46, 101–109.
http://dx.doi.org/10.1016/0304-3770(93)90039-Y
Grall, J. and Chauvaud, L. 2002. Marine eutrophication and benthos: the need for new approaches and concepts. Global Change Biology, 8, 813–830.
http://dx.doi.org/10.1046/j.1365-2486.2002.00519.x
Granéli, E. and Sundbäck, K. 1985. The response of planktonic and microbenthic algal assemblages to nutrient enrichment in shallow coastal waters, southwest Sweden. Journal of Experimental Marine Biology and Ecology, 85, 253–268.
http://dx.doi.org/10.1016/0022-0981(85)90161-3
Gray, J. S., Wu, R. S., and Or, Y. Y. 2002. Effects of hypoxia and organic enrichment on the coastal marine environment. Marine Ecology Progress Series, 238, 249–279.
http://dx.doi.org/10.3354/meps238249
Haahtela, I. 1984. A hypothesis of the decline of the Bladder Wrack (Fucus vesiculosus L.) in SW Finland in 1975–1981. Limnologica, 15, 345–350.
Hällfors, G., Kangas, P., and Niemi, A. 1984. Recent changes in the phytal at the south coast of Finland. Ophelia, 3, 51–59.
Havens, K. E., Hauxwell, J., Tyler, A. C., Thomas, S., McGlathery, K. J., Cebrian, J., et al. 2001. Complex interactions between autotrophs in shallow marine and freshwater ecosystems: implications for community responses to nutrient stress. Environmental Pollution, 113, 95–107.
http://dx.doi.org/10.1016/S0269-7491(00)00154-8
HELCOM. 2009. Manual for Marine Monitoring in the COMBINE Programme of HELCOM. Helsinki. http://helcom.fi/action-areas/monitoring-and-assessment/manuals-and-guidelines/ combine-manual (accessed 30.07.2014).
Howarth, R. W. 1988. Nutrient limitation of net primary production in marine ecosystems. Annual Review of Ecology, Evolution, and Systematics, 19, 89–110.
http://dx.doi.org/10.1146/annurev.es.19.110188.000513
Josefson, A. B. and Hansen, J. L. S. 2004. Species richness of benthic macrofauna in Danish estuaries and coastal areas. Global Ecology and Biogeography, 13, 273–288.
http://dx.doi.org/10.1111/j.1466-822X.2004.00091.x
Karez, R., Engelbert, S., Kraufvelin, P., Pedersen, M. F., and Sommer, U. 2004. Biomass response and changes in composition of ephemeral macroalgal assemblages along an experimental gradient of nutrient enrichment. Aquatic Botany, 78, 103–117.
http://dx.doi.org/10.1016/j.aquabot.2003.09.008
Kautsky, H. 1988. Factors structuring phytobenthic communities in the Baltic Sea. Doctoral thesis. Department of Zoology, University of Stockholm.
Kautsky, H. and van der Maarel, E. 1990. Multivariate approaches to the variation in phytobenthic communities and environmental vectors in the Baltic Sea. Marine Ecology Progress Series, 60, 169–184.
http://dx.doi.org/10.3354/meps060169
Kautsky, H., Kautsky, U., and Nellbring, S. 1988. Distribution of flora and fauna in an area receiving pulp mill effluents in the Baltic Sea. Ophelia, 28, 139–155.
http://dx.doi.org/10.1080/00785326.1988.10430808
Kautsky, H., Martin, G., Mäkinen, A., Borgiel, M., Vahteri, P., and Rissanen, J. 1999. Structure of phytobenthic and associated animal communities in the Gulf of Riga. Hydrobiologia, 393, 191–200.
http://dx.doi.org/10.1023/A:1003510105274
Kotta, J. and Ólafsson, E. 2003. Competition for food between the introduced exotic polychaete Marenzelleria viridis and the resident native amphipod Monoporeia affinis in the Baltic Sea. Journal of Sea Research, 342, 27–35.
http://dx.doi.org/10.1016/S1385-1101(03)00041-8
Kotta, J. and Orav, H. 2001. Role of benthic macroalgae in regulating macrozoobenthic assemblages in the Väinameri (north-eastern Baltic Sea). Annales Zoologici Fennici, 38, 163–171.
Kotta, J. and Witman, J. 2009. Regional-scale patterns. In Marine Hard Bottom Communities (Wahl, M., ed.), pp. 89–99. Ecological Studies, 206. Springer-Verlag.
Kotta, J., Paalme, T., Martin, G., and Mäkinen, A. 2000. Major changes in macroalgae community composition affect the food and habitat preference of Idotea baltica. International Review of Hydrobiology, 85, 697–705.
http://dx.doi.org/10.1002/1522-2632(200011)85:5/6<697::AID-IROH697>3.0.CO;2-0
Kotta, J., Torn, K., Martin, G., Orav-Kotta, H., and Paalme, T. 2004. Seasonal variation of invertebrate grazing on Chara connivens and C. tomentosa in Kõiguste Bay, NE Baltic Sea. Helgoland Marine Research, 58, 71–76.
http://dx.doi.org/10.1007/s10152-003-0170-2
Kotta, J., Orav-Kotta, H., Paalme, T., Kotta, I., and Kukk, H. 2006. Seasonal changes in situ grazing of the mesoherbivores Idotea baltica and Gammarus oceanicus on the brown algae Fucus vesiculosus and Pylaiella littoralis in the central Gulf of Finland, Baltic Sea. Hydrobiologia, 554, 117–125.
http://dx.doi.org/10.1007/s10750-005-1011-x
Kotta, J., Lauringson, V., and Kotta, I. 2007. Response of zoobenthic communities to changing eutrophication in the northern Baltic Sea. Hydrobiologia, 580, 97–108.
http://dx.doi.org/10.1007/s10750-006-0462-z
Kotta, J., Lauringson, V., Martin, G., Simm, M., Kotta, I., Herkül, K., and Ojaveer, H. 2008. Gulf of Riga and Pärnu Bay. In Ecology of Baltic Coastal Waters (Schiewer, U., ed.), pp. 217–243. Ecological Studies, 197. Springer-Verlag.
Kotta, J., Aps, R., and Orav-Kotta, H. 2009. Bayesian inference for predicting ecological water quality under different climate change scenarios. WIT Transactions on Ecology and the Environment. WIT Press, 127, 173–184.
Kraufvelin, P., Lindholm, A., Pedersen, M. F., Kirkerud, L. A., and Bonsdorff, E. 2010. Biomass, diversity and production of rocky shore macroalgae at two nutrient enrichment and wave action levels. Marine Biology, 157, 29–47.
http://dx.doi.org/10.1007/s00227-009-1293-z
Krause-Jensen, D., Carstensen, J., and Dahl, K. 2007a. Total and opportunistic algal cover in relation to environmental variables. Marine Pollution Bulletin, 55, 114–125.
http://dx.doi.org/10.1016/j.marpolbul.2006.08.019
Krause-Jensen, D., Middelboe, A., Carstensen, J., and Dahl, K. 2007b. Spatial patterns of macroalgal abundance in relation to eutrophication. Marine Biology, 152, 25–36.
http://dx.doi.org/10.1007/s00227-007-0676-2
Lauringson, V. and Kotta, J. 2006. Influence of the thin drift algal mats on the distribution of macrozoobenthos in Kõiguste Bay, NE Baltic Sea. Hydrobiologia, 554, 97–105.
http://dx.doi.org/10.1007/s10750-005-1009-4
Lauringson, V., Mälton, E., Kotta, J., Kangur, K., Orav-Kotta, H., and Kotta, I. 2007. Environmental factors influencing the biodeposition of the suspension feeding bivalve Dreissena polymorpha (Pallas): comparison of brackish and fresh water populations in the Northern Baltic Sea and Lake Peipsi. Estuarine, Coastal and Shelf Science, 75, 459–467.
http://dx.doi.org/10.1016/j.ecss.2007.05.037
Mäkinen, A., Kääriä, J., and Rajasilta, M. 1988. Factors controlling the occurrence of Furcellaria lumbricalis (Huds.) Lamour. and Phyllophora truncata (Pallas) Zinova in the upper littoral of the Archipelago of the SW Finland. Kieler Meeresforschung, 6, 140–146.
McGahee, C. F. and Davis, G. J. 1971. Photosynthesis and respiration in Myriophyllum spicatum L. as related to salinity. Limnology and Oceanography, 16, 826–829.
http://dx.doi.org/10.4319/lo.1971.16.5.0826
Morand, P. and Briand, X. 1996. Excessive growth of macroalgae: a symptom of environmental disturbance. Botanica Marina, 39, 491–516.
http://dx.doi.org/10.1515/botm.1996.39.1-6.491
Orav-Kotta, H. and Kotta, J. 2004. Food and habitat choice of the isopod Idotea baltica in the northeastern Baltic Sea. Hydrobiologia, 514, 79–85.
http://dx.doi.org/10.1023/B:hydr.0000018208.72394.09
Orav-Kotta, H., Kotta, J., Herkül, K., Kotta, I., and Paalme, T. 2009. Seasonal variability in the grazing potential of the invasive amphipod Gammarus tigrinus and the native amphipod Gammarus salinus in the northern Baltic Sea. Biological Invasions, 11, 597–608.
http://dx.doi.org/10.1007/s10530-008-9274-6
Paalme, T., Kukk, H., Kotta, J., and Orav, H. 2002. “In vitro” and “in situ” decomposition of nuisance macroalgae Cladophora glomerata and Pilayella littoralis. Hydrobiologia, 475/476, 469–476.
http://dx.doi.org/10.1023/A:1020364114603
Pavia, H., Carr, H., and Aberg, P. 1999. Habitat and feeding preferences of crustacean mesoherbivores inhabiting the brown seaweed Ascophyllum nodosum (L.) Le Jol. and its epiphytic macroalgae. Journal of Experimental Marine Biology and Ecology, 236, 15–32.
http://dx.doi.org/10.1016/S0022-0981(98)00191-9
Pedersen, M. F. 1995. Nitrogen limitation of photosynthesis and growth: comparison across aquatic plant communities in a Danish estuary (Roskilde fjord). Ophelia, 41, 261–272.
Pedersen, M. F. and Borum, J. 1996. Nutrient control of algal growth in estuarine waters. Nutrient limitation and the importance of nitrogen requirements and nitrogen storage among phytoplankton and species of macroalgae. Marine Ecology Progress Series, 142, 261–272.
http://dx.doi.org/10.3354/meps142261
Pihl, L., Svenson, A., Moksnes, P.-O., and Wennhage, H. 1999. Distribution of green algal mats throughout shallow soft bottoms of the Swedish Skagerrak archipelago in relation to nutrient sources and wave exposure. Journal of Sea Research, 41, 281–294.
http://dx.doi.org/10.1016/S1385-1101(99)00004-0
Posey, M. H., Alphin, T. D., Cahoon, L., Lindquist, D., and Becker, M. E. 1999. Interactive effects of nutrient additions and predation on infaunal communities. Estuaries, 22, 785–792.
http://dx.doi.org/10.2307/1353111
Puttman, R. J. 1986. Grazing in Temperate Ecosystems: Large Herbivores and the Ecology of the New Forest. Croom Helm, London.
http://dx.doi.org/10.1007/978-94-011-6081-0
R Core Team. 2013. R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna. http://www.R-project.org/ (accessed 15.12.2013).
Rönnberg, C. 2001. Effects and Consequences of Eutrophication in the Baltic Sea. Licentiate thesis, Department of Biology, Environmental and Marine Biology, Abo Akademi University, Finland.
Sand-Jensen, K. and Borum, J. 1991. Interactions among phytoplankton, periphyton and macrophytes in temperate freshwaters and estuaries. Aquatic Botany, 47, 137–175.
http://dx.doi.org/10.1016/0304-3770(91)90042-4
Valiela, I., McClelland, J., Hauxwell, J., Behr, P. J., Hersh, D., and Foreman, K. 1997. Macroalgal blooms in shallow estuaries: controls and ecophysiological and ecosystem consequences. Limnology and Oceanography, 42, 1105–1118.
http://dx.doi.org/10.4319/lo.1997.42.5_part_2.1105
Wallentinus, I. 1984a. Comparisons of nutrient uptake rates for Baltic macroalgae with different thallus morphologies. Marine Biology, 80, 215–225.
http://dx.doi.org/10.1007/BF02180189
http://dx.doi.org/10.1007/BF00027703r