Macroalgae in the Baltic Sea -responses to low salinity and nutrient enrichment in Ceramium and Fucus

The brackish Baltic Sea is a marginal environment for both marine and freshwater species. The rate of ecological differentiation is presumably high due to strong selection pressure from a gradient of decreasing salinity that has been present in its current state for only about 3 000 years. Even more recently, increased nutrient loading due to human activities has affected the growth rate of species, with potential effects on their competitive interactions and responses to other regulating factors. I have investigated the potential effects of low salinity and nutrient enrichment on the distributional ranges of two marine macroalgae with a wide distribution in the Baltic Sea, the red alga Ceramium tenuicorne (Kütz.) Wærn and the brown alga Fucus vesiculosus L. A field study in the northern Baltic Sea indicated a strong relationship between the community structure of macroalgae and abiotic factors even on a small, local scale. The abiotic factors are potentially modulated by eutrophication, which may have a strong effect on the depth distribution and abundance of macroalgae. On a regional scale, laboratory experiments suggested that nutrient enrichment is unlikely to affect the distribution of Ceramium and Fucus along the salinity gradient. Growth in Ceramium from the Baltic Proper was enhanced by nitrate and phosphate, but the response did not override growth constraints due to low salinity. Ceramium from the Gulf of Bothnia had an inherently lower growth rate that was not positively affected by nitrate and phosphate increase. In Fucus vesiculosus, reproductive performance was impaired by nitrate and phosphate levels corresponding to ambient levels in eutrophicated areas of the Baltic Sea, when measured by their effect on zygote attachment, germination, and rhizoid development. The wide distribution of Ceramium in the inner Baltic Sea is probably related to local adaptation, rather than a generalized tolerance of different salinity levels. Ecotypic differences were observed when comparing strains from the Baltic Proper (salinity 7 psu) and the Gulf of Bothnia (4 psu). A high rate of vegetative reproduction was evident, although sexual reproduction was occasionally observed in salinity 4. In Fucus vesiculosus, genetic and morphological analyses of sympatric and allopatric populations of the common, vesicular, morphotype and a dwarf morphotype, characteristic for the Gulf of Bothnia, showed that the dwarf morphotype represents a separate evolutionary lineage. Also, vegetative reproduction was observed in Fucus for the first time, as supported by genetic and experimental data. The results show that the biota of the inner Baltic Sea may have unique adaptive and genetic properties, and that it is highly relevant to consider subspecies diversity in Baltic Sea management.