Distribution and regulation of the originally tropical cyanobacterium Cylindrospermopsis raciborskii at its northern limits
Cylindrospermopsis raciborskii is a filamentous freshwater cyanobacterium of tropical and subtropical origin that spread out up to northern temperate zone during the last decades. Belonging to the order Nostocales it can differentiate two types of nonvegetative cells: i) heterocysts, in which it can fix molecular nitrogen, and ii) akinetes, which are resting stages, that allow the species to pass-by unfavourable growth conditions in the sediment. In the tropics and subtropics, C. raciborskii is a perennial species, which often occurs in bloom densities, and which is prominent for the synthesis of various toxins including the potent hepatotoxin cylindrospermopsin (CYN) and neurotoxic paralytic shellfish poisoning toxins. In Europe, its northernmost populations were found in North German lakes. Here, it occurs only during the summer months and has not been observed to form mass developments. The highest documented biomass it attained was 23 % of the total phytoplankton biomass. To investigate the spatial occurrence and relative frequency of C. raciborskii, we undertook a systematic survey, sampling 142 water bodies in the lowland region of Northeast Germany from June till September 2004. The cyanobacteria species present were analysed qualitatively and semi-quantitatively. C. raciborskii was a common species. It was present in 39 of the samples, of which 62 % were new findings. Further, we detected two additional thermophilic Nostocales that have not been reported from Germany before: Anabaena bergii and Aphanizomenon aphanizomenoides. These are the most northerly reports of both species so far. A. bergii occurred in 13 and A. aphanizomenoides in 19 of the 142 water bodies sampled. To elucidate the regulation mechanisms of the population dynamics of C. raciborskii and its abiotic boundaries in the newly invaded habitats furthermore long-term data series (1993 - 2005) of two polymictic shallow lakes in the study area were analyzed. Population sizes largely varied between years without any distinct long-term trend. In the annual course, filaments of the species occurred in the pelagial at temperatures above 15 - 17 °C, i.e. akinetes must have been germinated at temperatures below 15 °C. Population growth started at high rates (1.5 - 2.8 d-1) that declined continuously over the season. Growth rates were not significantly correlated with temperature but they were significantly positive correlated with the mean photosynthetic active radiation in the mixed water column (Imix). At the time population starts growing, Imix has exceeded its annual maximum, and growth rate declines with decreasing Imix. The time of akinete germination is therefore of crucial importance for the success of the population: as earlier akinetes germinate the higher the growth rate will be, resulting in a greater population size, a higher number of akinetes produced, and vice versa. Therefore, we conclude that an earlier increase of the water temperature due to global warming is a major cause for the spreading of C. raciborskii into the temperate zone.