Diversity of wetland vegetation in the Bulgarian high mountains, main gradients and context-dependence of the pH role

We fill a gap in understanding wetland vegetation diversity and relationship with environmental determinants in Bulgarian high mountains. A total of 615 phytosociological samples were taken from springs, mires, wet meadows and tall-forb habitats throughout Bulgaria, of which 234 relevés are from mire and spring vegetation above timberline. The vegetation was classified by TWINSPAN and the resulting vegetation types were reproduced by the formal definitions using the combination of Cocktail species groups based on phi-coefficient of joint co-occurrence of the species. Nine vegetation types of springs and fens have been clearly delimited above the timberline. All vegetation types include Balkan endemic species, the representation of which varies. Fens generally harbour more Balkan endemics than do springs, with the exception of species-poor high-altitude Drepanocladetum exannulati. The gradient structure of the vegetation was revealed by DCA and by CCA with forward selection of environmental factors. The major determinants of vegetation variation strongly differ above and below the timberline and likewise between springs and fens. The base-richness gradient controls the floristic variation of Bulgarian submontane fens, whereas the complete data set including both submontane and subalpine fens is governed by the altitude gradient from lowland and basin fens to subalpine fens rich in Balkan endemics. When focusing on sites above the timberline only, the first DCA axis separates fens from springs without organic matter. The major species turnover in springs follows the variation in water pH and mineral content in water, whereas fen vegetation variation is primarily controlled by succession gradient of peat accumulation. Altitude remains an important factor in all cases. Weak correlation between water pH and conductivity was found. This correlation was even statistically insignificant in fens above the timberline. Water pH is not influenced by mineral richness in Bulgarian high mountains, since it is buffered by decomposition of organic matter in fens. In springs, pH reaches maximum values due to strong aeration caused by water flow. The plant species richness decreases significantly with increasing altitude. The increase of species richness towards circumneutral pH, often found in mires, was not confirmed in Bulgarian high mountains. The correlation between species richness and pH was significant only when arctic-alpine species and allied European high-mountain species were considered separately. The richness of boreal species was independent on pH. Some of them had their optima shifted to more acidic fens as compared to regions below the timberline. Our results suggest that subalpine spring and fen vegetation should be analysed separately with respect to vegetation-environment correlations. Separate analysis of fens below and above timberline is quite appropriate.