Succession in inland sand ecosystems: which factors determine the occurrence of the tall grass species Calamagrostis epigejos (L.) Roth and Stipa capillata L.?
In many successional series, grass species play an important role in mid-successional stages. Using calcareous sand ecosystems in the northern upper Rhine valley (Germany) as a model, we analyzed successional trajectories and the factors that determine their direction. A 5- to 7-year study of 23 permanent plots is presented. Polar ordination revealed that succession starts with pioneer communities (dominated by Corynephorus canescens and cryptogams) and can eventually lead to relatively stable Calamagrostis epigejos or Stipa capillata tall grass stands. At some sites, Cynodon dactylon can play a major role, but we focused on the two previously mentioned species. Gradient analyses of their ecological amplitudes by means of principal components and regression analysis were carried out in order to elucidate the factors determining the successional trajectories. Soil analyses of 71 plots established significantly positive linear relations for C. epigejos with respect to total nitrogen, extractable phosphate and potassium, as well as soil moisture indicator values. C. epigejos is not able to spread if the stress factors nutrient deficiency and dryness are combined. Lower tolerance limits are presented. In contrast, S. capillata shows negative relations to phosphate, declining strongly at P contents > 20 mg kg(-1). At a moisture indicator value of around 3.2 (indicating dry sites), S. capillata reaches its optimum. S. capillata is a model for a tall grass species which, even with low nutrient levels and dry soil conditions, can be successful in mid-successional phases; but it is displaced by C. epigejos if soil moisture increases or phosphate/potassium supply is higher. Regarding a general framework, the successional phases are characterized by processes such as replacement of species, facilitation, and inhibition.