Competition and coexistence of rhizomatous perennial plants along a nutrient gradient

I studied competition and coexistence of three tall clonal perennial plant species, Calamagrostis epigejos (L.) Roth, Solidago canadensis L., and Tanacetum vulgare L. along a gradient of soil productivity over five years. A replacement series field experiment was conducted with high, moderate and low fertility levels in 1mx1m plots. There were significant effects of soil type on ramet density (P < 0.001), mean height (P < 0.01), and total biomass (P < 0.01). Ramet density, mean height, and total biomass increased with increasing soil fertility. There were also significant effects of mixture on ramet density (P < 0.01), but not on mean height and total biomass for all species. Significant neighbor effects on ramet density and total biomass (P < 0.01) were found for Solidago, showing that it is important whether Tanacetum or Calamagrostis is its neighbor within mixtures. During the five years there was only one case of competitive exclusion: Calamagrostis excluded Solidago on the most fertile substrate in the fifth growing season. In most cases species coexisted over the five years. Each of the three species was able to dominate in at least one combination of substrate type and mixture. The experiment showed that asymmetric competition for light on substrates of high fertility, symmetric competition for nutrients on nutrient-poor soil and positive interactions especially on substrates of intermediate fertility played a role. A founder effect was evident in aggregated mixtures of Calamagrostis and Solidago on the nutrient-rich substrate. A conceptual model of the relative importance of root competition for soil nutrients, shoot competition for light, and positive interactions along the fertility gradient is presented. The model emphasizes that positive interactions play an important role over a broad range of the productivity scale with a peak at intermediate levels of fertility. On the substrate of high productivity shoot competition for light is more important than positive interactions and root competition for soil nutrients as well. The competitive superiority of Calamagrostis on the most productive substrate was evident only in the long run. Rare events like extreme summer drought or selective herbivore pressure caused a switch in dominance in mixtures with Solidago, respectively Tanacetum. The guerrilla growth strategy of Calamagrostis and interference competition through a dense cover of aboveground biomass and litter could further cause competitive exclusion.