pH optima for Danish forest species compared with Ellenberg reaction values

Species distribution depends on the physiological and ecological niche where a species can exist and regenerate in resource competition with other species (niche limitation). The realized niche is influenced by local biotic processes that influence species behaviour and the shape of the response curves relative to environmental gradients. Processes on larger scales also influence the species niche through source-sink mechanisms (dispersal limitation) and the species richness of an area (pool limitation). Despite the growing evidence of skewed or irregular species response curves along gradients, many ecologists still assume symmetric, unimodal response curves along gradients in ecological interpretation. Ellenberg's indicator system is probably the most common example. However, the assumption is not ecologically or statistically valid, due to the many different processes affecting the distribution of plant species. Here I present the results of Huisman-Olff-Fresco (HOF) regressions for 209 Danish forest species. HOF modelling is chosen to avoid the classical drawbacks of assuming symmetric, unimodal response patterns. I calculate the optima for all species with unimodal responses to soil pH and compare these with the Ellenberg indicator values for reaction (R), which are often used as a substitute for soil pH measurements. I demonstrate that the assumption of symmetric, unimodal species behaviour is violated in 54% of the cases and that pH optima and R indicator values for species are not always compatible. Ellenberg reaction scale has been used by EWALD (Folia Geobot. 38: 357-366, 2003) as an indicator of which species are calcicole, i.e., whether they can grow and reproduce on calcareous soils. Such affinities of species, however, are related to both local niche properties and processes on large scales and cannot be generalized from a single empirical variable such as pH, nor from Ellenberg semi-ordinal indicator scale. I conclude that while the determination of whether species are calcicole or calcifuge requires more research, it is evident that Denmark contains a fairly balanced number of calciphytic and acidophytic species. This is probably due to the nearly equal areas with acidic and alkaline soils in Denmark, which also contribute to the high species richness of more than 500 vascular plant species in Danish forests.