Small-scale variability as a mechanism for large-scale stability in mountain grasslands

Abstract. Data from a 7-yr permanent plot study of grassland dynamics were used to address the relationship between processes at two levels of resolution (3.3 cm x 3.3 cm, 50 cm x 50 cm). Grasslands mown and manured in the traditional way in the Krkonose Mts. (Riesengebirge) were used as a model system. Spatial dynamics at the finest scale were very high, as demonstrated by turnover of individual species in 3.3 cm x 3.3 cm subplots and year-to-year transition matrices of the same subplots. The direction of these dynamics was not correlated with grassland treatment, although there was some correlation within years. An extrapolation of such year-to-year dynamics to larger time scales would result in big large-scale changes on the community level, and large shifts in species composition of the whole sward. However, dynamics at larger spatial or temporal scales were generally small. Some directional change occurred in manured plots, whereas little change occurred in unmanured plots. Large-scale dynamics were not correlated with small-scale dynamics in plots without manuring, but some correlation was detectable in manured plots. There are probably several processes that drive small-scale dynamics, such as non-linear interactions and environmental fluctuations. We argue that within certain limits these forces act on species composition so as to make small-scale dynamics non-directional. This results in both large-scale species diversity and apparent large-scale stability of these grasslands. However, if these forces are beyond these limits, the small-scale dynamics may become directional, resulting in rapid changes at larger spatial scales.