Below-ground spatial pattern of rhizomes in a grassland community and its relevance to above-ground spatial pattern

Studies of spatial patterns in grassland plant communities have focused on above-ground patterns, ignoring the fact that in clonal plant communities, such as those found in grasslands, above-ground spatial structure must reflect below-ground horizontal growth. The present study examines, at both a fine and a coarse spatial scale, relationships between rhizome and ramet distribution. At the coarse scale, the dominance of species differed between above- and below-ground; some species dominated only above- or below-ground, and others dominated in both layers. At the fine scale, a single species' ramet aggregation above- ground significantly differed from its rhizome aggregation below-ground, for many species. Even for a given species, quantitative relationships between above- and below-ground dominance varied among localities. The variation in spatial pattern among species can be explained by attributes of clonal growth form, including rhizome size, rhizome origin and pattern of above- ground ramet aggregation. Using these parameters of clonal growth, four major space occupation patterns were described for mountain grassland species. For species with a high abundance of evenly distributed rhizomes, ramets either i) reflect rhizome distribution, or ii) do not fully reflect rhizome distribution, but rather are spatially aggregated, and rhizomes are typically developed below-ground. For species with a low abundance of rhizomes, ramets either iii) reflect rhizome distribution and rhizomes are typically of above- ground origin, or iv) do not reflect rhizome distribution and are aggregated only at the growing tips of rhizomes. Spatial correlation above- ground among species was the same as below-ground for some pairs of species but was significantly different for other pairs.