LIGHT-FIELD HETEROGENEITY AMONG TUSSOCK GRASSES - THEORETICAL CONSIDERATIONS OF LIGHT-HARVESTING AND SEEDLING ESTABLISHMENT IN TUSSOCKS AND UNIFORM TILLER DISTRIBUTIONS

Although the tussock growth caespitose graminoids is widespread, the effect of this growth form on light interception and carbon gain of tillers has received little attention. Daily incident photosynthetic photon flux density (PFDinc) and carbon gain in monospecific stands of tussock grasses were compared with those of a hypothetical distribution with the equivalent tiller density per total ground area, but evenly distributed rather than clumped in tussocks. This was computed for two tussock grasses Pseudoroegneria spicata (Pursh) A. Love (bluebunch wheatgrass) and Agropyron desertorum (Fisch, ex Link) Schult. (crested wheatgrass) at different plant densities. Daily PFDinc and net photosynthesis (A) were greater if tillers were distributed uniformly rather than clumped in tussocks, except when the density of tussocks was so great as to approach a uniform canopy. When tussock density per ground area was low much of the difference between tussock and uniform tiller densities in PFDinc and A was due to shading within the tussocks; up to 50-60% of the potential carbon gain was lost in A. desertorum due to shading within tussocks. In a matrix of tussocks, the light field for establishing seedlings was very heterogeneous; potential A ranged from 7 to 96% relative to an isolated seedling. The mean of daily PFDinc and A for seedlings in a tussock stand were nearly identical to the values in corresponding stands of uniform tiller distributions. It is hypothesized that the loss of A resulting from clumping tillers into tussocks is offset by benefits of protecting sequestered belowground resources from invasion by seedlings of competitors.