Interactions between neighboring algae and snail grazing in structuring microdistribution patterns of periphyton

The micro-distribution of periphyton (filamentous algae) on homogeneous substrates was examined in experimental tanks with and without the pressure of grazing snails. The growth of periphyton attached to artificial substrate was estimated at a small spatial scale (9.3 mm × 9.3 mm cells) by varying the number of grazers (0, 5, or 10 snails per tank), using image processing analysis without removing the periphyton. The results suggest that periphyton growth within a cell was negatively affected by the biomass of periphyton in the cell but was positively affected by the biomass of periphyton in neighboring cells. A semivariogram analysis indicated that spatial heterogeneity increased with increasing grazing pressure. The size of patches was not clearly related to the number of snails, but there was a tendency for relative patch size to increase with snail density. Computer simulations were also conducted to examine factors affecting the degree of spatial heterogeneity. The simulation studies indicated that snails should graze a site that was previously grazed in order to produce the observed spatial heterogeneity of periphyton. The results also indicated that the positive effects of neighboring periphyton on the growth of algae might create patches. The interactions among neighboring algae and snail grazing might be an important factor creating the spatial heterogeneity of periphyton even on homogeneous substrates.