Density effects in a colonial monoculture: experimental studies with a marine bryozoan (Membranipora membranacea L.)

Naturally occurring monocultures of plants and animals are not common, despite recent emphasis on the analysis of density effects in artificial plant monocultures. In natural populations, Membranipora membranacea, an encrusting marine bryozoan, usually forms monospecific, nearly even-aged stands on kelp blades. We experimentally manipulated the density of M. membranacea colonies and monitored the responses of individual colonies on settling panels. Colonies undergo a sub-annual cycle of growth, stasis and reproduction, shrinkage, and death. However, crowding by conspecifics accelerates the transition to stasis, triggers early onset of reproduction, and results in increased stage-specific mortality. Unlike many interactions involving colonial invertebrates, overgrowth rarely occurs at boundaries of M. membranacea colonies. Instead, colonies stop growing when they contact conspecifics; therefore more dense assemblages are populated with smaller individual colonies. At the peak in colony size during August, the mean size among colonies grown at high population densities was 300 mm² less than colonies grown at low densities or approximately 62% smaller. Mortality was concentrated in small size classes; at the end of the season colonies gradually shrank to the smallest size classes and then died. We summarized the demography of M. membranacea colonies on low- and high-density panels using size-classified transition matrices and used loglinear analysis to examine the effects of density and time on the transition patterns. As the amount of free space on panels declined, so did the frequency of upward size-class transitions. Our analysis revealed that free space declined more rapidly on panels in the high density treatment and that the transitional probabilities were sensitive to density of conspecifics and seasonal change, but only for some size classes and during some time periods.