Patch Dynamics and Stability of Some California Kelp Communities

Abstract

This paper considers three concepts of stability as they relate to the dynamics of distinctive patch types of algal canopy guilds in southern and central California kelp communities: (1) persistence of a patch through more than one generation of the dominant species, which was evaluated by using life tables and observations of patch borders; (2) inertia or the resistance of different patches to invasion or disturbance, which was evaluated by artificially enhancing gametophytes by transplanting sporogenic material, by removing canopy, and by evaluating some important disturbance processes; and (3) resilience or recoverability of a patch following a perturbation sufficient to allow invasion of different species, which was studied by defining some of the mechanisms of successful invasion or succession. By working in distinct habitats in southern (Pt. Loma and Santa Catalina Island) and central (Pt. Piedras Blancas) California, we could evaluate different types of physical stresses as they related to these stability concepts. Taller perennial canopy guilds were dominant competitors for light, but were more susceptible to physical wave stress. Dominance hierarchies in the competition for light appeared to be reversed in areas exposed to increasing wave stress. The main causes of mortality at Pt. Loma were entanglement with storm—dislodged Macrocystis plants and, in some areas, sea urchin grazing. Mortality in central California was due to winter storms. In most cases, distinct patches resisted invasion for >10 yr. The mechanisms of resistance involved (1) competition for light and, possibly, nutrients, and (2) limits to spore dispersal. When succession occurred, it was often mediated by many factors, including seasonality of spore production, which coincided with winter storm—related mortalities; mechanisms of kelp dispersal, which were most effective via drifting plants and fragments of fertile material held against the substrate by invertebrates; and survivorship of gametophytes and small sporophytes, which was influenced by local scour and grazing. Appropriate spatial scales, stability, and succession studies in these kelp communities were determined by the size of the disturbed area, which varied from the free space resulting from detachment of single plants to the free space resulting from catastrophies such as overgrazing or unusual storms. Temporal scales were influenced by seasonality of disturbance and algal reproductive condition and aperiodic episodes of cool, nutrient—rich water advected into the patch. There appeared to be conflicting morphological adaptations of the canopy guilds: exploitation of light was enhanced at higher canopy levels, whereas the lower canopy levels were better adapted to tolerate stress from wave surge. The adaptations of the algae appeared to form four distinct groups of tactics: (1) ruderals or plants, such as Nereocystis and Desmarestia, with opportunistic life histories; (2) kelps, such as Macrocystis, adapted to exploitative competition for light and nutrients; (3) kelps (Eisenia, Dictyoneurum) adapted to physical stress such as wave surge; and (4) those algae, such as corallines and Agarum, adapted to heavy grazing. Within any given area, the relative patch stability was determined by biological relationships; between areas, the patch stability patterns were attributable to physical differences.