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The importance that frond crowding represents for the survival of fronds of the clonal intertidal alga Mazzaella cornucopiae (Postels et Ruprecht) Hommersand (Rhodophyta, Gigartinaceae) was investigated in Barkley Sound, British Columbia, Canada. Frond density is high for this species, up to 20 fronds·cm Ϫ2 in the most crowded stands. Frond crowding imposes a cost in the form of reduced net photosynthetic rates when fronds are fully hydrated as a result of reduced irradiance compared with experimental (not found naturally) low-density stands. However, the interaction between desiccation and irradiance alters this relationship between net photosynthetic rates and frond density. During a typical daytime low tide in spring, irradiance is 10-30 mol·m Ϫ2 ·s Ϫ1 below the canopy of fronds, and frond desiccation (relative to total water content) can reach 43% at the end of the low tide. In contrast to natural stands, fronds from experimentally thinned stands are subjected to irradiances up to 2000 mol·m Ϫ2 ·s Ϫ1 because of the spatial separation among fronds and can desiccate up to 81% at the end of the same low tide. Laboratory experiments showed that negative net photosynthetic rates occur between 40% and 80% desiccation at an irradiance of 515 mol·m Ϫ2 ·s Ϫ1 , and the literature suggests that strong bleaching could occur as a result. At 20 mol·m Ϫ2 ·s Ϫ1 of irradiance and desiccation levels up to 40%, simulating understory conditions of natural stands, net photosynthetic rates are never negative. Experimental thinning of stands of M. cornucopiae done during spring effectively resulted in a stronger extent of frond bleaching compared with natural stands. Therefore, the cost of reduced net photosynthetic rates at high frond densities when fronds are fully hydrated is counterbalanced by the protective effects of frond crowding against extensive bleaching, essential for survival at the intertidal zone. Future research will have to demonstrate the possible relationship between the frequency and duration of negative net photosynthetic rates and the extent of frond bleaching.In the intertidal zone, the regular alternation of periods of submergence and emergence results in dramatic changes in the magnitude of abiotic and biotic parameters, such as water and nutrient avail-1

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In populations of the Gigartinaceae (Rhodophyta), gametophytes often predominate numerically over tetrasporophytes. Several hypotheses have been proposed to explain this dominance, based on the usually implicit assumption that the stable ratio between gametophytes and tetrasporophytes (G:T ratio) should be 1 if both reproductive phases are ecologically similar. We developed demographic models to test this assumption, for which we considered that both phases are ecologically similar. Defining ecologic similarity for most demographic rates is relatively straightforward, except for rates of spore output. The first set of models considered the same spore output per thallus of both phases as representing ecologic similarity. Model iterations led to stable G:T ratios of 1 for triennial and for perennial thalli, regardless of the initial G:T ratio, but not for annual thalli with initial G:T ratios different from 1. However, equal spore output may not represent ecologic similarity, due to size differences between carpospores and tetraspores. The second set of models considered the lowest possible spore output for each phase, according to the life history of this family: only one carposporangium, with one carpospore, is produced from every two gametophytes and only one tetrasporangium, with four tetraspores, is produced by every tetrasporophyte. Model iterations led to stable G:T ratios of 2.8 for most cases, a ratio of 1 being obtained only every 2 years for annual thalli with an initial G:T ratio of 1. Increasing absolute spore output, without altering the relative output between phases and incorporating density‐independent mortality through a matrix model, given the same mortality rate for both phases, did not modify results. We suggest that the combination of both modeling and field research may uncover more rapidly than otherwise the most relevant ecologic differences between phases, if any, that underlie the G:T ratio observed for a given population.