A competitive hierarchy model integrating roles of physiological competence and competitive ability does not provide a mechanistic explanation for the zonation of three intertidal Fucus species in Europe

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It has been clear for the last 20 yr that both competitive ability and physiological competence on the intertidal gradient of rocky shores determine the zoned distribution of fucoid seaweeds in the North Atlantic. However to this point, there has been no concept integrating these functions for seaweed vegetation in a single mechanistic explanation. Keddy's competitive hierarchy model, which has not been tested in marine systems (i.e. not on seaweed vegetation) provides an explanation for the zonation of plant species on environmental gradients. The model proposes that competitive abilities of species on a stress or resource gradient are inversely related to fundamental niche breadths. We tested 2 assumptions of the model to determine whether it provided a comprehensive explanation of zonation of 3 Fucus species on the island of Helgoland in the North Sea. The 2 assumptions translate into specific predictions for the Fucus assemblage (where F. serratus occurs on the low shore, F. spiralis on the high shore, and F. vesiculosus in between): 1. competitive ranking is F. serratus > F. vesiculosus > F. spiralis; 2. fundamental niche breadth rankings are the reverse of competitive ability rankings. Pairwise competition experiments were done in the field. A highly modified substitutive design was used, in order to take into account the shortcomings of this approach. The empirically derived competitive hierarchy did not fit prediction 1. Transplant experiments with adults and juveniles provided results contrary to prediction 2. Since none of the assumptions of the model fit, it cannot be used to provide a mechanistic explanation for the zonation of Fucus species on Helgoland. Unlike other important models of competition, Keddy's approach does not claim universal validity in all communities. This means that it requires empirical testing in each species assemblage, and more importantly, that the negative outcome reported here does not invalidate the concept in general.

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