Competition and habitat suitability: small-scale segregation underpins large-scale coexistence of key species on temperate rocky shores.

Identifying the mechanisms that underpin species richness is one of the central issues of community ecology. On rocky shores in north-western Europe, two key limpet species coexist on the mid- and low shore but are segregated at small scales with respect to habitat. Shortterm and long-term experiments were done to test whether habitat suitability or habitat-specific competition drives the small-scale segregation of these species and therefore underpins their coexistence at larger spatial scales. In a controlled short-term experiment, Patella vulgata was transplanted onto open rock and into pools that either contained Patella ulyssiponensis and/or their mucus or from which P. ulyssiponensis and/or their mucus had been removed. After 2 days, P. vulgata remained in all experimental plots in similar numbers irrespective of treatment indicating that there was no negative response to P. ulyssiponensis, mucus or the pool habitat. In a long-term experiment, cage enclosures containing both species were set up in pools and on open rock over a 6-month period. P. vulgata grew equally well on both open rock and in pools but suffered higher mortality in pools. P. ulyssiponensis showed lower growth rate and higher mortality on open rock than in pools. P. ulyssiponensis exhibited increased growth in higher intraspecific densities on open rock and reduced growth in higher intraspecific densities in pools, indicating some degree of intraspecific facilitation on open rock and intraspecific competition in pools. There was no evidence of interspecific competition either in the short term or in the long term. Results revealed that habitat suitability was the mechanism causing segregation of these species at smaller spatial scales enabling them to coexist at larger spatial scales. Conflicting results in the short-term and long-term experiments highlight the importance of considering the correct temporal extent for experimental tests of hypotheses.