Asymmetric coexistence: bidirectional abiotic and biotic effects between goose barnacles and mussels

1. Species coexistence depends on the net effect of interacting species, representing the sum of multiple interaction components that may act simultaneously and vary independently depending on ambient environmental conditions. Consequently, for a comprehensive understanding of the compound nature of species interactions and coexistence, a mechanistic approach that allows a separate evaluation of each interaction component is required. 2. Two sessile filter-feeders, the goose barnacle Capitulum mitella and the mussel Septifer virgatus, coexist on moderately wave-exposed rocky shores in south-western Japan. In the upper intertidal, Capitulum positively influenced Septifer survivorship and growth through amelioration of thermal stress and of physical disturbance. On the other hand, these species are potential competitors as they have similar body sizes and modes of resource utilization. These opposite processes, facilitation and competition, are based on abiotic characteristics and biotic functions of the two species, respectively. 3. In order to quantify the bidirectional abiotic, biotic and net effects, a series of experimental manipulations was conducted involving the use of living neighbours with both abiotic and biotic effects, and artificial mimics to simulate abiotic effects without biotic effects. 4. Capitulum had strong positive abiotic effects on the mussel survivorship in most experimental periods, while the biotic effect was negligible or weakly negative, suggesting that the net effect of Capitulum on mussel survival was largely attributable to the abiotic effect. In contrast, a significantly negative biotic effect on the mussel growth rate was always present, though this was cancelled out by the larger, positive abiotic effect. In the case of Septifer, its abiotic and biotic effects on the survivorship of goose barnacles were negligible, while those on the growth rate showed temporal variation. 5. With respect to the relationship between species interaction and environmental conditions, the strength of abiotic facilitative effect of Capitulum on mussel survival increased with increasing abiotic stress, while the strength of biotic effect was negligible or weakly negative. As regards the effects of mussels on goose barnacles, our study indicated no obvious relationship. These results point to the importance of decomposing interaction for an accurate, mechanistic understanding of species relations and coexistence.