Territorial damselfish enhances multi-species co-existence of foraminifera mediated by biotic habitat structuring

The structures of benthic foraminiferal communities inside and outside the territory of the pomacentrid damselfish Stegastes nigricans on coral rocks in a moat of Sesoko Is. (26°38′N, 127°52′E) in Okinawa, Japan were compared. Inside the territory, an algal farm, i.e., a dense stand of a filamentous rhodophyte, Womersleyella setacea, was maintained throughout the year. Outside the territory, in areas subjected to intensive and continuous grazing by various grazers, the flora was composed of mat-like cyanophytes, a prostrate turf-form Padina sp., microscopically thin filaments and scattered filamentous rhodophytes. Algal biomass was greater inside the territory than outside the territory of the damselfish. These differences were reflected by the structure of the associated foraminiferal communities. We classified foraminiferal species based on their microhabitat use around algal communities into four life types: free-living type, crawling type, sedentary type, and sessile type. The abundance of foraminifera, especially the free-living type (e.g., Peneroplis pertusus and Quinqueloculina seminulum) and sedentary type (Rosalina globularis and Cymbaloporetta squammosa), was greater inside the territory of the damselfish than outside the territory. Species richness was also higher inside the territory, mainly due to an increase in species richness of the free-living and sedentary types. These increases in abundance and species richness were caused by habitat structuring. W. setacea is a relatively tall and complicated alga with tangled rhizoids. Its dense stand inside the territory trapped a larger amount of sediment, which provided free-living foraminifera with heterogeneous and stable habitats. In addition, the dense stand of W. setacea provided sedentary-type foraminifera with a large, complex substratum, providing refuge and food inside the territory. This assumption was confirmed by the results of plate experiments showing that foraminiferal community structures were controlled by associations between foraminifera and algae. We showed that habitat-conditioning is an important process in biotic habitat-structuring and that habitat-conditioning by territorial damselfish maintains and enhances multi-species coexistence of foraminifera on coral rocks in a coral reef.