Energy density patterns of nectar resources permit coexistence within a guild of Neotropical flower-visiting bats

Neotropical rainforests support guilds of nectar feeding bats (Phyllostomidae: Glossophaginae) with up to six coexisting species. To analyse guild structure and mechanisms of coexistence in a Costa Rican tropical lowland rainforest, the resource use and morphology of bats were compared to the energetic characteristics of preferred nectar resources and their spatio-temporal distribution. The relative abundance of nectar-feeding bats was determined from mistnet captures over 26 months. Food items were identified by analysis of pollen loads and faecal samples. Phenology, flower density and nectar sugar content of resource plants permitted quantitative estimations of resource availability expressed as energy density (kJ ha(-1) day(-1)) throughout the annual cycle. Four glossophagine bat species co-occurred at La Selva: two permanent residents (Glossophaga commissarisi, Hylonycteris underwoodi) and two seasonal species (Lichonycteris obscura, Lonchophylla robusta) that were found in small numbers during a period of high nectar availability. The two resident species differed in their abundance and in their temporal feeding strategies. After the main flowering peak, the common G. commissarisi shifted to a more frugivorous diet, while the rarer H. underwoodi fed on the few remaining bat-flowers. Resource plant species differed in their energy density by up to two orders of magnitude. Hylonycteris underwoodi visited more often plant species with a low energy yield than G. commissarisi. Because of its smaller body size and a wing morphology that promotes fast flight, H. underwoodi appears to be better adapted to low and scattered nectar resource levels. The two seasonal species differed greatly in body mass, which suggests different strategies for high-quality resource tracking. Large body mass in Lonchophylla robusta provides an energy buffer that permits daily commuting flights between a permanent roost and profitable foraging areas, while the small Lichonycteris obscura seems to track resources nomadically. It is proposed that energy density may be a major niche dimension that restricts access of species to certain habitats and that may profoundly influence the structure of nectar-feeding bat guilds.