Ecological outsourcing: a pitcher plant benefits from transferring pre‐digestion of prey to a bat mutualist

Abstract

Summary Mutualisms are interspecific interactions where each of the species involved gains net benefits from the other(s). The exchange of resources and/or services between mutualistic partners often involves tasks that species originally accomplished themselves but which have been taken over by or transferred to the more efficient partner during the evolution of the mutualism. Such ‘ecological outsourcing’ can be seen, for example, in several carnivorous plants that have transferred prey capture and digestion to animal partners. However, the outcome of this transfer and its fitness relevance has rarely been quantified. Using a digestive mutualism between a carnivorous pitcher plant (Nepenthes hemsleyana) and a bat (Kerivoula hardwickii) as a model, we tested the hypothesis that ecological outsourcing is a profitable strategy for the outsourcing partner. To evaluate the value of this mutualism, we conducted a series of field and glasshouse experiments. We measured the benefits of ecological outsourcing by comparing survival, growth, photosynthesis and nutrient content of N. hemsleyana plants fed with bat faeces to those fed with arthropods. To investigate the costs of such outsourcing processes, we repeated the experiment with the closest relative (Nepenthes rafflesiana) that is not adapted to digest bat faeces. We found that N. hemsleyana plants fed with faeces had increased survival, growth and photosynthesis compared to plants fed with arthropods only. On average, plants covered 95% of their nitrogen demand from faeces under strong nutrient deprivation. Despite N. rafflesiana's higher arthropod capture rate, faeces covered a large part of this species’ nutrient demand as well, suggesting low costs for outsourcing. Synthesis. Outsourcing prey capture and digestion to the mutualism partner seems to be a beneficial strategy for N. hemsleyana. It may explain the evolutionary trend of several carnivorous plants to lose their carnivorous traits while increasing their attractiveness to mutualistic partners. On a much broader scale, we propose that ecological outsourcing could be one of the major drivers for the evolution and maintenance of mutualisms.