ABSTRACT
Mixotrophic organisms combine light, mineral nutrients, and prey as supplementary resources. Based on theoretical assumptions and field observations, we tested experimentally the hypothesis that mixotrophs may invade established plankton communities depending on the trophic status of the system, and investigated possible effects on food web structure, species diversity, and nutrient dynamics. To test our hypothesis, we inoculated the mixotrophic nanoflagellate Ochromonas tuberculata into established planktonic food webs, consisting of specialist phototrophs, specialist phagotrophs, and bacteria at different supplies of soluble inorganic nutrients and dissolved organic carbon. Oligotrophic systems facilitated the invasion of O. tuberculata in two different ways. First, the combination of photosynthesis and phagotrophy gave mixotrophs a competitive advantage over specialist phototrophs and specialist phagotrophs. Second, low nutrient supplies supported the growth of small plankton organisms that fell into the food size spectrum of mixotrophs. Conversely, high nutrient supplies prevented O. tuberculata from successfully invading the food webs. Two important conclusions were derived from our experiments. First, in contrast to a paradigm of ecology, specialization may not necessarily be the most successful strategy for survival under stable conditions. Indeed, the use of several resources with lower efficiency can be an equally, or even more, successful strategy in nature. Second, when limiting nutrients promote the growth of bacterio- and picophytoplankton, invading mixotrophs may have a habitat-ameliorating effect for higher trophic levels, gauged in terms of food quantity and quality. Using given resources more efficiently, O. tuberculata generated higher biomasses and expressed an increased nutritional value for potential planktivores, due to decreased cellular carbon to phosphorus (C:P) ratios compared to specialized plankton taxa. Our findings may help to explain why energy transfer efficiency between phytoplankton and higher trophic levels is generally higher in oligotrophic systems than in nutrient rich environments.