Fending for oneself or partnering up: Competition between mixo- and heterotrophic ciliates under dynamic resource supply

The outcome of species competition strongly depends on the traits of the competitors and associated trade-offs, as well as on environmental variability. Here we investigate the relevance of consumer trait variation for species coexistence in a ciliate consumer - microalgal prey system under fluctuating regimes of resource supply. We focus on consumer competition and feeding traits, and specifically on the consumers ability to overcome periods of resource limitation by mixotrophy, i.e. the ability of photosynthetic carbon fixation via algal symbionts in addition to phagotrophy. In a 48-day chemostat experiment, we investigated competitive interactions of different heterotrophic and mixotrophic ciliates of the genera Euplotes and Coleps under different resource regimes, providing prey either continuously or in pulses under constant or fluctuating light, entailing periods of resource depletion in fluctuating environments, but overall providing the same amount of prey and light. Although ultimate competition results remained unaffected, population dynamics of mixotrophic and heterotrophic ciliates were significantly altered by resource supply mode. However, the effects differed among species combinations and changed over time. Whether mixotrophs or heterotrophs dominated in competition strongly depended on the genera of the competing species and thus species-specific differences in the minimum resource requirements that are associated with feeding on shared prey, nutrient uptake, light harvesting and access to additional resources such as bacteria. Potential differences in the curvature of the species resource-dependent growth functions may have further mediated the species-specific responses to the different resource supply modes. In addition, while the presence of a heterotrophic competitor may have a direct negative effect on the growth rate of a mixotrophic species through grazing on a shared prey species, its presence may also have an indirect positive effect on the growth rate of the mixotroph by reducing competition between the autotroph and mixotroph for shared nutrients and light. Our study thus demonstrates that complex trophic interactions determine the outcome of competition, which can only be understood by taking on a multidimensional trait perspective.