Friendly competition: evidence for a dilution effect among competitors in a planktonic host-parasite system.

The "dilution effect" concept in disease ecology offers the intriguing possibility that clever manipulation of less competent hosts could reduce disease prevalence in populations of more competent hosts. The basic concept is straightforward: host species vary in suitability (competence) for parasites, and disease transmission decreases when there are more incompetent hosts interacting with vectors or removing free-living stages of a parasite. However, host species also often interact with each other in other ecological ways, e.g., as competitors for resources. The net result of these simultaneous, multiple interactions (disease dilution and resource competition) is challenging to predict. Nonetheless, we see the signature of both roles operating concurrently in a planktonic host-parasite system. We document pronounced spatiotemporal variation in the size of epidemics of a virulent fungus (Metschnikowia bicuspidata) in Midwestern U.S. lake populations of a dominant crustacean grazer (Daphnia dentifera). We show that some of this variation is captured by changes in structure of Daphnia assemblages. Lake-years with smaller epidemics were characterized by assemblages dominated by less suitable hosts ("diluters," D. pulicaria and D. retrocurva, whose suitabilties were determined in lab experiments and field surveys) at the start of epidemics. Furthermore, within a season, less suitable hosts increased as epidemics declined. These observations are consistent with a dilution effect. However, more detailed time series analysis (using multivariate autoregressive models) of three intensively sampled epidemics show the signature of a likely interaction between dilution and resource competition between these Daphnia species. The net outcome of this interaction likely promoted termination of these fungal outbreaks. Should this outcome always arise in "friendly competition" systems where diluting hosts compete with more competent hosts? The answers to this question lie at a frontier of disease ecology.