Chytrid infections of Daphnia pulicaria: Development, ecology, pathology and phylogeny of Polycaryum laeve

1. We combined ecological surveys, life table studies, microscopy and molecular sequencing to determine the development, ecology, pathology and phylogeny of Polycaryum laeve, an endoparasite of cladocerans. We report the first records of P. laeve from North America, where we have used a polymerase chain reaction primer and microscopic examination to confirm infections in 14 lakes. Infections are highly pathogenic and caused increased mortality, reduced growth, and reproductive castration in Daphnia pulicaria during life table studies. 2. Biweekly data from Allequash Lake (Wisconsin, U.S.A.) throughout 2003 indicated that fecundity and infection prevalence were inversely correlated. Infection prevalence was highest in late winter and early spring (up to 80%) and lowest during late summer. Epidemics were generally followed by sharp declines in host population density (up to 99%). 3. Within the haemocoel of its host, P. laeve forms thick-walled sporangia, which occur systemically in later stages of infection. Fungal thalli associate closely with muscle fibres and connective tissue, leading to degeneration as the infection becomes advanced. Following death of the host, flagellated zoospores are released through an exit papilla on the sporangium. Based on the infection-induced castration of the host and increases in infection prevalence with Daphnia size, we postulate that transmission is horizontal, but may be indirect through an additional host or free-living stage. 4. Molecular and morphological data clearly indicate that P. laeve belongs in the fungal phylum Chytriodiomycota, order Blastocladiales. Based on ribosomal RNA gene sequences and morphological features, we transfer the genus Polycaryum from the Haplosporidia to the Chytridiomycota, and designate a lectotype and epitype for P. laeve. Considering the high prevalence of P. laeve infection within Daphnia populations, the frequency with which we detected infections among lakes, and the keystone importance of large-bodied Daphnia in aquatic food webs, we suggest that P. laeve may exert a regulatory influence on Daphnia populations in lake ecosystems.