“…Holospora have been found at low infection prevalences in temperate or elevated locations around the globe ( Fokin et al, 1993 , 2006 , 2004 ; Hori and Fujishima, 2003 ; Serra et al, 2016 ), but can occasionally cause local epidemics ( Fokin and Görtz, 2009 ), which might give a strong selective advantage to resistant variants over susceptible ones. Previous cross-inoculation studies demonstrated the existence of such natural variation, showing that different strains of Paramecium caudatum (and several other species) have different qualitative infection phenotypes, with some strains appearing universally susceptible to infection with Holospora , while others are more difficult to infect or even entirely resistant ( Fujishima and Fujita, 1985 ; Rautian et al, 1993 , 1996 ; Skoblo et al, 1996 ; Fokin et al, 2003b ; Potekhin et al, 2018 ). Previous studies have also compared resistance across independent mating groups or “syngens.” Syngens have long been known for P. caudatum ( Gilman, 1941 ), and although reproductive isolation may not always be complete ( Tsukii and Hiwatashi, 1983 ; Johri et al, 2017 ), genetic analyses identified various clades in P. caudatum that can be considered as independent evolutionary units (IEUs) ( Barth et al, 2006 ; Hori et al, 2006 ; Johri et al, 2017 ; De Souza et al, 2020 ).…”