4.1.2. Hyperparasites of black mildews

An approximate 200 species of fungi are reported to be hyperparasitic on colonies of black mildews (Sordariomycetes: Meliolales). They include organisms from diverse systematic groups, and therefore comprise species producing a high diversity of reproductive structures, such as synnemata, pycnidia, apothecia, perithecia, and catathecia, among others (Bermúdez-Cova et al., 2022). The most common hyperparasites of black mildews are species of the genera AtractilinaSpiropes (Pezizomycotina incertae sedis ), Dimerosporiella (Sordariomycetes: Hypocreales), and Trichothyrium (Dothideomycetes: Microthyriales) (Bermúdez-Cova et al., 2022; Deighton and Pirozynski, 1972; Ellis, 1968; Pirozynski, 1977; Rossman, 1987; Rossman et al., 1999). Hyperparasites of Meliolales are contact-biotrophic fungi and prevent their host from producing spores and ascomata (Stevens, 1918; Toro, 1952). The current systematic position of almost all species of hyperparasitic fungi of Meliolales is unknown due to two reasons: the description of many of these predated the molecular era and technical problems make DNA extractions and PCR amplifications challenging (see 7. Molecular studies of hyperparasitic fungi).