4. Diversity of hyperparasitic fungi

Hyperparasitic fungi are found across the fungal tree of life (Fig. 4), from Cryptomycota to former ‘zygomycetes’ to Basidiomycota (Gleason et al., 2012; Jeffries, 1985; Lutz et al., 2004) (Fig. 5). The genus Trichoderma (Sordariomycetes: Hypocreales) includes the best studied mycoparasites, some of which are hyperparasites of plant pathogens (Brotman et al., 2010; Elad et al., 1980). Hypocreales is an order with 320 genera that are rich in hyperparasites of fungi parasitic on plants, animals, and other fungi (Sung et al., 2007; Wijayawardene et al., 2022) (Fig. 6). Akanthomyces lecanii is a member of this order that exploits hosts in two different kingdoms: the coffee rust fungus, Hemileia vastatrix (Pucciniomycetes: Pucciniales), and the coffee scale insect, Coccus viridis (Vandermeer et al., 2009). Having hosts that themselves are obligate associates with coffee plants as parasites, potentially enables this dynamic hyperparasite to maintain various reservoirs for dispersal through time and physical space in the environment (Jackson et al., 2016). Many other prominent and well-studied groups of hyperparasites are representatives of Dothideomycetes. Some examples of hyperparasites in Dothideomycetes are Ampelomyces spp. (Pleosporales) on powdery mildews (Kiss et al., 2004) and Cladosporium spp. (Capnodiales) on various parasitic hosts (Moricca et al., 2005).