Multiple hyperparasite species can be associated with a single Ophiocordyceps–ant species pair. Three species – Pseudogibellula formicarum, Torrubiella carnata/liberiana/pseudogibellulae (Hypocreales), and Sporothrix insectorum (Ophiostomatales) – were found on Ophiocordyceps paltothyrei, which infects Palthothyreus tarsatus ants in Ghana (Araújo et al., 2020). Additionally, two recently described hyperparasite species, Niveomyces coronatus and Torrubiellomyces zombiae, are associated with Ophiocordyceps camponoti-floridani, infecting and manipulating the carpenter ant Camponotus floridanus in Florida, USA (Araújo et al., 2022). This suggests that future work on the hyperparasites of zombie-ant fungi will likely reveal a wealth of undescribed species that could be mined for their abilities to affect animal-infecting fungi.
Beyond documenting their diversity, the extent to which hyperparasites affect the life span and transmission of their Ophiocordyceps hosts, as well as their molecular and cellular mechanisms remain to be investigated. A field study on O. camponoti-floridani suggests that both hyperparasites associated with this ant parasite co-occur in the same wilderness areas and harshly limit its transmission chances; when hyperparasitized, 10–40% of ant cadavers were observed with perithecia of O. camponoti–floridani, while this percentage was significantly higher (76%) in non-parasitized Ophiocordyceps (Will et al., 2022). However, only 4% of Ophiocordyceps-manipulated ant cadavers had visible hyperparasite growth within the 1-year timespan of the study. Moreover, while new T. zombiae infections were found year-round, N. coronatus appeared to have a more seasonal occurrence (Will et al., 2022). The disease dynamics of hyperparasites associated with Ophiocordyceps spp. might add a species-specific layer of complexity to the understanding of these multitrophic interactions.