Of mushrooms and chocolate trees: aetiology and phylogeny of witches' broom and frosty pod diseases of cacao

The troubled history of the two major diseases of the chocolate tree (Theobroma cacao) in South America, witches’ broom and frosty pod, is reviewed, concentrating on critical aspects of the aetiology as well as the phylogeny of the causal agents. Both diseases are caused by sister species within the genus Moniliophthora, belonging to the Marasmiaceae family of mushrooms. The witches’ broom pathogen, Moniliophthora perniciosa, evolved on the Amazonian side of the Andes and induces brooms not only in cacao and its relatives in the genera Theobroma and Herrania (Malvaceae), but also in species in the plant families Bignoniaceae, Malpighiaceae and Solanaceae, on which the mushrooms (basidiomata) are produced. Moniliophthora roreri, the type species of the genus, evolved as a pod pathogen on endemic Theobroma species on the western side of the northern Andean Cordillera. Because Moniliophthora was described originally as the asexual form of an unknown basidiomycete, the generic diagnosis is amended here to accommodate species with agaricoid basidiomata. In addition, the new variety M. roreri var. gileri is designated for the morphotype occurring on Theobroma gileri, in northwest Ecuador. Cytology studies indicate that the supposed conidia of M. roreri are, in fact, sexual spores (meiospores) and it is posited that the fruiting structure represents a much-modified mushroom. Finally, based on preliminary data from pathogenicity testing, it is hypothesized that the true causal agent of both diseases is an as yet unidentified infectious agent vectored into the host by the fungus.