Cacao diseases-the trilogy revisited.

ABSTRACT This paper reviews the significant advances by the diseases themselves, as well as by the scientists, in the intervening period since the disease trilogy was first delimited in 1989. The impact of these diseases, black pod, witches' broom, and frosty pod rot, has increased dramatically. In addition, there have been radical changes in the taxonomic profiles of these pathogens, which have been based on both traditional (morphological, cytological) and modern (molecular) approaches. Black pod is caused by a complex of Phytophthora species, in which P. palmivora still is the most important worldwide. However, recent invasion of the principal cacao-growing countries of West Africa by the more virulent P. megakarya has been cause for concern. The latter evolved in the ancient forests straddling the Cameroon-Nigerian border as a primary coloniser of fallen fruit. Conversely, frosty pod rot, caused by Moniliophthora roreri, and witches' broom, caused by M. (Crinipellis) perniciosa, both neotropical diseases, are hemibiotrophic, coevolved pathogens. Respectively, M. roreri arose on Theobroma gileri in submontane forests on the north-western slopes of the Andes, whereas M. perniciosa developed as a complex of pathotypes with a considerably wider geographic and host range within South America; the cacao pathotype evolved on that host in the Amazon basin. The inter-relationships of these vicariant species and their recent spread are discussed, together with control strategies.