Phylogenetic evidence reshapes the taxonomy of Cacao and its allies (Theobroma and Herrania; Malvaceae, Byttnerioideae)

Abstract

Theobroma (Malvaceae, Byttnerioideae), the cacao genus, has a taxonomic history spanning over two centuries. Currently, it comprises 23 species of trees from lowland forests from the Tropical Americas. Herrania, a closely related genus described later, includes 17 additional species commonly referred to as “wild cacaos.” Phylogenetic relationships and morphological circumscriptions between Theobroma and Herrania have been the subject of debate. While Herrania has traditionally been treated as a separate genus based on evident morphological differences in leaf and petal features, it shares similarities with Theobroma in terms of habit, inflorescence, and fruit types. Recent phylogenetic evidence, incorporating a broader taxonomic sampling and a total-evidence analysis, suggested that Theobroma is paraphyletic, with Herrania nested within it. This finding supports the restoration of a classical circumscription of Theobroma wherein Herrania is considered a section of the former genus. Here, we provide a detailed account of the taxonomic history at infrageneric levels and propose one new subsection, two names at new ranks (to better allocate the diversity within T. sect. Herrania), and nine new combinations encompassing this expanded circumscription of Theobroma. In our study, we delimit Theobroma with forty species divided into six sections: T. sect. Glossopetalum (14 spp.), T. sect. Herrania (17 spp.), T. sect. Oreanthes (5 spp.), T. sect. Rhytidocarpus (1 sp.), T. sect. Telmatocarpus (2 spp.), and T. sect. Theobroma (1 sp.). Furthermore, we recognize three subsections within T. sect. Herrania. Alongside these newly proposed changes, we present a section-level identification key and provide diagnostic characters for each taxon.