A dated phylogeny of the papaya family (Caricaceae) reveals the crop's closest relatives and the family's biogeographic history

Papaya (Carica papaya) is a crop of great economic importance, and the species was among the first plants to have its genome sequenced. However, there has never been a complete species-level phylogeny for the Caricaceae, and the crop's closest relatives are therefore unknown. We investigated the evolution of the Caricaceae based on sequences from all species and genera, the monospecific Carica, African Cylicomorpha with two species, South American Jacaratia and Vasconcellea with together c. 28 species, and Mexican/Guatemalan Jarilla and Horovitzia with four species. Most Caricaceae are trees or shrubs; the species of Jarilla, however, are herbaceous. We generated a matrix of 4711 nuclear and plastid DNA characters and used maximum likelihood (ML) and Bayesian analysis to infer species relationships, rooting trees on the Moringaceae. Divergence times were estimated under relaxed and strict molecular clocks, using different subsets of the data. Ancestral area reconstruction relied on a ML approach. The deepest split in the Caricaceae occurred during the Late Eocene, when the ancestor of the Neotropical clade arrived from Africa. In South America, major diversification events coincide with the Miocene northern Andean uplift and the initial phase of the tectonic collision between South America and Panama resulting in the Panamanian land bridge. Carica papaya is sister to Jarilla/. Horovitzia, and all three diverged from South American Caricaceae in the Oligocene, 27 (22-33) Ma ago, coincident with the early stages of the formation of the Panamanian Isthmus. The discovery that C. papaya is closest to a clade of herbaceous or thin-stemmed species has implications for plant breeders who have so far tried to cross papaya only with woody highland papayas (Vasconcellea).