Molecular phylogeny of the tribe Danaideae (Rubiaceae: Rubioideae): Another example of out-of-Madagascar dispersal

摘要

Extensive efforts have been made to resolve the phylogeny of the large coffee family (Rubiaceae) based on molecular data. As a result, several small tribes have been described, but the phylogenies and generic delimitations for many of these groups remain unclear. This study focuses on the small tribe Danaideae that belongs to subfamily Rubioideae and whose generic limits have not previously been addressed with molecular data. It is the sole rubiaceous tribe distributed almost entirely in the Western Indian Ocean region, with the exception of the East African Danais xanthorrhoea. The tribe consists of three genera: Danais, Payera (including the monotypic genus Coursiana), and Schismatoclada. We present the first molecular phylogenetic study of Danaideae including representatives from all three genera and using Bayesian and maximum parsimony methods and sequence data from nuclear DNA (nrITS) and chloroplast DNA (petD, psbA-trnH, rpl32-trnL(UAG), rps16). Our main objectives were to rigorously test the monophyly of Danaideae as currently circumscribed and assess phylogenetic relationships within the tribe. The findings of this study shed light on the colonization history of the tribe. Our analyses reaffirm the monophyly of Danaideae and Danais but reveal the paraphyly of Payera and Schismatoclada. The close relationship between the three Danaideae genera and Coursiana is supported. However, we found very little support for the inclusion of the latter genus in Payera as proposed earlier. The tribe is resolved in two morphologically distinct major lineages, the highly supported Damns clade with lianescent habit (= Danais sensu Buchner & Puff) and the Payera-Schismatoclada clade with arborescent habit. The Malagasy and Mauritian specimens of Danais fragrans are not closely related, and we restrict D. fragrans to the Mauritian taxa and resurrect Danais lyallii Baker to accommodate the Malagasy D. fragrans. According to our analysis. Madagascar is the origin of all species of Danaideae occurring in the Comoro archipelago, East Africa, and Mauritius. The Mauritian and East African Danais each is the result of a single colonization event, while there were at least two independent colonization events to the Comoros.