Phylogenetic relationships of Santalales with insights into the origins of holoparasitic Balanophoraceae

Abstract

AbstractTo date molecular data have not revealed the exact phylogenetic position of Balanophoraceae in relation to hemiparasitic Santalales. To elucidate the phylogeny of Santalales and the position of Balanophoraceae, three plastid genes (matK, rbcL, accD), three nuclear genes (SSU and LSU rDNA and RPB2) and one mitochondrial gene (matR) from 197 Santalales samples (including 11 Balanophoraceae species) were analyzed with parsimony, maximum likelihood and Bayesian inference methods. Our results demonstrate that Balanophoraceae is composed of two well‐supported clades: a relatively slow‐evolving one including Dactylantus, Hachettea, and Mystropetalon (Mystropetalaceae) and an extremely fast‐evolving one composed of the remaining Balanophoraceae s.str. Support for monophyly of the two clades was low, thus it appears holoparasitism has arisen twice independently in Santalales. These two clades appeared during a time of great change in the order (ca. 100 Ma) when several major evolutionary innovations emerged, e.g., the root hemiparasites of Santalaceae s.l., the first aerial parasites (Misodendraceae), herbaceous root parasites (Schoepfiaceae), root parasitic Loranthaceae (the ancestors of aerial parasitic mistletoes), as well as the holoparasites in Balanophoraceae and Mystropetalaceae.