From tree tops to the ground: reversals to terrestrial habit in Galeandra orchids (Epidendroideae: Catasetinae)

The colonization of the epiphytic niche of tropical forest canopies played an important role in orchids extraordinary diversification in the Neotropics. However, reversals to the terrestrial habit occurred sparsely in species of Epidendroideae. To better understand which factors might have been involved in reversals to terrestrial habits in the predominantly epiphytic Epidendroideae, we investigate Galeandra diversification in time and space. We hypothesized that the reversal to the terrestrial habitat is linked to the origin of open vegetation habitats in South America. We generated a time-calibrated phylogeny based on a matrix of 17 out of 20 species of Galeandra plus outgroups and seven DNA markers. We found that Galeandra originated towards end of the Miocene, about 10 Ma in Amazonia (including the Guiana Shield). The terrestrial clade originated synchronously with the rise of dry vegetation biomes, suggesting that aridification during the last 5 million years dramatically impacted the diversification of epiphytic lineages in the Neotropics. Habit is correlated with floral spur lengths and geographic range size. The longer spurs and narrower ranges are found in epiphytic species: probably adapted to a specialized pollination mode, associated to the long-tongued Euglossini bees, which also prefer forested habits. The terrestrial species presents variable floral spurs and wide distribution ranges, with evidence of self-pollination, suggesting the loss of specialized pollination system and concomitant range expansion.