Systématique évolutive et biogéographie de Angraecum (Orchidaceae, Angraecinae) à Madagascar

Angraecum is a group of tropical orchids that includes ca. 221 species distributed in Sub-Saharan Africa, the western Indian Ocean region, and Sri Lanka. At least half of the species are found in Madagascar, 90% being endemic to the island. Taxonomic studies of Angraecum have always been problematic because of the great morphological diversity of the group. To facilitate classification, sections have been proposed, the best-known system to date being that of Garay (1973) that subdivides the genus into 19 sections. Previous phylogenetic studies had shown that genus Angraecum and Garay’s sections are not monophyletic. However, no systematic review was made because of a reduced species sample in these analyses. Using a greater sampling from Madagascar and adding morphological characters to the analyses, we brought greater resolution to the phylogenetic reconstruction of the group. This resolution mainly concerns the deeper nodes representing different clades within Angraecum, which basically correspond to natural sections. By using these clades, we redefined 14 sections and recognized five new ones. Using this phylogeny of Angraecum, we evaluated species diversification using macroevolutionary methods, essentially the effect of floral traits in speciation. The great diversity of Angraecum species in Madagascar, the high endemicity, and the geology and paleoclimate histories allowed us to evaluate diversification patterns within the genus as well as sub-tribe Angraecinae. The model of diversification in Angraecinae follows that of most tropical rain forest taxa, which results from the gradual accumulation of species through time and not from a rapid adaptive radiation, as is often the case for Malagasy fauna lineages. Several morphological characters are involved in the diversification of Angraecum. The beginning of Angraecum diversification in Madagascar coincided with the progressive movement of the island northwards, the establishment of a monsoon regime in the northern part of the island during the Miocene, and the expansion of the Malagasy rainforest during that period. Our historical biogeographic study of Angraecinae suggests a Malagasy origin of the subtribe Angraecinae and Angraecum. We observed out-of Madagascar long-distance dispersal in Angraecum. The north-eastern Malagasy rainforest is the center of species radiation for the genus. The first dispersal event within the island started in the late Miocene. This event was a migration from the north to the central highland. The majority of long-distance dispersal events outside Madagascar occurred during the Pliocene-Pleistocene, originating from either the center or the North East of the island. There were multiple independent dispersals to East Africa and the Comoros, and to the Mascarenes. A single founder-effect event in section Hadrangis is observed in the Mascarenes. The cyclonic seasons play a significant role in long-distance dispersal of orchid seeds, as compared to prevailing winds in the western Indian Ocean region, essentially trade wind and monsoon. Ecological niche similarity favored the expansion of Angraecum species in the Comoros and Mascarene archipelagos.

Keyphrases: Angraecum, Angraecinae, Diversification, Accumulation of lineages through time, Angraecum, Angraecinae, Biogeography, Delimitation of sections, Diversification, Phylogeny, Systematic, Taxonomy