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Am J Bot. 2014 Jun 1;101(6):979-1001. Epub 2014 Jun 6.

Speciation via floral heterochrony and presumed mycorrhizal host switching of endemic butterfly orchids on the Azorean archipelago.

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Royal Botanic Gardens Kew, Richmond, Surrey, TW9 3AB, UK.
Royal Botanic Gardens Kew, Richmond, Surrey, TW9 3AB, UK Imperial College London, London, SW7 2AZ, UK.
Department of Biology, University Federico II of Naples, Naples, I-80126, Italy.
CNR-IBBR Institute of Bioscience and BioResources, Via Universita' 133 80055 Portici, Naples, Italy.
Department of Life Sciences, Natural History Museum, Cromwell Road, London, SW7 5BD, UK.
CIBIO Research Center in Biodiversity and Genetic Resources-Azores, Department of Biology, University of the Azores, Rua Mae de Deus 58, Apartado 1422, 9501-801 Ponta Delgada, Portugal.


• Premise of the study: Most orchid species native to the Macaronesian islands reflect immigration from western Europe or North Africa followed by anagenesis. The only putative exception is the butterfly orchids (Platanthera) of the Azores, where three species apparently reflect at least one cladogenetic speciation event. This multidisciplinary study explores the origin, speciation, phenotypic, and genotypic cohesion of these Azorean species and their mainland relatives.• Methods: Plants of Platanthera from 30 localities spanning all nine Azorean islands were compared with those of four continental European relatives for 38 morphometric characters; substantial subsets were also analyzed for plastid microsatellites, and for nrITS of both the orchids and their mycorrhizae.• Key results: Although the three Azorean and four mainland species are all readily distinguished morphometrically using several floral characters, and hybridization appears rare, divergence in ITS and especially plastid sequences is small. Despite occupying similar laurisilva habitats, the Azorean species differ radically in the identities and diversity of their mycorrhizal partners; specialism apparently increases rarity.• Conclusions: Although morphological evidence suggests two invasions of the islands from NW Africa and/or SW Europe, ITS data imply only one. As the molecular data are unable to distinguish among the potential mainland ancestors, two scenarios of relationship are explored that imply different ancestors. Both scenarios require both anagenetic and cladogenetic speciation events, involving homoplastic shifts in overall flower size and (often substantial) changes in the relative dimensions of individual floral organs. Limited genotypic divergence among the three species compared with greater phenotypic divergence suggests comparatively recent speciation. Mycorrhizae may be the most critical factor dictating the respective ecological tolerances, and thus the relative frequencies, of these species. The recent IUCN Red-List amalgamation of Azorean Platanthera taxa into a single species urgently requires reappraisal, as P. micrantha is an excellent indicator species of seminatural laurisilva forest and P. azorica is arguably Europe's rarest orchid.


Platanthera; anagenesis; character conflict; cladogenesis; heterochrony; homoplasy; internal transcribed spacer; migration; morphometrics; mycorrhiza; orchid; phylogeny; plastid haplotypes; speciation; species circumscription

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