No abstract

Isolated oceanic archipelagos are natural laboratories of evolution, ideally suited for in situ studies of speciation processes, and the recent availability of checklist data for many archipelagos worldwide (e.g. Izquierdo et al., 2004;

Background and Aims. The Macaronesian islands represent an excellent crucible for exploring speciation. This dominantly phenotypic study complements a separate genotypic study, together designed to identify and circumscribe Platanthera species (butterfly-orchids) on the Azores, and to determine their geographic origin(s) and underlying speciation mechanism(s).Methods. 216 individuals of Platanthera from 30 Azorean localities spanning all nine Azorean islands were measured for 38 morphological characters, supported by light and scanning electron microscopy of selected flowers. They are compared through detailed multivariate and univariate analyses with four widespread continental European relatives in the P. bifolia-chlorantha aggregate, represented by 154 plants from 25 populations, and with the highly misleading original taxonomic descriptions. Physiographic and ecological data were also recorded for each study population.Key Results. Despite limited genetic divergence, detailed phenotypic survey reveals not one or two but three discrete endemic species of Platanthera that are readily distinguished using several characters, most floral: P. pollostantha (newly named, formerly P. micrantha) occupies the widest range of habitats and altitudes and occurs on all nine islands; P. micrantha (formerly P. azorica) occurs on eight islands but is restricted to small, scattered populations in laurisilva scrub; the true P. azorica appears confined to a single volcanigenic ridge on the central island of São Jorge.Conclusions. Although hybridity seems low, the excess of phenotypic over genotypic divergence suggests comparatively recent speciation. The most probable of several credible scenarios is that Azorean Platantheras represent a single migration to the archipelago of airborne seed from ancestral population(s) located in southwest Europe rather than North America, originating from within the P. bifolia-chlorantha aggregate. We hypothesise that an initial anagenetic speciation event, aided by the founder effect, was followed by the independent origins of at least one of the two rarer endemic species from within the first-formed endemic species, via a cladogenetic speciation process that involved radical shifts in floral development, considerable phenotypic convergence, and increased mycorrhizal specificity. The recent amalgamation by IUCN of Azorean Platantheras into a single putative species on their Red List urgently requires overruling, as (a) P. azorica is arguably Europe’s rarest bona fide orchid species and (b) the almost equally rare P. micrantha is one of the best indicators of semi-natural laurisilva habitats remaining on the Azores. Both species are threatened by habitat destruction and invasive alien plants. These orchids constitute a model system that illustrates the general advantages of circumscribing species by prioritising field-based over herbarium-based morphological approaches.

• 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.

The current study illustrates the utility of MSG data for resolving relationships in lineages that have undergone recent, rapid diversification resulting in extensive ecological and morphological diversity. We suggest that a similar approach may prove generally useful for other rapid plant radiations where resolving phylogeny has been difficult.