Since the last classification of Orchidaceae in 2003, there has been major progress in the determination of relationships, and we present here a revised classification including a list of all 736 currently recognized genera. A number of generic changes have occurred in Orchideae (Orchidoideae), but the majority of changes have occurred in Epidendroideae. In the latter, almost all of the problematic placements recognized in the previous classification 11 years ago have now been resolved. In Epidendroideae, we have recognized three new tribes (relative to the last classification): Thaieae (monogeneric) for Thaia, which was previously considered to be the only taxon incertae sedis; Xerorchideae (monogeneric) for Xerorchis; and Wullschlaegelieae for achlorophyllous Wullschlaegelia, which had tentatively been placed in Calypsoeae. Another genus, Devogelia, takes the place of Thaia as incertae sedis in Epidendroideae. Gastrodieae are clearly placed among the tribes in the neottioid grade, with Neottieae sister to the remainder of Epidendroideae. Arethuseae are sister to the rest of the higher Epidendroideae, which is unsurprising given their mostly soft pollinia. Tribal relationships within Epidendroideae have been much clarified by analyses of multiple plastid DNA regions and the low-copy nuclear gene Xdh. Four major clades within the remainder of Epidendroideae are recognized: Vandeae/Podochileae/Collabieae, Cymbidieae, Malaxideae and Epidendreae, the last now including Calypsoinae (previously recognized as a tribe on its own) and Agrostophyllinae s.s. Agrostophyllinae and Collabiinae were unplaced subtribes in the 2003 classification. The former are now split between two subtribes, Agrostophyllinae s.s. and Adrorhizinae, the first now included in Epidendreae and the second in Vandeae. Collabiinae, also probably related to Vandeae, are now elevated to a tribe along with Podochileae. Malaxis and relatives are placed in Malaxidinae and included with Dendrobiinae in Malaxideae. The increased resolution and content of larger clades, recognized here as tribes, do not support the 'phylads' in Epidendroideae proposed 22 years ago by Dressler.
Epiphytes are a characteristic component of tropical rainforests. Out of the 25 000 orchid species currently known to science, more than 70% live in tree canopies. Understanding when and how these orchids diversified is vital to understanding the history of epiphytic biomes. We investigated whether orchids managed to radiate so explosively owing to their predominantly epiphytic habit and/or their specialized pollinator systems by testing these hypotheses from a statistical and phylogenetic standpoint. For the first approach, species numbers of 100 randomly chosen epiphytic and terrestrial genera were compared. Furthermore, the mean number of pollinators per orchid species within the five subfamilies was calculated and correlated with their time of diversification and species richness. In the second approach, molecular epiphytic orchid phylogenies were screened for clades with specific suites of epiphytic adaptations. Epiphytic genera were found to be significantly richer in species than terrestrial genera both for orchids and non-orchids. No evidence was found for a positive association between pollinator specialization and orchid species richness. Repeated associations between a small body size, short life cycle and specialized clinging roots of twig epiphytes in Bulbophyllinae and Oncidiinae were discovered. The development of twig epiphytism in the first group seems repeatedly correlated with speciation bursts.
Herbarium specimens provide verifiable and citable evidence of the occurrence of particular plants at particular points in space and time, and are vital resources for assessing extinction risk in the tropics, where plant diversity and threats to plants are greatest. We reviewed approaches to assessing extinction risk in response to the Convention on Biological Diversity's Global Strategy for Plant Conservation Target 2: an assessment of the conservation status of all known plant species by 2020. We tested five alternative approaches, using herbarium-derived data for trees, shrubs and herbs in five different plant groups from temperate and tropical regions. All species were previously fully assessed for the IUCN Red List. We found significant variation in the accuracy with which different approaches classified species as threatened or not threatened. Accuracy was highest for the machine learning model (90%) but the least data-intensive approach also performed well (82%). Despite concerns about spatial, temporal and taxonomic biases and uncertainties in herbarium data, when specimens represent the best available evidence for particular species, their use as a basis for extinction risk assessment is appropriate, necessary and urgent. Resourcing herbaria to maintain, increase and disseminate their specimen data is essential to guide and focus conservation action.This article is part of the theme issue ‘Biological collections for understanding biodiversity in the Anthropocene’.
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