Taxonomy of the Orchidaceae

The taxonomy of the Orchidaceae (orchid family) has evolved slowly during the last 250 years, starting with Carl Linnaeus who in 1753 recognized eight genera. De Jussieu recognized the Orchidaceae as a separate family in his Genera Plantarum in 1789. Olof Swartz recognized 25 genera in 1800. Louis Claude Richard provided us in 1817 with the descriptive terminology of the orchids. (See External links below). The next step was taken in 1830-1840 by John Lindley, who recognized four subfamilies. He is generally recognized as the father of orchid taxonomy. The next important step was taken by George Bentham with a new classification, recognizing subtribes for the first time. This classification was first presented in a paper that Bentham read to the Royal Society in 1881. Then it was published in 1883 in the final volume of Genera Plantarum. The next great contributors were Pfitzer (1887), Schlechter (1926), Mansfeld (1937), Dressler and Dodson (1960), Garay (1960, 1972), Vermeulen (1966), again Dressler (1981). and Burns-Balogh and Funk (1986). Dressler's 1993 book had considerable influence on later work. Genera Orchidacearum was published in 6 volumes over 15 years, from 1999 to 2014. It covers all of the known orchids, including a description of each genus. It reflects the considerable progress in orchid taxonomy that had been made since Dressler published his classification in 1993. In the 1990s, orchid taxonomy began to be influenced by molecular phylogenetics based on DNA sequences. The first molecular phylogenetic study to include a substantial sample of orchids was published in 1999. The first classification that was based on cladistic analysis of DNA data was published by Chase et alii in 2003. An update to that classification was published by Chase et alii in 2015. This classification takes a different approach from Genera Orchidacearum, by consolidating many of the tribes and subtribes, and by recognizing very widely circumscribed genera. As of 2015, Orchidaceae was not yet covered in The Families and Genera of Vascular Plants, though most of the vascular plant families had been covered by that time. The number of genera recognized in the family has varied from one classification to another. In Genera Orchidacearum, many genera were consolidated, reducing their number to 765, smaller than in any previous modern classification. In 2015, Chase et alii merged even more genera, reducing their number to 736. Useful resources include the World Checklist of Selected Plant Families. Wikispecies (Orchidaceae) closely follows this source with modifications as they become accepted. The Plant List also has lists of genera and species, but no other taxonomic information.

History The following taxonomy follows largely the classification system of Robert Louis Dressler, an orchid specialist and adjunct curator at the Florida Museum of Natural History. This classification, published in 1981 in the book The Orchids: Natural History and Classification, was widely accepted by botanists and growers before the publication of Genera Orchidacearum. The initial scheme of 1981 was modified in 1986, twice in 1990, and then again in 1993. This comprehensive classification relies heavily on morphology, especially a few key characters, such as anther configuration and pollinarium structure. Consequently, many of the taxa are not monophyletic. Cladistic analyses, especially those based on molecular data, provide a firmer basis for classification than intuition, and the certainty (or uncertainty) of conclusions can be quantified by measures of statistical support. While our understanding of orchid phylogeny has greatly improved in recent years, the elucidation of orchid relationships is still ongoing. When Dressler published his classification in 1993, the relationships of orchids to other monocots was still unknown. Some of the first molecular phylogenetic studies of monocots resolved the Orchidaceae as sister to the astelioid clade of the order Asparagales, but this result never had strong statistical support. It is now known that Orchidaceae is the most basal clade in Asparagales, with the astelioid clade diverging next. According to cladistic analyses based on morphological character states or on nucleotide sequences, the orchid family is a monophyletic group. The subfamilies recognized by Dressler, however, were not all monophyletic. Dressler's delimitation of subfamilies was contradicted by subsequent studies of mitochondrial, chloroplast, and nuclear DNA sequences. In 2003, a new phylogenetic classification divided Orchidaceae into five subfamilies: Apostasioideae, Vanilloideae, Cypripedioideae, Orchidoideae, and Epidendroideae. These five subfamilies were all strongly supported as monophyletic groups in subsequent studies. In 2003, the position of Vanilloideae remained equivocal. The subfamilies Orchidoideae and Epidendroideae clearly formed a monophyletic group and Dressler believed that their closest relative was Vanilloideae. In 2006, a study based on the plastid genes rbcL and atpB found the closest relative of this pair to be Cypripedioideae, rather than Vanilloideae. This result had only weak maximum parsimony jackknife support, but in a phylogenomic study in 2015, it received strong maximum likelihood bootstrap support. Since 2006, phylogenies of two of the subfamilies, Vanilloideae and Epidendroideae have been published. Phylogenies of several tribes and subtribes have also been published. Compared to previous classifications, more of the tribes and subtribes of Dressler were monophyletic, but not all of them were supported by subsequent studies. In the classification that was published in 2015, the authors expressed doubt about their division of the tribes Orchideae and Vandeae into subtribes. The placement of the genera Pachites, Holothrix, and Hederorkis is especially problematic. The monophyly of the subtribe Cranichidinae is also in doubt. These authors singled out the tribe Podochileae, as well as the subtribes Oncidiinae, Goodyerinae, and Angraecinae as being in special need of phylogenetic study. The basal epidendroids, especially the tribe Gastrodieae, remain poorly sampled in phylogenetic studies.

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