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Taxonomy of the Orchidaceae : ウィキペディア英語版
Taxonomy of the Orchidaceae
The taxonomy of the Orchidaceae (orchid family) has evolved slowly during the last 250 years, starting with Carolus Linnaeus who in 1753 recognized eight genera.〔Carolus Linnaeus (Carl von Linné). 1753. ''Species Plantarum'', 1st edition, vol. 2, pages 939-954. Holmiae: Impensis Laurentii Salvii (Lars Salvius). (A facsimile with an introduction by William T. Stearn was published by the Ray Society in 1957). (See ''External links'' below).〕 De Jussieu recognized the Orchidaceae as a separate family in his ''Genera Plantarum'' (Jussieu) in 1789.〔Antoine Laurent de Jussieu. 1789. "ORCHIDEAE" pages 64-66. In: '' Genera plantarum :secundum ordines naturales disposita,···'' (See ''External links'' below).〕 Olof Swartz recognized 25 genera in 1800.〔Olof Swartz. 1800. "Afhandling om Orchidernes Slägter och deras Systematiska indelning". ''Kongliga vetenskaps academiens nya handlingar'' 21:115-139. (See ''External links'' below).〕 Louis Claude Richard provided us in 1817 with the descriptive terminology of the orchids.〔Louis Claude Richard. 1817. ''De Orchideis Europaeis annotationes''. Parisiis, ex typographia A. Belin. (See ''External links'' below).〕 The next step was taken in 1830-1840 by John Lindley, who recognized four subfamilies.〔John Lindley. 1830-1840. ''The Genera and Species of Orchidaceous Plants''. Ridgeways, Piccadilly: London, UK.〕 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.〔George Bentham. 1881. page 288. In: "Notes on Orchideae". ''The Journal of the Linnean Society. Botany''. 18(110):281-367. (See ''External links'' below).〕〔Alec M. Pridgeon, Phillip J. Cribb, Mark W. Chase, and Finn N. Rasmussen. 1999-2014. ''Genera Orchidacearum'' Oxford University Press. ISBN 978-0-19-850513-6 (volume 1), ISBN 978-0-19-850710-9 (volume 2), ISBN 978-0-19-850711-6 (volume 3), ISBN 978-0-19-850712-3 (volume 4), ISBN 978-0-19-850713-0 (volume 5), ISBN 978-0-19-964651-7 (volume 6)〕 Then it was published in 1883 in the final volume of ''Genera Plantarum'' (Bentham & Hooker).〔George Bentham and Joseph Dalton Hooker. 1883. ''Genera Plantarum'' (Bentham & Hooker) volume 3, part 2, pages 460-488. L.Reeve & Co.; Williams & Norgate: London, UK. (See ''External links'' below).〕 The next great contributors were Pfitzer (1887), Schlechter (1926), Mansfeld (1937), Dressler and Dodson (1960), Garay (1960, 1972), Vermeulen (1966), again Dressler (1981).〔Robert L. Dressler. 1981. ''The Orchids: Natural History and Classification''. Harvard University Press. ISBN 978-0-674-87525-8. (See ''External links'' below).〕 and Burns-Balogh and Funk (1986). Dressler's 1993 book had considerable influence on later work.〔Robert L. Dressler. 1993. ''Phylogeny and Classification of the Orchid Family''. Cambridge University Press. ISBN 978-0-521-45058-4. 314 pages〕
''Genera Orchidacearum'' was published in 6 volumes over 15 years, from 1999 to 2014.〔 It covers all of the 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.〔Kenneth M. Cameron, Mark W. Chase, W. Mark Whitten, Paul J. Kores, David C. Jarrell, Victor A. Albert, Tomohisa Yukawa, Harold G. Hills and Douglas H. Goldman. 1999. "A phylogenetic analysis of the Orchidaceae: evidence from rbcL nucleotide sequences". ''American Journal of Botany'' 86(2):208-224. (See ''External links'' below).〕 The first classification that was based on cladistic analysis of DNA data was published by Chase et alii in 2003.〔Mark W. Chase, Kenneth M. Cameron, Russell L. Barrett, and John V. Freudenstein. 2003. "DNA data and Orchidaceae systematics: a new phylogenetic classification". pages 69-89. In: Kingsley W. Dixon, Shelagh P. Kell, Russell L. Barrett, and Phillip J. Cribb (editors). 2003. ''Orchid Conservation''. Natural History Publications, Kota Kinabalu, Sabah, Malaysia. ISBN 978-983-812-078-4. (See ''External links'' below).〕
An update to that classification was published by Chase et alii in 2015.〔Mark W. Chase, Kenneth M. Cameron, John V. Freudenstein, Alec M. Pridgeon, Gerardo A. Salazar, Cássio van den Berg, and André Schuiteman. 2015. "An updated classification of Orchidaceae". ''Botanical Journal of the Linnean Society'' 177(2):151-174. (See ''External links'' below).〕 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'', tho most of the vascular plant families had been covered by that time.
Changes in the taxonomy of orchids are so frequent that it is recommended that the regularly updated World Checklist of Selected Plant Families be consulted for the most current generic and species classifications. Wikispecies (Orchidaceae) closely follows this source with modifications as they become accepted. The Plant List shows lists of genera and species, but no other taxonomic information.〔The Plant List: Orchidaceae. (See ''External links'' below).〕
== 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.〔 (See ''External links'' below).〕〔Peter F. Stevens (2001 onwards). "Orchidaceae" At: Angiosperm Phylogeny Website. At: Missouri Botanical Garden Website. (see ''External links'' below)〕
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.〔Kenneth M. Cameron. 2006. "A comparison and combination of plastid atpB and rbcL gene sequences for inferring phylogenetic relationships within Orchidaceae". pages 447-464. In: J. Travis Columbus, Elizabeth A. Friar, J. Mark Porter, Linda M. Prince, and Michael G. Simpson (editors). ''Aliso'' 22(''Monocots: Comparative Biology and Evolution''). 735 pages. Rancho Santa Ana Botanic Garden. printed by Allen Press: USA.〕 This result had only weak maximum parsimony jackknife support, but in a phylogenomic study in 2015, it received strong maximum likelihood bootstrap support.〔Thomas J. Givnish, Daniel Spalink, Mercedes Ames, Stephanie P. Lyon, Steven J. Hunter, Alejandro Zuluaga, William J.D. Iles, Mark A. Clements, Mary T.K. Arroyo, James Leebens-Mack, Lorena Endara, Ricardo Kriebel, Kurt M. Neubig, W. Mark Whitten, Norris H. Williams, and Kenneth M. Cameron. 2015. "Orchid phylogenomics and multiple drivers of their extraordinary diversification". ''Proceedings of the Royal Society, series B'' (biological sciences) 282(1814):pages??. . 〕
Since 2006, phylogenies of two of the subfamilies, Vanilloideae〔Kenneth M. Cameron. 2011. "Vanilloid Orchids: Systematics and Evolution". pages 1-14. In: Eric Odoux and Michel Grisoni (editors). ''Medicinal and Aromatic Plants – Industrial Profiles''. CRC Press: Boca Raton FL, USA.〕〔Kenneth M. Cameron. 2009. "On the value of nuclear and mitochondrial gene sequences for reconstructing the phylogeny of vanilloid orchids (Vanilloideae, Orchidaceae)". ''Annals of Botany'' 104(3):377-385. .〕 and Epidendroideae〔John V. Freudenstein and Mark W. Chase. 2015. "Phylogenetic relationships in Epidendroideae (Orchidaceae), one of the great flowering plant radiations: progressive specialization and diversification". ''Annals of Botany'' 115(4):665-681. .〕 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 problematical. 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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