Editing Orchid (section)

== Taxonomy ==
{{Main|Taxonomy of the Orchid family}}

The [[Taxonomy (biology)|taxonomy]] of this family is in constant flux, as new studies continue to clarify the relationships between species and groups of species, allowing more [[taxa]] at several [[Taxonomic rank|ranks]] to be recognized. The Orchidaceae is currently placed in the [[Order (biology)|order]] [[Asparagales]] by the [[APG III system]] of 2009.<ref name=APGIII2009 />

Five [[subfamilies]] are recognised. The [[cladogram]] below was made according to the [[APG system]] of 1998. It represents the view that most botanists had held up to that time. It was supported by [[Plant morphology|morphological]] [[Research|studies]], but never received strong support in [[molecular phylogenetic]] studies.

{{clade|style=font-size:80%;line-height:75%
|1={{clade|1='''[[Apostasioideae]]''': 2 genera and 16 species, south-eastern [[Asia]]
|2={{clade|1='''[[Cypripedioideae]]''': 5 genera and 130 species, from the temperate regions of the world, as well as tropical [[Americas|America]] and tropical Asia|label2=Monandrae
|2={{clade|1='''[[Vanilloideae]]''': 15 genera and 180 species, humid tropical and subtropical regions, eastern [[North America]]
|2={{clade|1='''[[Epidendroideae]]''': more than 500 genera and more or less 20,000 species, cosmopolitan
|2='''[[Orchidoideae]]''': 208 genera and 3,630 species, cosmopolitan
}}}}}}}}}}

In 2015, a [[phylogenetic]] study<ref>{{Cite journal |author1=Guillaume Chomicki |author2=Luc P.R. Bidel |author3=Feng Ming |author4=Mario Coiro |author5=Xuan Zhang |author6=Yaofeng Wang |author7=Yves Baissac |author8=Christian Jay-Allemand |author9=Susanne S. Renner |name-list-style=amp | date = 2015 | title = The velamen protects photosynthetic orchid roots against UV-B damage, and a large dated phylogeny implies multiple gains and losses of this function during the Cenozoic | journal = New Phytologist | volume = 205 | issue = 3 | pages = 1330–1341 | doi=10.1111/nph.13106|pmid=25345817 |doi-access=free |bibcode=2015NewPh.205.1330C }}</ref> showed strong [[Resampling (statistics)|statistical support]] for the following [[topology]] of the orchid [[Phylogenetic tree|tree]], using 9 [[Base pair|kb]] of [[plastid]] and [[Cell nucleus|nuclear]] [[DNA]] from 7 [[gene]]s, a topology that was confirmed by a [[phylogenomic]] study in the same year.<ref>{{cite journal | last1 = Givnish | first1 = Thomas J. | last2 = Spalink | first2 = Daniel | last3 = Ames | first3 = Mercedes | last4 = Lyon | first4 = Stephanie P. | last5 = Hunter | first5 = Steven J. | last6 = Zuluaga | first6 = Alejandro | last7 = Iles | first7 = William J.D. | last8 = Clements | first8 = Mark A. | last9 = Arroyo | first9 = Mary T.K. | last10 = Leebens-Mack | first10 = James | last11 = Endara | first11 = Lorena | last12 = Kriebel | first12 = Ricardo | last13 = Neubig | first13 = Kurt M. | last14 = Whitten | first14 = W. Mark | last15 = Williams | first15 = Norris H. | last16 = Cameron | first16 = Kenneth M. | year = 2015 | title = Orchid phylogenomics and multiple drivers of their extraordinary diversification | journal = Proceedings of the Royal Society B: Biological Sciences | volume =  282| issue = 1814| page =  20151553| doi = 10.1098/rspb.2015.1553 | pmid = 26311671 | pmc = 4571710 | doi-access = free }}</ref>

{{clade|style=font-size:80%;line-height:75%
|1={{clade|1='''[[Apostasioideae]]'''
|2={{clade|1='''[[Vanilloideae]]'''
|2={{clade|1='''[[Cypripedioideae]]'''
|2={{clade|1='''[[Epidendroideae]]'''
|2='''[[Orchidoideae]]'''
}}}}}}}}}}

=== Evolution ===
[[File:The Orchid Tree of Life v.2.0 (en).jpg|thumb|Artist's interpretation of orchid phylogeny, showing the evolution of the family]]

It has been hypothesised that the origin of the orchids goes back much longer than originally expected.<ref name="Origin of the orchids">{{cite journal |author1=Santiago R. Ramírez |author2=Barbara Gravendeel |author3=Rodrigo B. Singer |author4=Charles R. Marshall |author5=Naomi E. Pierce | title=Dating the origin of the Orchidaceae from a fossil orchid with its pollinator| journal=Nature | date=30 August 2007| volume=448| pages=1042–5| doi=10.1038/nature06039 | pmid=17728756 | issue=7157|bibcode=2007Natur.448.1042R }}</ref> An extinct species of stingless bee, ''Proplebeia dominicana'', was found trapped in [[Miocene]] [[amber]] from about 15–20&nbsp;million years ago. The bee was carrying [[pollen]] of a previously unknown orchid taxon, ''[[Meliorchis caribea]]'', on its wings. This find is the first evidence of fossilised orchids to date<ref name="Origin of the orchids" /> and shows insects were active [[pollinator]]s of orchids then. This extinct orchid, ''M. caribea'', has been placed within the extant tribe [[Cranichideae]], subtribe [[Goodyerinae]] (subfamily [[Orchidoideae]]). An even older orchid species, ''[[Succinanthera baltica]]'', was described from the [[Eocene]] [[Baltic amber]] by [[George Poinar Jr.|Poinar]] & Rasmussen (2017).<ref>{{Cite journal|author1=George Poinar, Jr. |author2=Finn N. Rasmussen |year=2017 |title=Orchids from the past, with a new species in Baltic amber |journal=Botanical Journal of the Linnean Society |volume=183 |issue=3 |pages=327–333 |doi=10.1093/botlinnean/bow018 |doi-access=free }}</ref>

Genetic sequencing indicates orchids may have arisen earlier, 76 to 84&nbsp;million years ago during the [[Late Cretaceous]].<ref>[http://ir.lib.ncku.edu.tw/bitstream/987654321/108263/2/An%20overview%20of%20the%20Phalaenopsis%20orchid%20genome%20by%20BAC%20end%20sequence%20analysis.pdf "An overview of the ''Phalaenopsis'' orchid genome by BAC sequence analysis" (pdf format)] {{Webarchive|url=https://web.archive.org/web/20140320133049/http://ir.lib.ncku.edu.tw/bitstream/987654321/108263/2/An%20overview%20of%20the%20Phalaenopsis%20orchid%20genome%20by%20BAC%20end%20sequence%20analysis.pdf |date=20 March 2014 }}.</ref> According to [[Mark W. Chase]] ''et al.'' (2001), the overall biogeography and phylogenetic patterns of Orchidaceae show they are even older and may go back roughly 100&nbsp;million years.<ref>{{cite book |author=Mark W. Chase |year=2001 |chapter=The origin and biogeography of Orchidaceae |editor=A. M. Pridgeon |editor2=P. J. Cribb |editor3=M. W. Chase |editor4=F. Rasmussen |series=Genera Orchidacearum |volume=2 |title=Orchidoideae (Part 1) |pages=1–5 |publisher=[[Oxford University Press]] |isbn=978-0-19-850710-9|author-link=Mark W. Chase }}</ref>

Using the [[molecular clock]] method, it was possible to determine the age of the major branches of the orchid family. This also confirmed that the subfamily [[Vanilloideae]] is a branch at the basal dichotomy of the [[monandrous]] orchids, and must have evolved very early in the evolution of the family. Since this subfamily occurs worldwide in tropical and subtropical regions, from tropical America to tropical Asia, New Guinea and West Africa, and the continents began to split about 100&nbsp;million years ago, significant biotic exchange must have occurred after this split (since the age of ''Vanilla'' is estimated at 60 to 70&nbsp;million years). Recent biogeographic studies conducted on densely sampled phylogenies indicated that the [[most recent common ancestor]] of all extant orchids probably originated somewhere 83 million years ago in the supercontinent [[Laurasia]].<ref name="Perez-Escobar-2024">{{cite journal |last1=Pérez-Escobar |first1=Oscar A. |last2=Bogarín |first2=Diego |last3=Przelomska |first3=Natalia A. S. |last4=Ackerman |first4=James D. |last5=Balbuena |first5=Juan A. |last6=Bellot |first6=Sidonie |last7=Bühlmann |first7=Roland P. |last8=Cabrera |first8=Betsaida |last9=Cano |first9=Jose Aguilar |last10=Charitonidou |first10=Martha |last11=Chomicki |first11=Guillaume |last12=Clements |first12=Mark A. |last13=Cribb |first13=Phillip |last14=Fernández |first14=Melania |last15=Flanagan |first15=Nicola S. |last16=Gravendeel |first16=Barbara |last17=Hágsater |first17=Eric |last18=Halley |first18=John M. |last19=Hu |first19=Ai-Qun |last20=Jaramillo |first20=Carlos |last21=Mauad |first21=Anna Victoria |last22=Maurin |first22=Olivier |last23=Müntz |first23=Robert |last24=Leitch |first24=Ilia J. |last25=Li |first25=Lan |last26=Negrão |first26=Raquel |last27=Oses |first27=Lizbeth |last28=Phillips |first28=Charlotte |last29=Rincon |first29=Milton |last30=Salazar |first30=Gerardo A. |last31=Simpson |first31=Lalita |last32=Smidt |first32=Eric |last33=Solano-Gomez |first33=Rodolfo |last34=Parra-Sánchez |first34=Edicson |last35=Tremblay |first35=Raymond L. |last36=van den Berg |first36=Cassio |last37=Tamayo |first37=Boris Stefan Villanueva |last38=Zuluaga |first38=Alejandro |last39=Zuntini |first39=Alexandre R. |last40=Chase |first40=Mark W. |last41=Fay |first41=Michael F. |last42=Condamine |first42=Fabien L. |last43=Forest |first43=Felix |last44=Nargar |first44=Katharina |last45=Renner |first45=Susanne S. |last46=Baker |first46=William J. |last47=Antonelli |first47=Alexandre |title=The origin and speciation of orchids |journal=New Phytologist |date=April 2024 |volume=242 |issue=2 |pages=700–716 |doi=10.1111/nph.19580 |pmid=38382573 |bibcode=2024NewPh.242..700P |hdl=10550/97112 |hdl-access=free }}</ref> Despite their long evolutionary history on Earth, the extant orchid diversity is also inferred to have originated during the last 5 million years,<ref name="Perez-Escobar-2024" /> with the American and Asian tropics as the geographic areas exhibiting the highest speciation rates (i.e., number of speciation events per million years) on Earth.

Genome duplication occurred prior to the divergence of this taxon.<ref>{{cite journal|doi=10.1038/nature23897|pmid=28902843|title=The Apostasia genome and the evolution of orchids|journal=Nature|volume=549|issue=7672|pages=379–383|year=2017|last1=Zhang|first1=Guo-Qiang|last2=Liu|first2=Ke-Wei|last3=Li|first3=Zhen|last4=Lohaus|first4=Rolf|last5=Hsiao|first5=Yu-Yun|last6=Niu|first6=Shan-Ce|last7=Wang|first7=Jie-Yu|last8=Lin|first8=Yao-Cheng|last9=Xu|first9=Qing|last10=Chen|first10=Li-Jun|last11=Yoshida|first11=Kouki|last12=Fujiwara|first12=Sumire|last13=Wang|first13=Zhi-Wen|last14=Zhang|first14=Yong-Qiang|last15=Mitsuda|first15=Nobutaka|last16=Wang|first16=Meina|last17=Liu|first17=Guo-Hui|last18=Pecoraro|first18=Lorenzo|last19=Huang|first19=Hui-Xia|last20=Xiao|first20=Xin-Ju|last21=Lin|first21=Min|last22=Wu|first22=Xin-Yi|last23=Wu|first23=Wan-Lin|last24=Chen|first24=You-Yi|last25=Chang|first25=Song-Bin|last26=Sakamoto|first26=Shingo|last27=Ohme-Takagi|first27=Masaru|last28=Yagi|first28=Masafumi|last29=Zeng|first29=Si-Jin|last30=Shen|first30=Ching-Yu|pmc=7416622|bibcode=2017Natur.549..379Z|display-authors=29|url=https://biblio.ugent.be/publication/8533332/file/8533333.pdf}}</ref>

=== Genera ===
{{Main|List of Orchidaceae genera}}
There are around 800 genera of orchids. The following are amongst the most notable genera of the orchid family:<ref>{{cite book |doi=10.1016/B978-0-12-374380-0.50007-5 |chapter=Diversity and Classification of Flowering Plants |title=Plant Systematics |date=2010 |last1=Simpson |first1=Michael G. |pages=181–274 |isbn=978-0-12-374380-0 }}</ref>
{{div col|colwidth=9em}}
* ''[[Aa (plant)|Aa]]''
* ''[[Abdominea]]''
* ''[[Acampe]]''
* ''[[Acanthophippium]]''
* ''[[Aceratorchis]]''
* ''[[Acianthus]]''
* ''[[Acineta]]''
* ''[[Acrorchis]]''
* ''[[Ada (plant)|Ada]]''
* ''[[Aerangis]]''
* ''[[Aeranthes]]''
* ''[[Aerides]]''
* ''[[Aganisia]]''
* ''[[Agrostophyllum]]''
* ''[[Anacamptis]]''
* ''[[Ancistrochilus]]''
* ''[[Angraecum]]''
* ''[[Anguloa]]''
* ''[[Ansellia]]''
* ''[[Aorchis]]''
* ''[[Aplectrum]]''
* ''[[Arachnis (plant)|Arachnis]]''
* ''[[Arethusa (plant)|Arethusa]]''
* ''[[Armodorum]]''
* ''[[Ascoglossum]]''
* ''[[Australorchis]]''
* ''[[Auxopus]]''
* ''[[Barkeria]]''
* ''[[Bartholina]]''
* ''[[Beloglottis]]''
* ''[[Biermannia]]''
* ''[[Bletilla]]''
* ''[[Brassavola]]''
* ''[[Brassia]]''
* ''[[Bulbophyllum]]''
* ''[[Calanthe]]''
* ''[[Calypso (plant)|Calypso]]''
* ''[[Catasetum]]''
* ''[[Cattleya]]''
* ''[[Chiloschista]]''
* ''[[Cirrhopetalum]]''
* ''[[Cleisostoma]]''
* ''[[Clowesia]]''
* ''[[Coelogyne]]''
* ''[[Coryanthes]]''
* ''[[Cycnoches]]''
* ''[[Cymbidium]]''
* ''[[Cyrtopodium]]''
* ''[[Cypripedium]]''
* ''[[Dactylorhiza]]''
* ''[[Dendrobium]]''
* ''[[Disa (plant)|Disa]]''
* ''[[Dracula (plant)|Dracula]]''
* ''[[Encyclia]]''
* ''[[Epidendrum]]''
* ''[[Epipactis]]''
* ''[[Eria]]''
* ''[[Eulophia]]''
* ''[[Gastrochilus]]''
* ''[[Gongora]]''
* ''[[Goodyera]]''
* ''[[Grammatophyllum]]''
* ''[[Gymnadenia]]''
* ''[[Habenaria]]''
* ''[[Herschelia]]''
* ''[[Ionopsis]]''
* ''[[Laelia]]''
* ''[[Lepanthes]]''
* ''[[Liparis (plant)|Liparis]]''
* ''[[Ludisia]]''
* ''[[Lycaste]]''
* ''[[Masdevallia]]''
* ''[[Maxillaria]]''
* ''[[Meliorchis]]''
* ''[[Mexipedium]]''
* ''[[Miltonia]]''
* ''[[Mormodes]]''
* ''[[Odontoglossum]]''
* ''[[Oeceoclades]]''
* ''[[Oncidium]]''
* ''[[Ophrys]]''
* ''[[Orchis]]''
* ''[[Paphiopedilum]]''
* ''[[Papilionanthe]]''
* ''[[Paraphalaenopsis]]''
* ''[[Peristeria (plant)|Peristeria]]''
* ''[[Phaius]]''
* ''[[Phalaenopsis]]''
* ''[[Pholidota (plant)|Pholidota]]''
* ''[[Phragmipedium]]''
* ''[[Platanthera]]''
* ''[[Platystele]]''
* ''[[Pleione (plant)|Pleione]]''
* ''[[Pleurothallis]]''
* ''[[Pomatocalpa]]''
* ''[[Promenaea]]''
* ''[[Pterostylis]]''
* ''[[Renanthera]]''
* ''[[Restrepia]]''
* ''[[Restrepiella]]''
* ''[[Rhynchostylis]]''
* ''[[Roezliella]]''
* ''[[Saccolabium]]''
* ''[[Sarcochilus]]''
* ''[[Satyrium (plant)|Satyrium]]''
* ''[[Seidenfadenia]]''
* ''[[Selenipedium]]''
* ''[[Serapias]]''
* ''[[Sobralia]]''
* ''[[Spiranthes]]''
* ''[[Stanhopea]]''
* ''[[Stelis]]''
* ''[[Thrixspermum]]''
* ''[[Tolumnia (plant)|Tolumnia]]''
* ''[[Trias (plant)|Trias]]''
* ''[[Trichocentrum]]''
* ''[[Trichoglottis]]''
* ''[[Vanda]]''
* ''[[Vanilla (genus)|Vanilla]]''
* ''[[Yoania]]''
* ''[[Zeuxine]]''
* ''[[Zygopetalum]]''
{{div col end}}

=== Etymology ===

The [[type (biology)|type]] [[genus (biology)|genus]] (i.e. the genus after which the family is named) is ''[[Orchis]]''. The genus name comes from the [[Ancient Greek]] {{lang|grc|[[:wikt:ὄρχις|ὄρχις]]}} (''{{lang|grc|órkhis}}''), literally meaning "[[testicle]]", because of the shape of the twin tubers in some species of ''Orchis''.<ref>{{cite book |author=Joan Corominas |year=1980 |title=Breve Diccionario Etimológico de la Lengua Castellana |publisher=Ed. Gredos |isbn=978-84-249-1332-8 |page=[https://archive.org/details/brevediccionario00colo/page/328 328] |url-access=registration |url=https://archive.org/details/brevediccionario00colo/page/328 }}</ref><ref>{{Cite book |last1=Hyam |first1=R. |last2=Pankhurst |first2=R.J. |year=1995 |title=Plants and their names : a concise dictionary |location=Oxford |publisher=Oxford University Press |isbn=978-0-19-866189-4 |name-list-style=amp }}</ref><ref>{{cite web|url=https://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0057:entry=o)/rxis |title=ὄρχις |last1=Liddell |first1=Henry George|last2=Scott|first2=Robert |work=A Greek-English Lexicon |publisher=Perseus Digital Library |date= 1940 }}</ref> The term "orchid" was introduced in 1845 by [[John Lindley]] in ''School Botany'',<ref>[http://www.etymonline.com/index.php?term=orchid Online Etymology Dictionary, "orchid"].</ref> as a shortened form of ''Orchidaceae''.<ref>{{Cite book |last1=Grigson |first1=G. |year=1973 |title=A Dictionary of English Plant Names |location=London |publisher=Allen Lane |isbn=978-0-7139-0442-0 }}</ref>

In [[Middle English]], the name ''bollockwort'' was used for some orchids, based on "[[wikt:bollock|bollock]]" meaning testicle and "[[wikt:wort#Etymology 1|wort]]" meaning plant.<ref>{{Cite web|url=http://www.oed.com/view/Entry/21112|title=bollock, n. and adj.|website=[[Oxford English Dictionary]]|access-date=2018-01-19|url-access=subscription }}</ref>

=== Hybrids ===
Orchid species hybridize readily in cultivation, leading to a large number of hybrids with complex naming.  Hybridization is possible across genera, and therefore many cultivated orchids are placed into [[Hybrid name|nothogenera]].  For instance, the nothogenus [[× Brassocattleya|× ''Brassocattleya'']] is used for all hybrids of species from the genera ''[[Brassavola]]'' and ''[[Cattleya]]''.  Nothogenera based on at least three genera may have names based on a person's name with the suffix ''[[wikt:-ara|-ara]]'', for instance [[× Colmanara|× ''Colmanara'']] = ''[[Miltonia]]'' × ''[[Odontoglossum]]'' × ''[[Oncidium]]''.  (The suffix is obligatory starting at four genera.<ref>[[International Code of Nomenclature for Cultivated Plants]] 9th edition (2016), Article H.6 and H.7.</ref>)

Cultivated hybrids in the orchid family are also special in that they are named by using [[Grex (horticulture)|grex]] nomenclature, rather than nothospecies.  For instance, hybrids between ''[[Brassavola nodosa]]'' and ''[[Brassavola acaulis]]'' are placed in the grex ''Brassavola'' Guiseppi.<ref>[https://bluenanta.com/common/information/100023580/?family=Orchidaceae&role=pri Brassavola Guiseppi Casa Luna 1968] {{Webarchive|url=https://web.archive.org/web/20230524163603/https://bluenanta.com/common/information/100023580/?family=Orchidaceae&role=pri |date=24 May 2023 }}, BlueNanta.</ref> The name of the grex ("Guiseppi" in this example) is written in a non-italic font without quotes.<ref>[[International Code of Nomenclature for Cultivated Plants]] 9th edition, 2016.</ref>

=== Abbreviations ===
As a unique feature of the orchid family, a system of abbreviations exists that applies to names of genera and nothogenera.  The system is maintained by the [[Royal Horticultural Society]].<ref>{{cite web| url = https://www.rhs.org.uk/plants/pdfs/plant-registration-forms/orchid-name-abbreviations-list.pdf| title = Alphabetical List of Standard Abbreviations for Natural and Hybrid Generic Names}}</ref>  These abbreviations consist of at least one character, but may be longer.  As opposed to the usual one-letter abbreviations used for names of genera, orchid abbreviations uniquely determine the (notho)genus.  They are widely used in cultivation.  Examples are ''Phal'' for ''[[Phalaenopsis]]'', ''V'' for ''[[Vanda]]'' and ''Cleis'' for ''[[Cleisostoma]]''.