Gymnadenia

Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Monocots
Ordo: Asparagales

Familia: Orchidaceae
Subfamilia: Orchidoideae
Tribus: Orchideae
Subtribus: Orchidinae
Genus: Gymnadenia
Overview of species (26)

G. archiducis-joannis – G. austriaca – G. bicolor – G. bicornis – G. borealis – G. buschmanniae – G. carpatica – G. conopsea – G. corneliana – G. crassinervis – G. densiflora – G. dolomitensis – G. emeiensis – G. frivaldii – G. gabasiana – G. lithopolitanica – G. miniata – G. minor – G. nigra – G. odoratissima – G. orchidis – G. rhellicani – G. runei – G. stiriaca – G. taquetii – G. widderi

Nothospecies: (24)

G. × abelii – G. × borisii – G. × delphineae – G. × eggeriana – G. × fohringeri – G. × geigelsteiniana – G. × godferyana – G. × hedrenii – G. × heufleri – G. × hubertii – G. × intermedia – G. × jabornikana – G. × kaeseri – G. × petzenensis – G. × pyrenaeensis – G. × ritzbergeri – G. × schwerei – G. × suaveolens – G. × teppneri – G. × trummeriana – G. × truongiae – G. × turnowskyi – G. × wettsteiniana – G. × wucherpfennigiana
Name

Gymnadenia R.Br. in W.T.Aiton, Hortus Kew. 5: 191 (1813)

Type species: Gymnadenia conopsea (L.) R.Br. in W.T.Aiton, Hortus Kew. 5: 191 (1813)

Synonymy

Heterotypic
Nigritella Rich., De Orchid. Eur.: 26 (1817)
× Gymnigritella E.G.Camus, J. Bot. (Morot) 6: 484 (1892)

Distribution
Native distribution areas:

Europe
Northern Europe
Denmark, Finland, Great Britain, Ireland, Norway, Sweden.
Middle Europe
Austria, Belgium, Czechoslovakia, Germany, Hungary, Netherlands, Poland, Switzerland.
Southwestern Europe
Baleares, France, Portugal, Spain.
Southeastern Europe
Albania, Bulgaria, Greece, Italy, Romania, Sicilia, Yugoslavia.
Eastern Europe
Belarus, Baltic States, Krym, Central European Russia, East European Russia, North European Russia, South European Russia, Northwest European Russia, Ukraine.
Asia-Temperate
Siberia
Altay, Buryatiya, Chita, Irkutsk Krasnoyarsk, Tuva, West Siberia, Yakutiya.
Russian Far East
Amur, Khabarovsk, Kuril Islands, Primorye, Sakhalin.
Caucasus
North Caucasus, Transcaucasus.
Western Asia
Iran, Turkey.
China
China South-Central, Inner Mongolia, Manchuria, China North-Central, Qinghai, Tibet.
Mongolia
Mongolia.
Eastern Asia
Japan, Korea.
Asia-Tropical
Indian Subcontinent
East Himalaya, Nepal, Pakistan, West Himalaya.
Indo-China
Myanmar.

References: Brummitt, R.K. 2001. TDWG – World Geographical Scheme for Recording Plant Distributions, 2nd Edition
References
Primary references

Brown, R. 1813. Hortus Kewensis; or, a Catalogue of the Plants Cultivated in the Royal Botanic Garden at Kew. London (2nd ed.) 5: 191, 197.
Pridgeon, A.M., Cribb, P.J., Chase, M.W. & Rasmussen, F.N. (eds.) 2001. Genera Orchidacearum Volume 2: Orchidoideae (Part one); page 294 ff., Oxford: Oxford University Press. ISBN 0-19-850710-0

Links

'eFloras 2009. Gymnadenia in Flora of China vol. 25 Orchidaceae --> pdf Gymnadenia
Global Biodiversity Information Facility. 2023. GBIF Backbone Taxonomy. Checklist dataset. Taxon: Gymnadenia. Accessed: 2023 Jul 13.
Govaerts, R. et al. 2023. Gymnadenia in Kew Science Plants of the World Online. The Board of Trustees of the Royal Botanic Gardens, Kew. Published online. Accessed: 2023 Jul 13. Reference page.
Hassler, M. 2023. Gymnadenia. World Plants: Synonymic Checklists of the Vascular Plants of the World In: Roskovh, Y., Abucay, L., Orrell, T., Nicolson, D., Bailly, N., Kirk, P., Bourgoin, T., DeWalt, R.E., Decock, W., De Wever, A., Nieukerken, E. van, Zarucchi, J. & Penev, L., eds. 2023. Species 2000 & ITIS Catalogue of Life. Published online. Accessed: 2023 Jul 13. Reference page.
Hassler, M. 2023. World Plants. Synonymic Checklist and Distribution of the World Flora. . Gymnadenia. Accessed: 13 Jul 2023.
International Plant Names Index. 2023. Gymnadenia. Published online. Accessed: 13 Jul 2023. Reference page.
Tropicos.org 2023. Gymnadenia. Missouri Botanical Garden. Published online. Accessed: 13 Jul 2023.

Vernacular names

azərbaycanca: Dəmirsəhləb
تۆرکجه: سهلاب
башҡортса: Кәкүк әшәлсәһе
беларуская: Ядрушка
čeština: Pětiprstka
dansk: Trådspore
Deutsch: Händelwurzen
español: Orquídea olorosa
suomi: Kirkiruohot
hornjoserbsce: Kukawka
magyar: Bibircsvirág
հայերեն: Լերկաբշտիկ
italiano: Manina
日本語: テガタチドリ属
қазақша: Көкекшөп
перем коми: Гимнадения
коми: Гимнадения
lietuvių: Plauretis
кырык мары: Гимнадени
Nederlands: Muggenorchis
polski: Gółka
русский: Кокушник
slovenčina: Päťprstnica
svenska: Brudsporrar
удмурт: Гимнадения
українська: Билинець
中文: 手參屬, 手参属
中文（简体）: 手参属

Gymnadenia is a genus of flowering plants in the orchid family (Orchidaceae) containing 22 terrestrial species. The former genus Nigritella is now included in Gymnadenia.

They can be found in damp meadows, fens and marshes, and on chalk or limestone, often in alpine regions of Europe and Asia from Portugal to Kamchatka, including China, Japan, Mongolia, Siberia, the Himalayas, Iran, Ukraine, Germany, Scandinavia, Great Britain, etc.[2][3][4] The fragrant orchid (Gymnadenia conopsea) has been introduced into the USA and is reportedly naturalized in Connecticut.[5]

These hardy terrestrial orchids are deciduous. They survive the winter through two deep-cut tubers. Long lanceolate green leaves grow at the bottom of the stem. There are some small leaves at the stop of the stem.

They flower during the summer. The inflorescence is a dense cylindrical spike between 5 and 30 cm long. It can consists of up to 150 small pleasant-smelling flowers. It is recently discovered that eugenol and isoeugenol, floral volatile scent compounds, are catalyzed by single type of enzyme in Gymnadenia species and gene encoding for this enzyme is first functionally characterized gene in this species so far.[6] Their color can vary from pale purple to pink and white. The lip is wide with three lobes. The marginal petals are horizontal. There is a long, thin, threadlike spur.

Several species were formerly classified under Nigritella. The nothogeneric name ×Gymnigritella was used for hybrids between these two groups.
Species
Gymnadenia × heufleri
Gymnadenia rhellicani (syn. Nigritella rhellicani)

As of August 2023, Plants of the World Online accepted the following species and hybrids.[1] ([N] = formerly placed in Nigritella, [GN] = formerly placed in ×Gymnigritella.)

Gymnadenia archiducis-joannis (Teppner & E.Klein) Teppner & E.Klein [N] – Austria
Gymnadenia austriaca (Teppner & E.Klein) P.Delforge [N] – C. Europe to Pyrenees
Gymnadenia bicolor (W.Foelsche) W.Foelsche & O.Gerbaud
Gymnadenia bicornis Tang & K.Y.Lang – Tibet
Gymnadenia borealis (Druce) R.M.Bateman, Pridgeon & M.W.Chase
Gymnadenia buschmanniae (Teppner & Ster) Teppner & E.Klein [N] – Italy
Gymnadenia carpatica (Zapal.) Teppner & E.Klein [N] – E. Carpathians
Gymnadenia conopsea (L.) R.Br. – fragrant orchid, Europe to Temp. E. Asia
Gymnadenia corneliana (Beauverd) Teppner & E.Klein – SW. Alps
Gymnadenia crassinervis Finet – China
Gymnadenia densiflora (Wahlenb.) A.Dietr.
Gymnadenia dolomitensis Teppner & E.Klein [N] – S. Alps
Gymnadenia emeiensis K.Y.Lang – China (Sichuan)
Gymnadenia frivaldii Hampe ex Griseb. – Balkan Pen. to S. Carpathians
Gymnadenia gabasiana (Teppner & E.Klein) Teppner & E.Klein [N] – Pyrenees
Gymnadenia lithopolitanica (Ravnik) Teppner & E.Klein [N] – SE. Alps
Gymnadenia miniata (Crantz) Hayek
Gymnadenia minor (W.Foelsche & Zernig) W.Foelsche, Zernig & O.Gerbaud
Gymnadenia nigra (L.) Rchb.f. [N] – Europe, Israel
Gymnadenia odoratissima (L.) Rich. – Europe
Gymnadenia orchidis Lindl. – Himalaya to China
Gymnadenia rhellicani (Teppner & E.Klein) Teppner & E.Klein [N] – W. Alps
Gymnadenia runei (Teppner & E.Klein) Ericsson – Sweden
Gymnadenia stiriaca (Rech.) Teppner & E.Klein
Gymnadenia taquetii Schltr. – Korea
Gymnadenia widderi (Teppner & E.Klein) Teppner & E.Klein [N] – NE. Alps, C. Italy

Hybrids

Gymnadenia × abelii (Asch. & Graebn.) J.M.H.Shaw
Gymnadenia × borisii Stoj., Stef. & T.Georgiev
Gymnadenia × chanousiana G.Foelsche & W.Foelsche (G. rhellicani × G. conopsea[7]) [GN] – France
Gymnadenia × delphineae (M.Gerbaud & O.Gerbaud) M.Gerbaud & O.Gerbaud (G. corneliana × G. rhellicani[8]) GN) – France
Gymnadenia × eggeriana O.Gerbaud (G. austriaca var. gallica × G. rhellicani[9]) [N] – France
Gymnadenia × fohringeri (Griebl) J.M.H.Shaw
Gymnadenia × geigelsteiniana (B.Baumann & H.Baumann) J.M.H.Shaw
Gymnadenia × godferyana (G.Keller) W.Foelsche (G. conopsea × G. rubra[10]) [GN] – Alps
Gymnadenia × hedrenii (W.Foelsche) J.M.H.Shaw
Gymnadenia × heufleri (A.Kern.) Wettst. – Alps
Gymnadenia × hubertii (Griebl) J.M.H.Shaw
Gymnadenia × intermedia Peterm. – Europe
Gymnadenia × kaeseri (Braun-Blanq.) Oddone
Gymnadenia × moritziana (Brügger ex Nyman) Oddone
Gymnadenia × petzenensis (F.Fohringer & Redl) Oddone
Gymnadenia × pyrenaeensis W.Foelsche [GN] – Pyrenees
Gymnadenia × ritzbergeri (Lachmair) Eccarius
Gymnadenia × schwerei (G.Keller) J.M.H.Shaw
Gymnadenia × suaveolens (Vill.) Rchb.f.
Gymnadenia × teppneri (W.Foelsche) Oddone
Gymnadenia × trummeriana (W.Foelsche) J.M.H.Shaw
Gymnadenia × truongiae (Demares) W.Foelsche (G. conopsea × G. corneliana[11]) [GN] – France
Gymnadenia × turnowskyi (W.Foelsche) W.Foelsche (G. conopsea × G. lithopolitanica[12]) [GN] – Austria
Gymnadenia × wettsteiniana O.Abel [N] – Alps
Gymnadenia × wucherpfennigiana (W.Foelsche) Eccarius