Handelwurzen

Handelwurzen
	Dense-flowered Händelwurz ( Gymnadenia densiflora )
Systematics
Order :	Asparagales (Asparagales)
Family :	Orchids (orchidaceae)
Subfamily :	Orchidoideae
Tribe :	Orchideae
Sub tribus :	Orchidinae
Genre :	Handelwurzen
Scientific name
	Gymnadenia
	R.Br.

The gymnadenia ( Gymnadenia ) are a genus of flowering plants in the family of orchids (Orchidaceae). The genus includes, including the species of the earlier genus Nigritella (cabbage florets), about 26 species. The genus Gymnadenia s. l. is Palearctic , from the Iberian Peninsula in the west to China, Korea and Japan in the east.

There is a scientific controversy within botany about the inclusion of the cabbage florets (known as the genus Nigritella ) in the genus Gymnadenia . While some researchers believe that both should be united in the genus Gymnadenia (as illustrated in this article), others argue that both are sister groups and could be maintained as separate genera.

description

Illustration from Nordens Flora: A. Mücken-Haendelwurz ( Gymnadenia conopsea ) and B. Fragrant Haendelwurz ( Gymnadenia odoratissima )

Illustration from images of the basic forms of orchid species occurring in Germany and the neighboring areas , panel 40 of the black cabbage rose ( Gymnadenia nigra )

Illustration from Album des orchidées de l'Europe centrale et septentrionale, 1899 of the fragrant Handelwurz ( Gymnadenia odoratissima )

Inflorescence of the white form of the gnat ( Gymnadenia conopsea ) with zygomorphic flowers in detail, the spur is clearly visible

Vegetative characteristics

Gymnadenia species grow terrestrially as perennial herbaceous plants of medium height. The relatively thick, fleshy tubers are divided in the shape of a hand, each with some slender, somewhat fleshy roots . The independently upright stem is slim to strong, stem -round, fleshy and glabrous.

The leaves are arranged alternately on the stem. The simple leaf blades are linear-ribbon-shaped to elliptical and the longitudinally folded blade base envelops the stem. Leaf sheaths are often present at the base of the stem.

Generative characteristics

Many flowers stand together in terminal, head-shaped to elongated, racemose inflorescences . There are few lanceolate, sterile bracts . The fertile bracts are shorter to slightly longer than the ovary.

The seated under constant ovary is usually turned slightly and thus are the flowers resupinat (in the species of the former genus Nigritella not rotated). The bald ovary is narrow-cylindrical or spindle-shaped. The relatively small to medium-sized, hermaphrodite flowers are zygomorphic and threefold. The cylindrical spur is often slightly curved and shorter or longer than the ovary, with a blunt or bilobed end. The bracts are pink to dark purple or rarely white. The central, concave sepal and the straight, lateral petals often curve together like a helmet; whereby the two petals are slightly shorter than the sepal. The two free, lateral sepals are mostly spread out and bent back. The lip ( labellum ) is broad-rhombic to obovate and simple to clearly three-lobed. The column is short. The upright anthers are elongated or egg-shaped and have two counters . There are two club-shaped pollinia . The relatively small to elongated rostellum (adhesive gland) is indistinctly three-lobed. The two relatively small staminodes are almost spherical and are located laterally at the base of the fertile anthers. The two very large scars are on the side of the base on each side of the column.

The capsule fruits are upright.

ecology

In Gymnadenia TYPES is tuber geophytes . There is an endotrophic orchid-type mycorrhiza .

Attractants and pollinators

In two species ( Mosquito-Haendelwurz and Fragrant Haendelwurz ) gas chromatography identified 45 and 44 volatile aromatic substances that attracted butterflies , of which 37 and 25 species could be identified. Benzyl acetate , eugenol and benzoic acid benzyl ester were recognized as attractants in the case of the mosquito-handelwort, benzyl acetate, benzaldehyde , phenylethanal , 1-phenyl-2,3-butanedione, phenylethyl acetate, eugenol and an unidentified component acted as attractants in the case of fragrant Handelwurz. The smell of the mosquito-handelwurz, spread in the distribution area of the fragrant hendelwurz, could not attract pollinators for the mosquito-handelwurz. In a day-night rhythm , the examined flower types gave off different attracting aromas.

Archduke Johann Kohlröschen ( Gymnadenia archiducis joannis )

Austrian cabbage rose ( Gymnadenia austriaca )

Cornelia's cabbage rose ( Gymnadenia corneliana )

Gabas cabbage florets ( Gymnadenia gabasiana )

Gymnadenia orchidis

Common cabbage rose ( Gymnadenia rhellicani )

Aries cabbage rose ( Gymnadenia widderi )

Systematics and distribution

The genus Gymnadenia was in 1813 by Robert Brown in Hortus Kewensis; or, a Catalog of the Plants Cultivated in the Royal Botanic Garden at Kew. London. , 2nd edition, Volume 5, pages 191, 197 prepared. Type species is Gymnadenia conopsea (L.) R.Br. The generic name Gymnadenia is derived from the Greek gymnos: naked and aden: gland (ie “naked gland”), it refers to the often exposed adhesive disc (Viscidium) in the flowers.

The genus Gymnadenia belongs to subtribe Orchidinae from the tribe Orchideae in the family of Orchidaceae . For a while it was assumed that the genus Gymnadenia was only closely related to the genus Nigritella (cabbage florets). But now the latter is mostly included in the former.

The genus Gymnadenia s. l. is Palearctic , widespread in the west from the Iberian Peninsula across Europe, West and Central Asia to China (five species, three of which are only there), Korea and Japan in the east. Some species are endemic and few species are widespread.

The genus Gymnadenia includes, including the species of the earlier genus Nigritella Rich. , about 26 species:

Archduke Johann Kohlröschen ( Gymnadenia archiducis-joannis (Teppner & E. Klein) Teppner & E. Klein , Syn .: Nigritella archiducis-joannis Teppner & E. Klein ): It only occurs in the northeastern Limestone Alps in the Dead Mountains , Dachstein Mountains ( State of Salzburg), in Hochobir (Karawanken, Carinthia), in the Styrian Koralpe and in the Triglav National Park in Slovenia.
Austrian cabbage rose ( Gymnadenia austriaca (Teppner & E. Klein) P.Delforge , Syn .: Nigritella nigra subsp. Austriaca Teppner & E. Klein ): There have been two varieties since 1998. It occurs from Central Europe to the Pyrenees and the Carpathians .
Two-colored cabbage florets ( Gymnadenia bicolor (W.Foelsche) W.Foelsche & O.Gerbaud , Syn .: Nigritella bicolor W.Foelsche ): It occurs in the Alps and in the southern Carpathians .

Gymnadenia bicornis Tang & KYLang : It occurs in eastern and south-eastern Tibet .

Gymnadenia borealis (Druce) RMBateman, Pridgeon & MWChase (Syn .: Gymnadenia conopsea subsp. Borealis (Druce) F.Rose ): It occurs in Central and Northern Europe . This rare species appears only in a few isolated locations in the United Kingdom and Germany.

Buschmann's cabbage florets ( Gymnadenia buschmanniae (Teppner & Ster) Teppner & E. Klein , Syn .: Nigritella buschmanniae Teppner & Ster , Nigritella rubra subsp. Buschmanniae (Teppner & Ster) H. Baumann & R.Lorenz ): This endemic only occurs in the northeast Italy before.

Carpathian cabbage rose ( Gymnadenia carpatica (Zapal.) Teppner & E. Klein , Syn .: Nigritella nigra subsp. Carpatica (Zapal.) H.Baumann & R.Lorenz ): This endemic occurs only in the eastern Carpathians.

Mosquito Händelwurz ( Gymnadenia conopsea (L.) R.Br. , Syn .: Gymnadenia vernalis Dworschak ): It is widespread from Europe to Western Asia and Russia to China and Japan.

Cornelias Kohlröschen ( Gymnadenia corneliana (Beauverd) Teppner & E. Klein , Syn .: Nigritella nigra subsp. Corneliana Beauverd , Nigritella corneliana (Beauverd) Gölz & HRReinhard ): This endemic occurs only in the French and Italian western Alps .

Gymnadenia crassinervis Finet : It occurs from Tibet to western Sichuan and northwestern Yunnan .

Gymnadenia densiflora (Wahlenb.) A.Dietr. : It is common in Europe.

Dolomite cabbage florets ( Gymnadenia dolomitensis Teppner & E. Klein , syn .: Nigritella rubra subsp. Dolomitensis (Teppner & E. Klein) Wenker & S.Wenker ): This endemic occurs only in the southeastern Alps .

Gymnadenia emeiensis K.Y.Lang : It only thrives at an altitude of around 3,100 meters in the southwestern part of the Chinese province of Sichuan .

Balkan Haendelwurz or Frivalds Haendelwurz ( Gymnadenia frivaldii Hampe ex Griseb. , Syn .: Pseudorchis frivaldii (Hampe ex Griseb.) PFHunt ): It comes from the Balkan Peninsula from Montenegro and Bulgaria to northern Greece and occurs in the southern Carpathians.

Gabas cabbage rose ( Gymnadenia gabasiana (Teppner & E. Klein) Teppner & E. Klein , Syn .: Nigritella gabasiana Teppner & E. Klein ): It occurs from the French Pyrenees to northern Spain .

Gymnadenia hygrophila (W.Foelsche & Heidtke) W.Foelsche, Heidtke & O.Gerbaud (Syn .: Nigritella hygrophila W.Foelsche & Heidtke ): This endemic occurs only in the southeastern Alps.

Steineralpen cabbage florets ( Gymnadenia lithopolitanica (Ravnik) Teppner & E. Klein , Syn .: Nigritella lithopolitanica Ravnik , Nigritella nigra subsp. Lithopolitanica (Ravnik) H.Baumann & R.Lorenz ): This endemic occurs only in the Southeastern in the Koralpe as well in Karawanken in Austria and Kamnik-Savinja in northern Slovenia.

Red cabbage rose ( Gymnadenia miniata (Crantz) Hayek , Syn .: Nigritella miniata (Crantz) Janch. , Nigritella rubra (Wettst.) K. Richt. ): It occurs in the Central Alps, the Southeast Alps and the southern Carpathians.

Black cabbage rose ( Gymnadenia nigra (L.) Rchb. F. , Syn .: Nigritella nigra (L.) Rchb.f. , Satyrium nigrum L. , Orchis nigra (L.) Scop. , Habenaria nigra (L.) R. Br. , Sieberia nigra (L.) Spreng. , Nigritella angustifolia Rich. , Nigritella fragrans Saut. Rchb ex. , orchis atropurpurea exchange , orchis reichenbachii Mutel , Nigritella suaveolens W.DJKoch , Nigritella brachystachya A.Kern. , Nigritella megastachya A. Kern. , Nigritella hybrida Schur , Orchis moritziana Brügger )

Fragrant Händelwurz ( Gymnadenia odoratissima (L.) Rich. ) It occurs in the Limestone Alps , in northern Spain , in central Germany , in southern Sweden , in the Baltic States , in Belarus and in the Ukraine .

Gymnadenia orchidis Lindl. : It iswidespreadin India in the Himalayas from Arunachal Pradesh via Sikkim to Kashmir , in Nepal and Pakistan, in the eastern and southern Tibetan autonomous region and in the Chinese provinces of southeastern Gansu , western Hubei , southern Qinghai , southern Shaanxi , western Sichuan , northwestern Yunnan .

Common cabbage florets ( Gymnadenia rhellicani (Teppner & E. Klein) Teppner & E. Klein , Syn .: Nigritella rhellicani Teppner & E. Klein , Nigritella nigra subsp. Rhellicani (Teppner & E. Klein) H. Baumann, Künkele & R. Lorenz ): It occurs in the Alps (France, Germany, Austria, Switzerland, Italy), in the Appennines and in the Carpathians, in Bosnia and Herzegovina and in northern Greece.

Bride's cabbage rose ( Gymnadenia runei (Teppner & E. Klein) Ericsson , Syn .: Nigritella runei (Teppner & E. Klein) Kreutz ): It occurs only in Sweden .

Styrian cabbage rose ( Gymnadenia stiriaca (Rech.) Teppner & E. Klein , Syn .: Nigritella rubra subsp. Stiriaca (Rech.) H.Baumann & R.Lorenz ): This endemic occurs only in the Salzkammergut , Dachstein region and in the Graz mountainous region.

Gymnadenia taquetii Schltr. : This endemic occurs only in the Korean province of Jeju-do .

Spring Händelwurz ( Gymnadenia vernalis Dworschak ): It is a synonym of Gymnadenia conopsea (L.) R.Br. . This endemic occurs only in Upper Bavaria.

Ram's cabbage rose ( Gymnadenia widderi (Teppner & E. Klein ) Teppner & E. Klein , Syn .: Nigritella widderi Teppner & E. Klein , Nigritella rubra subsp. Widderi (Teppner & E. Klein ) H.Baumann & R.Lorenz ): It occurs in the Limestone Alps in the Eastern Alps in Bavaria as well as in the Austrian state of Styria and in the Italian Abruzzo .

Gymnadenia × heufleri

There are also a large number of hybrids:

Gymnadenia × abelii (Asch. & Graebn.) JMHShaw = Gymnadenia odoratissima × Gymnadenia rubra
Gymnadenia × borisii Stoj., Stef. & T.Georgiev = Gymnadenia frivaldii × Gymnadenia nigra
Gymnadenia × chanousiana G.Foelsche & W.Foelsche = Gymnadenia conopsea × Gymnadenia rhellicani
Gymnadenia × delphineae (M.Gerbaud & O.Gerbaud) M.Gerbaud & O.Gerbaud = Gymnadenia corneliana × Gymnadenia rhellicani
Gymnadenia × eggeriana O. Gerbaud = Gymnadenia austriaca var. Gallica × Gymnadenia rhellicani
Gymnadenia × fohringeri (Griebl) JMHShaw = Gymnadenia lithopolitanica × Gymnadenia miniata
Gymnadenia × geigelsteiniana ( B. Baumann & H.Baumann) JMHShaw = Gymnadenia odoratissima × Gymnadenia widderi

Gymnadenia × godferyana (G.Keller) W.Foelsche = Gymnadenia conopsea × Gymnadenia rubra

Gymnadenia × hedrenii (W. Foelsche) JMHShaw = Gymnadenia conopsea × Gymnadenia hygrophila

Gymnadenia × heufleri (A.Kern.) Wettst. = Gymnadenia nigra × Gymnadenia odoratissima

Gymnadenia × hubertii (Griebl) JMHShaw = Gymnadenia bicolor × Gymnadenia conopse

Gymnadenia × intermedia Peterm. = Gymnadenia conopsea × Gymnadenia odoratissima

Gymnadenia × kaeseri (Braun-Blanq.) Oddone = Gymnadenia conopsea × Gymndenia nigra × Gymnadenia rhellican

Gymnadenia × moritziana (Bruges ex Nyman) Oddone = Gymnadenia conopsea × Gymnadenia nigra

Gymnadenia × petzenensis (F.Fohringer & Redl) Oddone = Gymnadenia lithopolitanica × Gymnadenia rhellicani

Gymnadenia × pyrenaeensis W. Foelsche = Gymnadenia conopsea × Gymnadenia gabasiana

Gymnadenia × heavy (G.Keller) JMHShaw = Gymnadenia conopsea × Gymnadenia nigra × Gymnadenia odoratissima

Gymnadenia × teppneri (W. Foelsche) Oddone = Gymnadenia hygrophila × Gymnadenia rhellicani

Gymnadenia × trummeriana (W. Foelsche) JMHShaw = Gymnadenia odoratissima × Gymnadenia stiriaca

Gymnadenia × truongiae (Demares) W. Foelsche = Gymnadenia conopsea × Gymnadenia corneliana

Gymnadenia × turnowskyi (W.Foelsche) W.Foelsche = Gymnadenia conopsea × Gymnadenia lithopolitanica

Gymnadenia × wettsteiniana O. Abel = Gymnadenia nigra × Gymnadenia rubra

Individual evidence

↑ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t ^u ^v ^w ^x ^y ^z ^aa ^ab ^ac ^ad Rafaël Govaerts (Ed.): Gymnadenia. In: World Checklist of Selected Plant Families (WCSP) - The Board of Trustees of the Royal Botanic Gardens, Kew . Retrieved October 13, 2018.
↑ Richard M. Bateman, Alexander RM Murphy, Peter M. Hollingsworth, Michelle L. Hart, Ian Denholm, Paula J. Rudall (2018): Molecular and morphological phylogenetics of the digitate-tubered clade within subtribe Orchidinae ss (Orchidaceae: Orchideae) . Kew Bulletin 73: 54 doi: 10.1007 / S12225-018-9782-1
↑ Marie K. Brandrud, Ovidiu Paun, Richard Lorenz, Juliane Baar, Mikael Hedrén (2019): Restriction-site associated DNA sequencing supports a sister group relationship of Nigritella and Gymnadenia (Orchidaceae). Molecular Phylogenetics and Evolution 136: 21-28. doi: 10.1016 / j.ympev.2019.03.018
↑ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k Xinqi Chen, Stephan W. Gale, Phillip J. Cribb: Gymnadenia R. Brown in WT Aiton. , Pp. 133–135 - the same text online as the printed work , In: Wu Zheng-yi, Peter H. Raven, Deyuan Hong (eds.): Flora of China. Volume 25: Orchidaceae. Science Press and Missouri Botanical Garden Press, Beijing and St. Louis 2009, ISBN 978-1-930723-90-0 .
↑ ^a ^b ^c ^d Franz K. Huber, Roman Kaiser, Willi Sauter, Florian P. Schiestl: Floral scent emission and pollinator attraction in two species of Gymnadenia (Orchidaceae). In: Oecologia , Volume 142, 2005, pp. 564-575, doi: 10.1007 / s00442-004-1750-9 ( PDF ).
^ Gymnadenia at Tropicos.org. Missouri Botanical Garden, St. Louis, accessed October 13, 2018.
↑ Herwig Teppner, Erich Klein: Etiam atque etiam - Nigritella versus Gymnadenia: new combinations and Gymnadenia dolomitensis spec, nova (Orchidaceae-Orchideae). In: Phyton , Vol. 38, No. 1, 1998, pp. 220-224.
↑ Alec M. Pridgeon: Genera Orchidacearum: Orchidoideae (part 1). Oxford University Press, 2001, ISBN 978-0-19-850710-9 (77th genus Gymnadenia on page 294).
↑ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ Gymnadenia in the IUCN Red List of Threatened Species 2018. Accessed on 2018-10-13.
^ Gymnadenia at Tropicos.org. In: Flora of Pakistan . Missouri Botanical Garden, St. Louis

Web links

Commons : Handelwurzen ( Gymnadenia ) - album with pictures, videos and audio files

Wikispecies: Haendelwurzen ( Gymnadenia ) - species directory

Gymnadenia at Tropicos.org. In: IPCN Chromosome Reports . Missouri Botanical Garden, St. Louis
Data sheet with the species occurring in Italy at Schede di Botanica .
Thomas Meyer: Data sheet genus: Haendelwurz (Gymnadenia) with identification key and photos at Flora-de: Flora von Deutschland (old name of the website: Flowers in Swabia ).