Orchids ( Dactylorhiza )

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Orchids
Dactylorhiza incarnata subsp. coccinea

Dactylorhiza incarnata subsp. coccinea

Systematics
Order : Asparagales (Asparagales)
Family : Orchids (orchidaceae)
Subfamily : Orchidoideae
Tribe : Orchideae
Sub tribus : Orchidinae
Genre : Orchids
Scientific name
Dactylorhiza
Neck. ex Nevski

The orchids ( Dactylorhiza ), also finger Wurzen or cuckoo flower called, form a comprehensive plant genus in the family of orchids (Orchidaceae). The 40 or so species are widespread in large parts of Europe, in the far north of Africa, in northern and central Asia; one species reaches western North America.

The orchids of the genus Dactylorhiza were not differentiated from the orchids of the genus Orchis until the middle of the 20th century . With regard to the delimitation from the genus Coeloglossum , systematic questions remain unanswered, as do the distinction between individual species. The use of the tubers as salep poses problems for species protection ; the use of individual species as ornamental plants is based on horticultural propagation.

description

Illustration of the spotted orchid ( Dactylorhiza maculata )

Vegetative characteristics

Orchid species are perennial herbaceous plants . As persistence organs, they form underground tubers that are 10 to 20 centimeters long and are composed of a 0.5 to 1.5 centimeter short shoot and the adjacent root tissue. The side of the tuber is compressed, in some species deeply to five-part fingered with relatively thin, long sections, in other species less subdivided and more like a rounded, spindle-shaped tuber. Since there are often several steles , these tubers could have arisen from several roots that have grown together. Every year a new tuber is formed that lasts through the winter while the old one dies. The roots emerge on the shoot above the tuber . Only in the Crimean orchid ( Dactylorhiza iberica ) are there subterranean runners .

The upright, bare stem axis becomes hollow as the stem grows. The stem axis is constructed in cross-section as follows: under the cuticle and epidermis is followed by a three to six cell layers thick chlorenchyma with large intercellular spaces, then a three to five cell layers thick sclerenchyma that the vascular bundles comprises, finally in the middle of a parenchyma which dissolves, which forms the cavity in the stem. At the base of the stem axis are some lower leaves arranged in two rows , three of which contain a renewal bud, the top of which sprouts.

The leaves are in a loose rosette or are distributed in the lower area of ​​the shoot. They are pure green or green with reddish spots. The tip of the leaf often ends fused into a small cap. The leaf base includes the stem axis. The leaves of many species are broadly oval, but species that are widespread in high altitudes and northern areas have narrow leaves that are folded along the midrib or curved into a sickle shape. Another adaptation of the foliage leaves, which occurs in arid regions and high altitudes, is a reduced number of leaves with a rosette-like accumulation at the base of the stem axis. The leaf margin has a toothed structure with a folded cuticle that is only visible under the microscope. The stomata are located on the underside of the leaf and on the stem axis ; these have no secondary cells.

  • Tuber, spotted orchid ( Dactylorhiza maculata )

  • Large, reddish-spotted leaves at the base, spotted orchid

  • Small leaves scattered on the stem, spotted orchid

  • Immaculate leaf, flesh-colored orchid ( Dactylorhiza incarnata )

Inflorescences and flowers

Dactylorhiza incarnata nothosubsp. versicolor

Above the leaves, the stem axis continues as a racemose inflorescence . The flowers are usually close together. The bracts are leaf-like and often longer than the flower. The sessile ovary is cylindrical to spindle-shaped and twisted, so that the flowers are resupinated .

The hermaphrodite flowers are zygomorphic and threefold. The colors of the bracts are pink, purple, yellow or rarely white. The upward-pointing sepal and the side petals adhere to each other and form a hood over the flower. The lateral sepals are spread out to the rear. The lip is clearly three-lobed to almost entire, at the base with a spur that does not contain any nectar. The lip is often provided with darker stripes or spots on a light background. The column is short, with two barely visible staminodes on the side , the fertile stamen contains two club-shaped pollinia . The pollinia hang over a small stalk (caudicel) on a sticky gland (viscidium), which is surrounded by a thin cover (bursicula). The pollinia are made up of several parts ( massulae ), each of which consists of many pollen grains glued together and reach a length of 140 to 300 µm with a width of 80 to 200 µm. The individual pollen grains have a diameter of 10 to 20 µm, they are firmly glued together, the outer layer (exine) of the pollen grains is only formed on the outside of the massulae, it is missing inside between the glued pollen grains. The separating tissue between the stamen and stigma (rostellum) extends between the two pollinia.

Fruits and seeds

The capsule fruit contains 2000 to 5000 seeds. The capsule fruits are spindle-shaped, 350 to 1100 µm long with a diameter of 150 to 300 µm. The seed coat is compared to the genus Orchis of many small cells. These butt against each other without gaps, they have species-specific patterns on the surface. The seeds retain their ability to germinate for several years, which is a relatively long time for an orchid.

  • Inflorescence of Dactylorhiza majalis subsp. baltica

  • Zygomorphic flowers of the spotted orchid ( Dactylorhiza maculata )

  • Infructescence of the spotted orchid

  • Two color morphs of elder orchid ( Dactylorhiza sambucina )

Protocorm

The seeds only germinate with the help of mushrooms, from which they get nutrients. A proto shape is formed that is elongated and conical in shape, somewhat like a turnip.

ingredients

The leaves of some examined species contain various flavonoids . When spotted orchid ( Dactylorhiza fuchsii ) were kaempferol and quercetin found in the plant probably as glycosides are present. When flesh-colored orchid ( Dactylorhiza incarnata ) are in the leaves flavone glycosides present, the Elder-flowered Orchid ( Dactylorhiza sambucina ) are known various quercetin glucosides as Isoquercitrin .

The flowers contain anthocyanins . These are available as cyanidine glucosides, cyanine and seranine as diglucosides and other dyes - orchicyanine I and II, ophrysanine and serapianine, which are named after orchids - which contain oxalic acid via an acyl group .

cytology

The basic chromosome number in Dactylorhiza is x = 20. There is mostly diploidy with a chromosome number 2n = 40. There are several polyploid species with 2n = 4x = 80 and 2n = 6x = 120 chromosomes. There are hybrids between diploid and tetraploid individuals (they are usually sterile) and then have a chromosome number of 2n = 60, a number that also occurs in the island orchid ( Dactylorhiza insularis ). In some populations, such as the family of the elder orchid ( Dactylorhiza sambucina ) and Dactylorhiza aristata , plants with a chromosome set of 2n = 42 occur regularly. In the Roman orchid ( Dactylorhiza romana ) specimens with an aneuploid chromosome set of 2n = 40 + 1B were found. Other irregular chromosome numbers, which only occur rarely, are 2n = 100, 120, 122 ( Dactylorhiza maculata , Dactylorhiza russowii ) and 2n = 41, 78 ( Dactylorhiza maculata, Dactylorhiza fuchsii , Dactylorhiza umbrosa ). The chromosomes are generally small and are similar in this aspect of the boy herbs of the genus orchis , but possess a number from 2n = 42nd Metacentric chromosomes of roughly equal size are typical of Dactylorhiza .

Life cycle

Life course

The seeds of Dactylorhiza usually germinate in the autumn of the year of their distribution, for further differentiation they need a cold period. Young protocorms that live mycotrophically have been found in several species in summer. The first root forms in autumn, and the first shoot with leaves can be formed in the next spring. Then the plant goes to sympodial growth, the rhizome becomes free of fungi, which are then limited to the roots. The time until the first leaves appear differs for different species: this can happen in the first growing season or it can take several years. A flower does not yet appear in the young plants (as is the case with the short-lived green hollow tongue ( Coeloglossum viride )), but only after further vegetation periods, which are spent purely vegetatively. The times from germination to the first flowering range from two to 16 years. Flowering plants usually bloom in the following growing season again and for several years enduring . A population consists to a large extent (the figures range from 50 to 85%) of adult, flowering plants. Especially in unfavorable environmental conditions, fully grown plants can fall back into the purely vegetative stage and stop flower formation. The so-called secondary dormancy also rarely comes into play, in which adult plants only survive for one or more years with their subterranean organs. Many species are quite long-lived and also reproduce vegetatively to a limited extent, but overall the populations depend on regular reproduction by seeds for their maintenance.

Seasonal rhythm

The leaves emerge from the perennial tuber in spring, and in the case of Mediterranean species even in autumn. Simultaneously with the above-ground development of the leaf rosette, a new tuber, rarely two, is planted underground, which is filled with reserve substances during the growing season. It blooms in spring or early summer, followed by the ripening of the seeds. While in the south the growth activity ends with the onset of the summer drought, further north the seed ripeness shifts into autumn. The old shoot, including the tuber from which it sprouted, die at the end of the growing season. The unfavorable season (dry summer in the Mediterranean area, cold winter further north) is survived by means of the newly formed tuber.

ecology

These geophytes hibernate with the help of a hand-shaped tuber. The colored flowers are visited by insects, but contain no nectar .

Mycorrhiza

In the protocorm stage, the plant feeds mycotrophically . Also later, in plants that have leaves, fungi can be found in the roots and occasionally in the ends of the finger-shaped divided tubers. Fungi of the genus Ceratobasidium and Thanetephorus orchidicola , Thanatephorus cucumeris and Tulasnella calospora were isolated from the roots of adult plants . Under laboratory conditions, the seeds of the orchids will germinate with some of the fungi isolated from adult plants. However, the germination rates and the further development of the Protokorm are very different for different fungal strains.

pollination

The flowers are wide open and, with the spreading lip, offer a landing place for insects. There are usually lines or points on the lip that indicate the wide opening of the spur. The sides of the scar often have a darker stripe pointing towards the spur. The lip is also drawn in high contrast in ultraviolet light. Fine papillae that become denser towards the spur could serve as tactile guidelines for the insects. Deviating from these general properties, the flower of the green hollow tongue is built: It contains nectar that is located in the spur and at the base of the lip, on both sides of the entrance to the spur. The entrance to the spur is narrowed by two side slats. The flowers of the green hollow tongue smell slightly of honey.

If an insect visits the flower and looks for nectar in the spur, it touches the cover (bursicula) around the adhesive discs with its head or thorax , breaks this cover at a predetermined breaking point, pushes it backwards and exposes the sticky viscidia. These stick to the insect, which pulls the pollinia out of the stamen with further movements. The size of the flower and the size of the mainly pollinating insects must match; only then are the pollinaries glued to the center of the insect. Pollinaries sticking to the side can be removed more easily from the insect by cleaning. 20 to 40 seconds after being pulled out of the stamen, the stalks of the pollinaria dry up and bend forward. Only then do the pollinia come into a position in which they can touch the scar. The insect has usually already left the flower by this time, and geitonogamy is prevented. When the flowers are visited again, the pollinia stick to the stigma and some parts (massulae) of the pollinia stick there. The bond strength of scars liquid must be greater than the cohesion of the Massulae (by elastic viscin filaments) with one another. Since only parts of the pollinium remain on the stigma, an insect can pollinate several flowers.

The orchids are often pollinated by bumblebee queens ( bombus ), who are still inexperienced and who look for nectar in the flowers. The variable drawing and coloring of the flowers within a species could help to prevent the insects from learning quickly. The pollinators are not species-specific, so that there are frequent hybrids. In Fuchs's orchid ( Dactylorhiza fuchsii ) and the spotted orchid ( Dactylorhiza maculata ) longhorn beetles (Cerambycidae) have also been observed as pollinators, they eat papillae on the lip. Honey bees ingest sugar-containing stigma from the flowers of Fuchs's orchid. Without a visit from the insects, the flowers do not set any fruit, so they are dependent on cross- pollination .

distribution

Area

Distribution area of ​​the genus Dactylorhiza

The Dactylorhiza species are mainly found in Europe in the temperate areas , as well as around the Mediterranean, including North Africa and Asia Minor . Dactylorhiza hatagirea as the most south-eastern species has a distribution area on the southern slope of the Himalayas, Dactylorhiza aristata reaches the Aleutian Islands via eastern Russia , Madeira orchid ( Dactylorhiza foliosa ) marks the south-western limit of distribution, while Iceland is reached in the northwest .

Dactylorhiza species grow from sea level to altitudes of 2500 meters.

Averyanov suspects ancestors of the genus Dactylorhiza in the Poltava flora from the Paleogene . Due to the warmer climate, these plants would not have had any underground tubers. The storage organs could then have developed during the uplift of the Alps as an adaptation to the climate of the high elevations. Towards the end of the Pliocene , when the climate in Europe was generally cooler, the ancestors of Dactylorhiza settled far lower altitudes. The summer-dry Mediterranean region could also be reached in this way, made possible by the ability to withstand cooler and drier conditions acquired in the high altitudes. During the frequently changing climatic conditions of the ice ages , the areas of individual species fluctuated. Populations were separated, allopatric speciation occurred . During favorable periods, the genus was able to extend its distribution across Siberia to the Pacific; since then, the increasingly drier climate in the interior of Asia has created gaps in the area. As refuges during the Ice Ages, Greece, the Iberian Peninsula and North Africa come into question. Central Europe was settled from there after the Ice Ages, so the genetic variability of the northern plants is only a part of those found in the Mediterranean area.

If one measures the diversity on the basis of the number of described species, then northwest Europe forms a center of biodiversity. For example, in an area that includes the British Isles and the south of Scandinavia and stretches south to the Alps, there are numerous widespread species on the one hand, and clans with a narrowly delimited area on the other. The endemic clans present here are of recent origin: arose on the one hand through the variable glaciation during the Ice Age, on the other hand through hybridization. The more widespread species mainly radiate to the east, while the Eastern European and Western Siberian species mostly originate in northwestern Europe. The connections to the south to the Mediterranean area are less pronounced, there are mostly different species than in northwestern Europe.

A high biodiversity can also be found in the Carpathian Mountains and the Balkans. There are a number of species that are also found in northwestern Europe, but also a number of endemic clans. A third center of diversity is in Asia Minor: The species that occur here have little relation to the European Dactylorhiza species, they mostly occur exclusively in Asia Minor and the Caucasus. The endemics of these two areas are of older origin; the species have not changed or expanded their areas as much as some European ones.

In Central Asia there are two more regions that are home to several Dactylorhiza species: the Hindu Kush and the Pamir . In addition to the widespread species with a European center, there are other endemic species.

However, the distribution of genetic diversity differs from the distribution of the number of species. The eastern Mediterranean, the Caucasus and the Crimea have the greatest genetic diversity, although not that many species have been described in this area. In north-western Europe, where numerous species are known, there is little diversity at the genome level ; in this area many closely related clans and hybrids have been described as distinct species.

Locations

The locations are mostly in open, sunny situations or in partial shade. These can be different habitats such as dry grass , dunes , meadows , swamps, moors , light shrubbery and light forests. While some species such as Dactylorhiza maculata and Dactylorhiza fuchsii were found in both moist and dry locations, other species are tied to specific biotopes . The demands of the individual species also differ with regard to the pH of the soil .

Endangerment and species protection

A decline can be observed in many populations of the orchid species. The reasons for this are the draining of swamps and moors as well as an intensification of agriculture with fertilization of meadows and pastures. Changed use of pastureland, which leads to bushes and shading, can also lead to a decline in orchids. In Asia, the collection of the tubers is a threat, especially Dactylorhiza hatagirea from the Himalayas is particularly threatened. Only a few species, such as Fuchs's orchid ( Dactylorhiza fuchsii ) and the overlooked orchid ( Dactylorhiza praetermissa ), colonize disturbed locations and can stay there for a certain time.

Systematics

Herbarium of Dactylorhiza sambucina

Botanical history

The genus Dactylorhiza was established by Noël Martin Joseph de Necker in Sergei Arsenjewitsch Newski .

The orchids of the genus Dactylorhiza were not distinguished from those of the genus Orchis until late . Although the current scientific name was coined by Necker in 1790 , it was forgotten and is not considered a valid first description (Necker's Elementa botanica is listed in the ICBN as an opus utique opressum , a work that cannot be used ). Even the work of Nevski , who published the first description in the Flora of the USSR in 1937, still received no response. So in 1947 Vermeulen used the new name Dactylorchis in his dissertation Studies on Dactylorchids , which was a fundamental work for the study of the genus . The separation from the genus Orchis was thus widely recognized, the name Dactylorhiza found widespread use in the following years, for example in the work Problems of the genus Dactylorhiza published by Karlheinz Senghas in 1968 .

The botanical genus name Dactylorhiza is derived from the Greek  δάκτυλος dactylos for finger and ρίζα rhiza for root and describes the tubers, which are compressed and multi-part hand-shaped. The most common German name "Knabenkraut" originated when the current genera Orchis and Dactylorhiza were not yet differentiated. It is still used for the species of the genus Dactylorhiza . The name "cuckoo flowers", another old name for the orchid, was proposed by Werner Rothmaler in 1958 as the name for the genus Dactylorhiza . Occasionally, to distinguish it from the genus Orchis, the use of the name "Fingerwurzen" is recommended, which is the literal translation of the botanical name.

External system

The genus Dactylorhiza belongs to the tribe Orchideae in the subfamily Orchidoideae within the family Orchidaceae . The closest relationship exists to the genus of the Händelwurzen ( Gymnadenia ) and to the green hollow tongue ( Coeloglossum viride ). Most of the related genera also occur mainly in Europe, with the exception of the clade of forest hyacinths ( Platanthera ) and Galearis , which have centers of distribution in East Asia and North America. The relationships can be represented with a cladogram as follows:




Orchids ( Dactylorhiza ) + Green Hollow Tongue ( Coeloglossum viride )


   

Handelwurzen ( Gymnadenia )



   


Höswurzen ( Pseudorchis )


   

Forest hyacinths ( Platanthera )


   

Galearis




   

Sphere Orchis ( Traunsteinera )


   

Dwarf stendel ( Chamorchis alpina )





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Relationship to the green hollow tongue

The green hollow tongue, for which a separate genus, Coeloglossum , was established is particularly closely related to Dactylorhiza . Initial studies of ribosomal DNA placed the green hollow tongue within the genus Dactylorhiza . It was consequently renamed from Coeloglossum viride to Dactylorhiza viridis . The generic name Dactylorhiza , which is more recent than Coeloglossum , has been proposed as a nomen conservandum . An examination of the DNA of the chloroplasts did not reveal any resolution of the closely related species. The result of a more recent study, which included further DNA segments, sees the green hollow tongue as a sister taxon to the genus Dactylorhiza and still as an independent genus.

Coeloglossum viride as a sister group to Dactylorhiza Coeloglossum viride within Dactylorhiza



Dactylorhiza incarnata - group


   

remaining Dactylorhiza



   

Coeloglossum viride



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Dactylorhiza incarnata - group


   

Coeloglossum viride


   

remaining Dactylorhiza




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Internal system

In the genus Dactylorhiza there are diploid and tetraploid plants, whereby the assignment of the diploid to individual species is not as controversial as the assessment of the tetraploid clans. The duplication of the chromosome set can be present as autopolyploidy, which means that only one initial species was involved in the formation of the tetraploid species, or as allopolyploidy, in which the tetraploid plants were created by hybridizing two diploid parent species. From the same combination of two parent species, hybrids repeatedly emerged at different times and in different places, which can differ outwardly and in their ecological demands, but show few genetic differences. The delineation of the tetraploid orchid species in particular is difficult, as they show a large range of variation that mediates between the original species. In addition to old, stabilized hybrid flocks, there are often spontaneous hybrids that are sometimes more numerous than the parent species. The deciphering of the relationships by molecular genetic methods is made more difficult by the repeated hybridization and by the small genetic differences within the genus.

Leonid Averyanov presented a division of the genre into four sections. Two of them contained only one species each, the Crimean orchid ( Dactylorhiza iberica ) in the section Iberanthus and Dactylorhiza aristata in the section Aristatae . The Sambucinae section was also poor in species, the majority of the species were in the Dactylorhiza section , which was subdivided even more finely by subsections. Molecular genetic studies could largely not support the subdivision proposed by Averyanov. In addition to the actually relatively isolated three species-poor sections of Averyanov ( Dactylorhiza iberica , Dactylorhiza aristata as well as Dactylorhiza romana and Dactylorhiza sambucina ) there are three other large groups: two mainly diploid groups around Dactylorhiza incarnata and Dactapylorhiza maculata between these two groups of the tetrloid. In the representation of a classical cladogram, the relationships cannot be represented by hybridization, so the two following only contain a selection of diploid species:

Pillon et al. 2006 Pillon et al. 2007








Dactylorhiza foliosa


   

Dactylorhiza maculata



   

Dactylorhiza fuchsii



   

Dactylorhiza saccifera



   

Dactylorhiza aristata



   

Dactylorhiza romana


   

Dactylorhiza sambucina


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Dactylorhiza iberica



   

Coeloglossum viride


   

Dactylorhiza incarnata


   

Dactylorhiza euxina





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Dactylorhiza foliosa


   

Dactylorhiza maculata



   

Dactylorhiza fuchsii



   

Dactylorhiza saccifera



   

Dactylorhiza sambucina



   

Dactylorhiza romana



   

Dactylorhiza aristata



   

Coeloglossum viride



   

Dactylorhiza incarnata


   

Dactylorhiza euxina



   

Dactylorhiza iberica


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Dactylorhiza alpestris
Dactylorhiza aristata
Dactylorhiza cordigera subsp. pindica
Dactylorhiza elata
Dactylorhiza foliosa
Dactylorhiza hatagirea
Dactylorhiza insularis
Dactylorhiza lapponica
Dactylorhiza maculata
Dactylorhiza majalis
Dactylorhiza romana
Dactylorhiza russowii
Dactylorhiza saccifera
Dactylorhiza traunsteineri
Dactylorhiza viridis
Dactylorhiza × venusta

List of Dactylorhiza species and their distribution

Alphabetical list of Dactylorhiza species:

  • Dactylorhiza alpestris (Pugsley) Aver. (Syn .: Dactylorhiza majalis subsp. Alpestris (Pugsley) Senghas ): It occurs in European mountains: in the Alps , the Pyrenees and the Carpathians .
  • Dactylorhiza aristata (Fisch. Ex Lindl.) Soó : It occurs from eastern China to southern Alaska .
  • Dactylorhiza atlantica Kreutz & Vlaciha : This endemic occurs only in the High Atlas of Morocco.
  • Dactylorhiza baumanniana J.Hölzinger & Künkele (incl. Dactylorhiza smolikana B.Willing & E.Willing ): It occurs from Bulgaria to northern Greece .
  • Dactylorhiza cordigera (Fr.) Soó : There are four subspecies:
    • Dactylorhiza cordigera subsp. bosniaca (Beck) Soó : It occurs on the northern Balkan Peninsula .
    • Dactylorhiza cordigera (Fr.) Soó subsp. cordigera (Syn .: Dactylorhiza cordigera var. graeca (H.Baumann) Presser ): It occurs from Southeastern Europe to the Ukraine .
    • Dactylorhiza cordigera subsp. pindica (B.Willing & E.Willing) H.Baumann & R.Lorenz : It occurs from Romania to northwestern Greece .
    • Dactylorhiza cordigera var. Rhodopeia Presser : It occurs in northern Greece.
  • Dactylorhiza cyrnea W. Foelsche & Cord-Landwehr : It wasfirst describedfrom Corsica in 2012.
  • Dactylorhiza czerniakowskae Aver. : It occurs in Central Asia .
  • Dactylorhiza durandii (Boiss & Reut..) M.Laínz (Syn .: Dactylorhiza maculata subsp. Maurusia (Emb & Maire) Soó. , Dactylorhiza maurusia (Emb & Maire) Holub. ): It comes in Morocco and northern Algeria before.
  • Dactylorhiza elata (Poir.) Soó : It occurs from Western Europe to the western Mediterranean . There are two subspecies:
    • Dactylorhiza elata (Poir.) Soó subsp. elata (Syn .: Dactylorhiza elata subsp. mauritanica B. Baumann & H.Baumann ): It occurs in southwestern Spain and in northwestern Africa .
    • Dactylorhiza elata subsp. sesquipedalis (Willd.) Soó : It occurs in western and south- western Europe .
  • Dactylorhiza euxina (Nevski) Czerep. : It occurs from northeastern Turkey to the Caucasus . There are two subspecies:
    • Dactylorhiza euxina subsp. armeniaca (Hedrén) Kreutz : It occurs in eastern Turkey.
    • Dactylorhiza euxina (Nevski) Czerep. subsp. euxina : It occurs from north-eastern Turkey to the Caucasus .
  • Dactylorhiza foliosa (Rchb.f.) Soó : It occurs in Madeira .
  • Dactylorhiza fuchsii (Druce) Soó : There are six subspecies:
    • Dactylorhiza fuchsii subsp. carpatica (Batoušek & Kreutz) Kreutz : It only occurs in the Carpathian Mountains .
    • Dactylorhiza fuchsii (Druce) Soó subsp. fuchsii : It is widespread from Europe to Mongolia .
    • Dactylorhiza fuchsii subsp. hebridensis (Wilmott) Soó : It occurs only in Great Britain and in western Ireland .
    • Dactylorhiza fuchsii subsp. okellyi (Druce) Soó : It occurs in western Great Britain and western Ireland.
    • Dactylorhiza fuchsii subsp. psychrophila (Schltr.) Holub (Syn .: Dactylorhiza maculata var. psychrophila (Schltr.) Vöth , Dactylorhiza psychrophila (Schltr.) Aver. ): It is widespread from Europe to Siberia .
    • Dactylorhiza fuchsii subsp. sooana (Borsos) Borsos : It occurs only in the Carpathian Mountains.
  • Dactylorhiza graggeriana (Soó) Soó : It occurs in the western Himalayas .
  • Dactylorhiza hatagirea (D.Don) Soó : It occurs from Mongolia to the Himalayas.
  • Dactylorhiza iberica (M.Bieb. Ex Willd.) Soó : It occurs from Greece to Iran .
  • Dactylorhiza incarnata (L.) Soó : It occurs from Europe to north-western China. There are eight subspecies:
    • Dactylorhiza incarnata subsp. coccinea (Pugsley) Soó : It occurs in Great Britain and Ireland.
    • Dactylorhiza incarnata subsp. cruenta ( Blood-Red Orchid ) (OFMüll.) PDSell : It is widespread from Europe to north-western China.
    • Dactylorhiza incarnata subsp. gemmana (Pugsley) PDSell : It occurs in Western Europe .
    • Dactylorhiza incarnata (L.) Soó subsp. incarnata : It is widespread from Europe to Central Asia.
    • Dactylorhiza incarnata subsp. jugicrucis Akhalk., R. Lorenz & Mosul. : It occurs in Transcaucasia .
    • Dactylorhiza incarnata subsp. lobelii (Verm.) HAPedersen : It occurs in Norway , Denmark and the Netherlands .
    • Dactylorhiza incarnata subsp. ochroleuca ( straw-yellow orchid ) (Wüstnei ex Boll) PFHunt & Summerh. : It occurs in Europe.
    • Dactylorhiza incarnata subsp. pulchella (Druce) Soó : It occurs in Europe.
  • Dactylorhiza insularis (Sommier) Ó.Sánchez & Herrero : It occurs in Morocco , Portugal , Spain , France , Sardinia , Corsica and Italy .
  • Dactylorhiza isculana Seiser : This endemic wasfirst describedfrom Austria in 2002.
  • Dactylorhiza kafiriana Renz : There are two subspecies:
    • Dactylorhiza kafiriana subsp. baumgartneriana (B. Baumann & al.) Sczep. & Kreutz : It only occurs in Azerbaijan .
    • Dactylorhiza kafiriana Renz subsp. kafiriana : It occurs in Afghanistan , Pakistan and in the western Himalayas.
  • Dactylorhiza kalopissii E. Nelson : There are three subspecies:
    • Dactylorhiza kalopissii E. Nelson subsp. kalopissii : It occurs from Bulgaria to northern Greece.
    • Dactylorhiza kalopissii subsp. macedonica (J.Hölzinger & Künkele) Kreutz : It occurs in northern Greece and in northern and central Romania .
    • Dactylorhiza kalopissii subsp. pythagorae (Gölz & HRReinhard) Kreutz (Syn .: Dactylorhiza pythagorae Gölz & HRReinhard ): It occurs only on islands in the eastern Aegean .
  • Dactylorhiza kerryensis (Wilmott) PFHunt & Summerh. (Syn .: Dactylorhiza majalis subsp. Occidentalis (Pugsley) PDSell ): It occurs only in southern and western Ireland and in northern Great Britain.
  • Dactylorhiza kulikalonica Chernyak. : It occurs in Central Asia .
  • Dactylorhiza lapponica (Laest. Ex Hartm.) Soó : There are two subspecies.
    • Dactylorhiza lapponica subsp. angustata (Arv.-Touv.) Kreutz : It occurs only in France .
    • Dactylorhiza lapponica (Laest. Ex Hartm.) Soó subsp. lapponica : It occurs in Northern Europe and in Central Europe.
  • Dactylorhiza maculata (L.) Soó : There are nine subspecies:
    • Dactylorhiza maculata subsp. battandieri (Raynaud) H.Baumann & Künkele : This endemic occurs only in northern Algeria.
    • Dactylorhiza maculata subsp. caramulensis Verm. : It occurs only in Portugal, Spain and France.
    • Dactylorhiza maculata subsp. elodes (Griseb.) Soó (Syn .: Dactylorhiza montellii (Verm.) P.Delforge ): It occurs in Europe.
    • Dactylorhiza maculata subsp. ericetorum (EFLinton) PFHunt & Summerh. : It occurs in Western Europe.
    • Dactylorhiza maculata subsp. islandica (Á.Löve & D.Löve) Soó : This endemic occurs only in Iceland .
    • Dactylorhiza maculata (L.) Soó subsp. maculata : It is widespread from Europe to Siberia .
    • Dactylorhiza maculata subsp. podesta (Landwehr) Kreutz : This endemic occurs only in the Netherlands .
    • Dactylorhiza maculata subsp. savogiensis (D.Tyteca & Gathoye) Kreutz : It occurs in the Pyrenees and in the Western Alps.
    • Dactylorhiza maculata subsp. transsilvanica (Schur) Soó : It occurs in eastern Central Europe and in Southeastern Europe.
  • Dactylorhiza magna (Czerniak.) Ikonn. (Syn .: Dactylorhiza baldshuanica Chernyak. ): It occurs in Central Asia.
  • Dactylorhiza majalis (Rchb.) PFHunt & Summerh. : There are three subspecies:
    • Dactylorhiza majalis subsp. baltica ( Baltic orchid ) (blade) H.Sund. (Syn .: Dactylorhiza longifolia (Neuman) Aver. ): It is distributed from Finland and Germany to Central Asia.
    • Dactylorhiza majalis (Rchb.) PFHunt & Summerh. subsp. majalis : It occurs in Europe.
    • Dactylorhiza majalis subsp. sphagnicola (Höppner) HAPedersen & Hedrén : It occurs in Norway , Sweden , Denmark , Belgium , the Netherlands , France and Germany .
  • Dactylorhiza nieschalkiorum H.Baumann & Künkele : It occurs only in northern Turkey .
  • Dactylorhiza osmanica (blade) PFHunt & Summerh. : It occurs from Turkey to Syria . There are two subspecies:
    • Dactylorhiza osmanica var. Anatolica (E. Nelson) Renz & Taubenheim : It occurs in Turkey.
    • Dactylorhiza osmanica (blade) PFHunt & Summerh. var. osmanica : It occurs from Turkey to Syria .
  • Dactylorhiza praetermissa (Druce) Soó : It occurs in western and northwestern Europe. There are the following subspecies and varieties:
    • Dactylorhiza praetermissa var. Junialis (Verm.) Senghas : It occurs in Western Europe.
    • Dactylorhiza praetermissa subsp. osiliensis (Pikner) Kreutz : It occurs only on the islands of the Baltic Sea .
    • Dactylorhiza praetermissa (Druce) Soó subsp. praetermissa : It occurs in western and north-western Europe.
    • Dactylorhiza praetermissa subsp. schoenophila R.M.Bateman & Denholm : This endemic was first described from Great Britain in 2012
  • Dactylorhiza purpurella (T.Stephenson & TAStephenson) Soó : It occurs in western and north-western Europe. There are two varieties:
    • Dactylorhiza purpurella var. Cambrensis (RHRoberts) RMBateman & Denholm (Syn .: Dactylorhiza majalis var. Cambrensis (RHRoberts) RMBateman & Denholm ): It occurs only on the coasts of Great Britain and Denmark .
    • Dactylorhiza purpurella (T.Stephenson & TAStephenson) Soó var. Purpurella : It occurs from northern Ireland to northern and north-western Great Britain, in southern Norway and on the island of Schiermonnikoog .
  • Dactylorhiza romana (Sebast.) Soó : There are three subspecies:
    • Dactylorhiza romana subsp. georgica (blade) Soó ex Renz & Taubenheim (Syn .: Dactylorhiza flavescens (K.Koch) Holub ): It occurs from Turkey to Central Asia .
    • Dactylorhiza romana subsp. guimaraesii (EGCamus) HAPedersen : It occurs in Morocco, Algeria, Portugal and Spain.
    • Dactylorhiza romana (Sebast.) Soó subsp. romana (Syn .: Dactylorhiza markusii (Tineo) H.Baumann & Künkele ): It occurs from Sicily to the eastern Mediterranean.
  • Dactylorhiza russowii (blade) Holub (Syn .: Dactylorhiza lapponica subsp. Russowii (blade) H.Baumann & R.Lorenz ): It occurs from Central Europe to Siberia.
  • Dactylorhiza saccifera (Brongn.) Soó : There are two subspecies:
    • Dactylorhiza saccifera subsp. gervasiana (death.) Kreutz : It occurs in Algeria and from Sicily to southern Italy.
    • Dactylorhiza saccifera (Brongn.) Soó subsp. saccifera (Syn .: Dactylorhiza urvilleana subsp. bithynica (H.Baumann) H.Baumann & R.Lorenz ): It occurs in the Mediterranean region.
  • Dactylorhiza salina (Turcz. Ex Lindl.) Soó : It occurs from the Caucasus to the Amur region.
  • Dactylorhiza sambucina (L.) Soó : It occurs in Europe.
  • Dactylorhiza sudetica (Poech ex Rchb. F.) Aver. : It is common from Europe to Siberia.
  • Dactylorhiza traunsteineri (Saut. Ex Rchb.) Soó : There are seven subspecies:
    • Dactylorhiza traunsteineri subsp. curvifolia (F.Nyl.) Soó : It occurs in Northern Europe and in the Czech Republic .
    • Dactylorhiza traunsteineri subsp. irenica (FMVázquez) Kreutz (Syn .: Dactylorhiza irenica F.M.Vázquez ): It occurs in Spain.
    • Dactylorhiza traunsteineri subsp. rhaetica (H.Baumann & R.Lorenz) F.Benoît (Syn .: Dactylorhiza lapponica subsp. rhaetica H.Baumann & R.Lorenz ): It occurs only in northern Italy.
    • Dactylorhiza traunsteineri subsp. schurii (blade) Kreutz (Syn .: Dactylorhiza maculata subsp. schurii (blade) Soó ): It occurs only in the southern and eastern Carpathians .
    • Dactylorhiza traunsteineri (Saut. Ex Rchb.) Soó subsp. traunsteineri : It occurs from northern Europe and central Europe to western Siberia.
    • Dactylorhiza traunsteineri subsp. turfosa (F.Proch.) Kreutz : It occurs in the Alps and in the western Carpathians .
    • Dactylorhiza traunsteineri subsp. vosagiaca Kreutz & P.Wolff : It occurs in Central Europe.
  • Dactylorhiza traunsteinerioides (Pugsley) Landwehr : It occurs only in Great Britain and Ireland .
  • Dactylorhiza umbrosa (Kar. & Kir.) Nevski : It iswidespreadfrom West Asia and Central Asia to Siberia.
  • Dactylorhiza urvilleana (Steud.) H.Baumann & Künkele : It occurs from northern and northeastern Turkey to Iran .
  • Dactylorhiza viridis (L.) RMBateman, Pridgeon & MWChase (Syn .: Coeloglossum viride (L.) Hartm. ): There are two varieties:
    • Dactylorhiza viridis var. Virescens (Muhl. Ex Willd.) Baumbach : It occurs from Russia's Far East to Japan and in North America.
    • Dactylorhiza viridis var. Viridis : It occurs in the subarctic and in the subalpine areas of the northern hemisphere.

List of natural Dactylorhiza hybrid species

Alphabetical list of Dactylorhiza - Nothospecies :

Superstition, use, conservation

In superstition, the bulbs dug up on St. John's Day ("St. John's hand") were considered to be good luck charms and sexual enhancers.

The tubers of the Orchis and Dactylorhiza species and some other orchids were used in paediatrics as a mucus drug ("Tubera Salep ") to treat irritation of the gastrointestinal tract, when there were no better medicines and remedies. Today, orchid species are no longer considered effective medicinal herbs.

The following important information is missing from this article or section:
There is no evidence for the following statement, here and in orchis .
Help Wikipedia by researching and pasting it .

All species of the genera Orchis and Dactylorhiza are under the strictest nature protection, especially the underground parts of the plant.

literature

  • Leonid V. Averyanov: A Review of the Genus Dactylorhiza . In: Orchid Biology. Reviews and Perspectives . tape 5 . Timber Press, Portland, Oregon 1990, ISBN 0-88192-170-X , pp. 159-206 .
  • Fritz Füller: The genera Orchis and Dactylorhiza. Orchids of Central Europe, 3rd part . In: The New Brehm Library . tape 286 . A. Ziemsen Verlag, 1983, ISSN  0138-1423 .
  • Alec M. Pridgeon, Phillip Cribb, Mark W. Chase, Finn Rasmussen (Eds.): Genera Orchidacearum. Orchidoideae (Part one) . Oxford University Press, New York / Oxford 2001, ISBN 0-19-850710-0 , pp. 279-284 .
  • Problems of the genus Dactylorhiza . In: Karlheinz Senghas, Hans Sundermann (Hrsg.): Annual reports of the natural science association in Wuppertal . tape 21/22 . Wuppertal 1968.

Individual evidence

  1. ↑ a b MG Vakhrameeva, IV Tatarenko, TI Varlygina, GK Torosyan, MN Zagulskii: Orchids of Russia and Adjacent Countries . ARG Gantner, Ruggell 2008, ISBN 978-3-906166-61-2 , p. 58 ff., 211 ff .
  2. ↑ a b c d e f g h i j k l Leonid V. Averyanov: A Review of the Genus Dactylorhiza . In: Orchid Biology. Reviews and Perspectives . tape 5 . Timber Press, Portland, Oregon 1990, ISBN 0-88192-170-X , pp. 159-206 .
  3. ↑ a b c d e f g h i j k l m n o p q r s Alec M. Pridgeon, Phillip Cribb, Mark W. Chase, Finn Rasmussen (eds.): Genera Orchidacearum. Orchidoideae (Part one) . Oxford University Press, New York / Oxford 2001, ISBN 0-19-850710-0 , pp. 279-284 .
  4. ↑ a b c Hanne N. Rasmussen: Terrestrial orchids from seed to mycotrophic plant . Cambridge University Press, Cambridge 1995, ISBN 0-521-04881-8 , pp. 295-303 .
  5. ^ Hanne N. Rasmussen: Terrestrial orchids from seed to mycotrophic plant . Cambridge University Press, Cambridge 1995, ISBN 0-521-04881-8 , pp. 121 .
  6. ^ A b c Jean Claessens, Jacques Kleynen: The flower of the European orchid. Form and function . Self-published, Geulle 2011, ISBN 978-90-90-25556-9 , pp. 232-249 (English).
  7. ↑ a b c d Yohan Pillon, Michael F. Fay, Alexey B. Shipunov, Mark W. Chase: Species diversity versus phylogenetic diversity: A practical study in the taxonomically difficult genus Dactylorhiza (Orchidaceae) . In: Biological Conservation . tape 129 , no. 1 , 2006, p. 4-13 , doi : 10.1016 / j.biocon.2005.06.036 .
  8. ↑ Fritz Füller: The genera Orchis and Dactylorhiza. Orchids of Central Europe, 3rd part . In: The New Brehm Library . tape 286 . A. Ziemsen Verlag, 1983, ISSN  0138-1423 .
  9. ↑ Nicolas Devos, Olivier Raspé, Anne-Laure Jacquemart, Daniel Tyteca: On the monophyly of Dactylorhiza Necker ex Nevski (Orchidaceae): is Coeloglossum viride (L.) Hartman a Dactylorhiza? In: Botanical Journal of the Linnean Society . tape 152 , no. 3 , 2006, p. 261-269 , doi : 10.1111 / j.1095-8339.2006.00561.x .
  10. ↑ a b c Yohan Pillon, Michael F. Fay, Mikael Hedrén, Richard M. Bateman, Dion S. Devey, Alexey B. Shipunov, Michelle van der Bank, Mark W. Chase: Evolution and temporal diversification of western European polyploid species complexes in Dactylorhiza (Orchidaceae) . In: Taxon . tape 56 , no. 4 , 2007, p. 1185-1208 , doi : 10.2307 / 25065911 .
  11. ^ Richard M. Bateman, Peter M. Hollingsworth, Jillian Preston, Luo Yi-Bo, Alec M. Pridgeon, Mark W. Chase: Molecular phylogenetics and evolution of Orchidinae and selected habenariinae (Orchidaceae) . In: Botanical Journal of the Linnean Society . tape 142 , no. 1 , 2003, p. 1-40 , doi : 10.1046 / j.1095-8339.2003.00157.x .
  12. ↑ 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 ae af ag ah ai aj ak al am an ao ap aq ar as at au av aw ax ay az ba bb bc bd be bf bg bh bi bj bk bl bm bn bo bp bq br bs bt bu bv bw bx by bz ca cb cc cd ce cf cg ch ci cj ck cl cm cn co cp cq cr cs ct cu cv cw cx cy cz da db dc Rafaël Govaerts (Ed.): Dactylorhiza. In: World Checklist of Selected Plant Families (WCSP) - The Board of Trustees of the Royal Botanic Gardens, Kew . Retrieved May 7, 2020.
  13. ↑ Orchid was considered a sexual enhancer on ksta.de, accessed on May 26, 2013.
  14. ↑ Schleimstoffe on exkotours.de, accessed on May 26, 2013.

See also

Web links

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