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Dactylorhiza Neck. ex Nevski

Family Orchidaceae published in Trudy Botanicheskogo Instituta Akademii Nauk S S S R. Ser. 1, Flora i Sistematika Vysshikh Rastenii. Moscow & Leningrad 4: 332. 1937. This genus name is correct Distribution: Heavily represented in Western and Central Europe. Subtropical to Arctic Eurasia and mountainous North Africa; one species (Dactylorhiza aristata) also occurs in Alaska and one (Dactylorhiza praetermissa) in Newfoundland. 1) Horticultural abbr. : D. or Dact. Closely related to: Gymnadenia sensu lato
2)

Looks like: species in Orchis, Stachys officinalis and Lupinus often mislead beginning orchid hunters. Common names: in two distinct groups: Marsh orchids & Spotted orchids Associated groups: Terrestrial orchids Associated groups: Terrestrial orchids

Introduction to the genus

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Dactylorhiza sambucina

Dactylorhiza is a fairly large genus of terrestrial orchids with a firm standard design, the shape and size of the lip is the most useful characteristic for identification. The name is derived from the Greek Dacty (finger) and Rhiza (root), referring to the lobed tubers. Pretty much all species are highly variability in the number of flowers per inflorescence, their pigmentation and base colour, and the pigmentation of the leaves. The discussion on what's a species in this genus is far from settled and there are bound to be revisions in the future. A number of species are polymorphic (D. incarnata, D. sambucina, D. romana) and these flower morphs often grow mixed in the same population. 3) Indeed, Dacts are a difficult genus to tackle when you have a go at identification - always look at a population from a statistical point of view and discard the outliers. Absence or density of spots on the leaves is the least useful parameter in identification. 4) In addition, many species readily hybridise and produce fertile offspring where populations collide. Identification of hybrids is virtually impossible without decent knowledge of the species growing in the vicinity. Flora of Great Britain and Ireland: Butomaceae-Orchidaceae by Peter D. Sell and Gina Murrell, p340 Perennial herbs with 2-3 root tubers, which are usually palmately and deeply 2- to 3-lobed, the lobes narrowed and prolonged distally. Leaves several, the lower sheathing the stem, the upper transitional to the bracts and not sheathing, though sometimes clasping. Emerging spike not enclosed in spathe-like leaves. Bracts always herbaceous. Outer lateral perianth segments more or less erect or spreading, not contributing t0 the helmet over the column. Inner perianth segments connivent and forming a helmet over the column. Labellum not lobed, or more or less 3-lobed. porrect or slightly decurved. Spur usually less than 10 mm, downpointed, mostly rather wide. Column erect. Anther adnate to the top of the column: pollinia 2, narrowed below to the caudicle and attached by a basal disc to one of the viscidia; pollen in packets of tetrads united by elastic threads. Stigma more or less 2-lobed, roofing the entrance to the spur. Rostellum 3-lobed, the middle lobe short, lamelliform. Capsule erect. Bursicle present. Molecular studies
9)10)11)12)13) 5)6)7)8)

support five groups within the genus. The use of these groups has become popular compared to the four sections and seven subsections which were supported in previous monographs.

Dactylorhiza incarnata group: D. euxina and D. ulbrosa (diploids) Dactylorhiza maculata group: D. fuchsii, D. saccifera, D. foliosa (diploids), and D. maculata (autotetraploid) Dactylorhiza majalis group: D. angustata, D. elata, D. brennensis, D. sphagnicola, D. traunsteineri, D. baltica, D. russowii, D. praetermissa, D. purpurella, D. majalis, and associated infraspecific taxa (allotetraploids) Dactylorhiza sambucina group: D. romana and D. flavescens (all diploid) primitive diploids group: D. aristata, D. viridis (= Coeloglossum viride), and D. iberica An alternative classification relies on chromosome numbers: all those with the same chromosome number are a species (the Marsh orchids) and split the others (Spotted orchids) into two species, each with a different chromosome count. The Marsh orchids are then split up based on the shape of the lip, maintaining ecological and geographical populations as subspecies or varieties 14) . It's been suggested that Western European Dactylorhiza allotetraploids have evolved by hybridization between four main diploid lineages. The allotetraploid species D. elata, D. brennensis, and D. sphagnicola have originated from the autotetraploid D. maculata together with the diploid D. incarnata, while D. majalis, D. traunsteineri, and D. angustata seem to have evolved by hybridization between the D. fuchsii s.str and D. incarnata lineages. Finally, the diploid D. saccifera lineage seems to have been involved together with the D. incarnata lineage in the formation of the allotetraploid D. praetermissa. 15) Dactylorhiza was separated from Orchis in 1935 by Nevski, they stand out because of palmate tubers, green bracts and different chromosome number 16) . The first hint was already given by Linnaeus in his Species Plantarum 17) when he divided the genus Orchis in three parts based on the morphology of the roots: Bulbis indivisis, Bulbis palmatis and Bulbis fasciculatis. Necker 18) raised these parts to genus level, the first became Orchis L. while the latter two were joined in Dactylorhiza. 19) Vermeulen was apparently unaware of Nevski's work and seperated the genus Dactylorchis from Orchis in 1947, the name is therefore not accepted but often encountered in literature. For a run-down on the history of Dact taxonomy we advise to read Dactylorhiza Nevski, the correct generic name of the Dactylorchids by P.F. Hunt and V.S. Summerhayes. Dacts are typical for boreal and temperate meadows, headlands and grasslands with a moist subsoil in Europe. They're often among the first orchids to be spotted in areas with weak (to significant) swamping or over the edges of lowland and transient bogs. 20) Many populations are in decline because of a change in the water level, due to drainage and land development. In East Europe a number of Dactylorhiza species are still harvested from the wild in large quantities to make sahlep.

General cultivation techniques

Dacts are mostly used as garden plants in full soil, but are sporadically grown in container culture. Hybrids are the preferred choice for full soil cultivation, they show more vigorous growth and commercially propagated plants have a much richer inflorescence. Secondly, Dactylorhiza is still dug up from the wild and offered for sale - even though most species are loosing foothold. When introducing them in the garden it's worth the effort of looking up the natural occurrence (of the parents in case of hybrids): although all species are considered hardy, the ones which grow in Southern regions and D. foliosa 21) are best compared to Serapias and the less hardy Ophrys in winter care. 22) The opposite is also true: northern species such as D. aristata won't appreciate heath in Southern gardens. The majority of species are summergreen, only a few species have winterfoliage (e.g. D. sambucina & D. romana). The tuber pushes out a shoot in early Spring from which the basal rosette emerges, adults subsequently grow a single impressive flower spike. Shortly after the leaves have emerged a new tuber will start to form below ground, this tuber swells until Autumn while the previous tuber decays. Most species flower in Spring to mid-Summer (e.g. D. incarnata, D. majalis) or Summer (D. fuchsii, D. maculata). After the peak of flowering the bottom flowers will have faded and the seedpods swell. In pure species this often happens when the top of the inflorescence is still developing. Commercial hybrids on the other hand normally have an inflorescence whose flowers are all open during peak of blooming, and which fades rather fast when the plant starts producing seeds. In late Summer the leaves start to turn yellow and fade, don't cut the leaves back as they are still pumping nutrients in the new tuber. In Autumn the leaves and roots will have died back and all that remains is the new tuber (often more than 1) which is the orchid's way to survive Winter. Dacts rely heavily on seasonal triggers to coordinate their growth phases. 23) Whether grown in full soil outdoor or in pot indoor you need to provide a minimum of 3 months of cold treatment to induce shoot formation and healthy flowering the following season. They all need seasonal triggers, e.g. vernalisation, to coordinate their growth phases. They flower best after a cold winter so make sure they are well exposed to winter weather. Fertilisers should be avoided for this genus, they are rather hungry plants but the nutrition must be provided by a rich organic soil. For most terrestrial orchids this kind of soil mix combined with constant humidity normally causes tuber rot. Dacts won't suffer as long as their subterranean parts are not sitting in a waterlogged location. Always mix in drainage material to evacuate excess water around the plants, this can require preparing their spot (e.g. heavy clay soils) by digging to the depth of a foot and mixing it with perlite, sand and/or grit. Potted plants are often lost because of tuber rot due to overwatering in the winter, when they're dormant. The other reason is that a plant in the garden and in the wild is normally sitting in a rather inorganic soil with an organic top-layer. In pot culture the while container is often filled with an equal distribution of organic material. In nature the tuber isn't in contact with the rot-inducing topsoil but is positioned just below it, the new roots sprout from the top of the tuber and radiate outwards horizontally, tapping into the rich top layer.

in the Garde n
Good vs. Bad flowering quality

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Spike should bloom completely before setting seed

Hybrid: D fuchsii x praetermissa

Pure species: D. fuchsii

A few exceptions aside Dacts are not that strict in soil pH as their habitat often makes you believe, they do well in slightly acidic or slightly lime soils. But what they do demand is a soil which isn't allowed to go dry. They grow in a soil which is wet/moist or constantly damp from Spring to end of Summer and damp during Winter. Keeping the soil damp can be achieved by mulchin or by establishing a bed of ground-hugging companion plants before introducing them in the garden. Stagnant water is best avoided - especially for younger plants - as it leads to rot, exceptions such as D. majalis and D. praetermissa can be planted near the edge of a pond. D. baltica, D. sphagnicola and D. ericetorum can take a very wet soil in full light as long as the water isn't stagnant. The soil is a mix of washed coarse sand (drains excess moisture), peat (moistureretentive) and leaf mold. D. fuchsii and D. praetermissa however hate acidic soil, add dolomite limestone or an equivalent instead of peat to establish an alkaline or neutral pH. Most species appreciate dappled and light shade, but a position in full sunlight will be rewarded if the soil is not allowed to go dry. The tuber gives a hint towards the soil dryness they can withstand. The finger-like roots on the tuber are longer in species that are known to survive in drier conditions and shorter in those that show no chance of survival. A long-fingered tuber will grow its roots upwards in a moist soil, while growing them straight down in a dry setting. These long tuber-roots should not be seen as a possibility to grow them in an arid environment, but as an ability to withstand a freak period of drought. The following list sums up successful applications of pure species in the garden. It's intended to give the reader an idea of what parentage to look for when shopping for Dact cultivars and hybrids in the garden. Note that most Dacts enjoy direct sunlight, but their location must provide a damp soil. This is why we take the opposite approach: list suitable spots in the garden and species which can be established. Moist open woodland, e.g. orchard: species which enjoy a humus-rich top-layer with dappled shade and soil which remains damp or moist all year round: D. foliosa, D. sambucina and D. romana. Paris quadrifolia is an ideal companion plant. Direct sunlight in the morning is often the key factor for successful blooming. Moist border: D. grandis, D. foliosa, D. fuchsii or D. elata grows with Primula japonica or as a companion plant for Peltoboykinia watanabei 24) . They enjoy sun in the morning but require a spot which is shaded from noon till the evening. D. elata can take dry Summers if trickle irrigation is provided. D. fuchsii can take some Summer dryness without Southern heat 25) and is the ideal plant to give it a try as pure species in the garden, it's an often used parent in hybrids. Full sun in moist soil, lawn with a moist subsoil and wet meadows with a rich soil: D. elata, D. fuchsii, D. pardalina, D. purpurella and D. incarnata. More and more hybrids are becoming available which are suited for the lawn. When grown in the lawn care must be taken from early Spring on to not defoliate the plants when cutting back the grass. Vigorous grasses are best cut back when the Dactylorhiza shoots start to appear to give them a healthy start. This becomes less of an issue after a few years when a dense clump starts to form. It's not uncommon to find seedlings in the garden a few years after they've been allowed to set seed. If you planted commercial hybrids it's quite possible that these seedlings will not resemble the parents at all when they are mature. Remove them if you find the quality of the inflorescence inferior. If you're not interested in seedlings and just want to restrict the plants to their designated place it's advised to deadhead the inflorescence after flowering. Simply cut the flowering stalk halfway between the lowest flower and soil level, next year's flowers will benefit from this as seed development normally requires a fair bit of energy.

in Pots
Dacts in pots are not grown as successful as those in the garden, four possible reasons: the soil in the container is mixed as would be for other plants, growers mix all the ingredients and then fill the pot. Since dacts require a good deal of organic components the tuber sits in a rich medium, contrary to the garden situation where the tuber is buried in a rather inorganic soil with an organic layer on top. When planting them in a pot, make two mixes: one without organic material which fills the pot up to a few centimeters from the edge. Plant the tuber in this medium and fill the rest of the container with a mix with organic material. soil humidity during dormancy is tricky to control in pots, it's easy to overwater them. The easiest trick is to place the pots in a shallow plastic container filled with damp soil. Watering this soil will keep the pots damp enough, without having to water the tuber directly. Dactylorhiza - like most terrestrial orchids - are dependant on mycorrhiza. Repotting might have an adverse affect, but this is speculation. inadequate vernalization: dormant tubers need to be have a period of 3 months cold (3-10C depending on hardiness) to induce healthy shoot growth and flowering. For container-bound plants you can use general purpose potting soil (e.g. John Innes n3) but add a healthy serving of sand and grit for drainage. The top few centimeters are the same soil which has a generous serving of humus rich compost (e.g. beach/birch leaf mold). You can repot every year, preferably during dormancy - this allows you to inspect the tuber. Some people report that repotting in early Spring (before the shoot appears) or after flowering (when the plant fades) is a reliable alternative. Plants in pots can be fed with 20% strength foliar feed, but this is normally only done to reduce the drain during seed development.

In vitro culture

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D. fuchsii rhizoids (top) and protocorms (bottom)

Seed germination in vitro is inhibited by the carapace, sterilise with 1% NaOCl for 15 minutes to break this barrier. Stratification has a good effect on the germination rate of a lot of species, chill the cultures for 3-4 months at 4C.
27)

26)

The stratification is not always beneficial, as was shown by Znaniecka and ojkowska 28)

Symbiotic
Dactylorhiza has a low specificity concerning their mycorrhizal partners and interacts with species of the Rhizoctonia group. Thanatephorus orchidicola - strains which have been isolated from adult roots. Swelling and germination can be expected after 2 weeks, protocorm development should have started by week 6.
29)

Seedlings have been established on cultures of Ceratobasidium, Tulasnella calospora and

Asymbiotic
Modified Malmgren: amino acids plus 20 ml pine-apple juice/liter medium.
30)

A dark cycle with (optional) chilling for 2 months at 4C improves germination rates. Dactylorhiza can often be sown onto an MS medium with a fifth (1/5) of the regular concentration. Jeast-extract [1g/L], sugar [10g/L], and activated carbon [0,5g/L] can also be added. A maintenance medium is formulated with more B-vitamins (jeast-extract [500mg] and banan pulp [50g/L]) and may contain slightly more nutrient-salts. 31) This medium is often referred to as the Wetsteyn modified MS and can be found in detail at the OSP [members.cox.net].

Hybridisation
Hybridisation within the genus appears to be limitless. Species with the same ploidy produce fertile offspring and crosses from parents with a different ploidy are not completely sterile even if they result in triploid hybrids 32) . Hybrids are very varied, even from selfed parentage. Intergeneric hybrids with the genera Gymnadenia, Orchis, Platanthera and Coeloglossum have been established. It's uncertain what the possibilities are, but some conclusions: Platanthera x Dactylorhiza (= X Dactylanthera P.F. Hunt & Summerh.) was first established as X Rhizanthera chevallieriana (E.G. Camus) So but this turned out to be an abnormal D. maculata 33) . The intergeneric hybrid has been created in culture and also spotted in the wild. Platanthera is very dominant in both size of the plant as in flowers, labellum is slightly enlarged. 34) 35) Coeloglossum x Dactylorhiza (= X Dactyloglossum) has flowers which inherit most of their pigment from the Dactylorhiza parentage, but with a green overlay or tinge. Habit, leaves and perianth shape is variably intermediate depending on parentage. 36) The flowers are tilted forward with a typical hood as in Coeloglossum, but they're more open and generally have a larger lip. Fertility unknown. Young plants grow faster than regular Dacts but will always remain smaller in the adult stage. Colour spectrum: mainly purples, some yellows, a few red

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D. traunsteineri

D. iberica D. maculata subsp. maurusia

D. maculata leaves

yellow + purple D. romana

D. incarnata subsp. coccinea

D. fuchsii f. alba

D. insularis

D. sambucina, dark yellow morph

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Ex-vitro propagation
Many species clump readily (e.g. D. elata, D. fuchsii) but it's best not to divide them too often, they make an impressive display in group and they don't appreciate uprooting. Wait till Autumn when they're dormant to dig up the shallow tubers and keep as much soil around the tubers as possible. Plants in pots can off course be repotted every year to harvest new tubers. Forced tuber multiplication as explained in Ophrys seems to work. Gently uproot the plant after the flowers have faded, when the seedpods are swelling. Twist off the new (smaller, pale looking) tuber(s). Most growers also deadhead the plants to minimize loss of energy. Plant them back carefully and in most cases they'll start to produce new tubers to replace the ones you removed. Those which you harvested can be planted as well. An overview of this procedure is shown at the SRGC Bulb Log Diary [srgc.org.uk]. Plants in the garden often seed out to a corner which is sheltered and constantly damp. You can increase your luck by sowing the seed around the parent plant immediately after harvesting it - preferably just before a few days of rain. Seedlings normally pop up in patches which aren't disturbed and have low growing plants covering the soil (cushion plants). Dacts growing in a bog or peat soil seem to be particularly good at seeding out next to the motherplant. The SRGC Bulb Log Diary has an interesting entry on Dact seedlings [srgc.org.uk] which reports that healthy tubers have a strange smell of wet horse which is not present in unhealthy and infected tubers.

Biosis Pests & Diseases

slugs and snails fungal attacks: isolate affected plants, don't treat your other plants with fungicide as it might have an adverse effect on its mycorrhizal partner.

Prints & Books

Terrestrial orchids: from seed to mycotrophic plant by Hanne N. Rasmussen Cambridge University Press ISBN-10: 0521451655 ISBN-13: 978-0521451659 Orchids of Britain & Europe Pierre Delforge, 1995. Collins Photo Guide. ISBN 0 00 2200244. Hardy orchids Phillip Cribb and Christopher Bailes, 1989. Timber Pr. ISBN 0 88192 147 5. Die Orchideen Mitteleuropas und der Alpen Presser Helmut, 1995. ecomed, ISBN 3 609 65600 X. Das Moor im eigenen Garten: Moorgrten anlegen,gestalten und pflegen Erich Maier, 2000. Parey, ISBN 3 8263 3301 2 Orchideen im Garten: Verwendung, Pflege und Vermehrung Gerd Kohls und Ulrich Khler, 1993.Parey, ISBN 3 489 636244. Orchideen fr den Garten: Europische und tropische Erdorchideen Alfons Brger, 1992. Ulmer, ISBN 3 8001 64876.

Commercial sources
Rareplants.co.uk [rareplants.co.uk] offers a diverse list MyOrchids.de [myorchids.de] in Germany Easy Orchids [easyorchids.eu] in Europe

External pages
Gnther Blaich maintains an extensive list of Dact. hybrids in literature: part 1 [guenther-blaich.de] & part 2 [guenther-blaich.de] Dacts in a garden [srgc.org.uk] at the Scottish Rock Garden Club

Correct species
Dactylorhiza abantiana H.Baumann & Knkele Dactylorhiza aldenii H.Baumann Dactylorhiza alpestris (Pugsley) Aver. Dactylorhiza altobracensis (Coste & Souli) So Dactylorhiza aristata (Fisch. ex Lindl.) So Dactylorhiza aschersoniana (Hausskn.) Borsos & So Dactylorhiza baicalica Aver. Dactylorhiza balabaniana H.Baumann Dactylorhiza baldshuanica Chernyak. Dactylorhiza baumanniana J.Hlzinger & Knkele Dactylorhiza bayburtiana H.Baumann Dactylorhiza beckeriana (Hppner) So Dactylorhiza bohemica Businsk Dactylorhiza boluiana H.Baumann Dactylorhiza bourdonii D.Tyteca & Gathoye Dactylorhiza braunii (Halcsy) Borsos & So Dactylorhiza breviceras Renz & Taubenheim Dactylorhiza carnea So Dactylorhiza claudiopolitana (Simonk.) Borsos & So Dactylorhiza cordigera (Fr.) So Dactylorhiza csatoi (So) So Dactylorhiza czerniakowskae Aver. Dactylorhiza daunia W.Rossi & al. Dactylorhiza delamainii (G.Keller ex T.Stephenson) So Dactylorhiza dinglensis (Wilmott) So Dactylorhiza dubreuilhii (G.Keller & Jeanj.) So Dactylorhiza dufftiana (M.Schulze) So Dactylorhiza dufftii (Hausskn.) Peitz Dactylorhiza elata (Poir.) So Dactylorhiza euxina (Nevski) Czerep. Dactylorhiza flixensis (Gsell) So Dactylorhiza foliosa (Rchb.f.) So Dactylorhiza formosa So Dactylorhiza fourkensis B.Willing & E.Willing Dactylorhiza fuchsii (Druce) So Dactylorhiza gabretana (Fuchs) So Dactylorhiza genevensis (Klinge ex Schulze) So Dactylorhiza graggeriana (So) So Dactylorhiza grandis (Druce) P.F.Hunt Dactylorhiza guillaumeae C.Bernard Dactylorhiza gustavssonii H.Baumann Dactylorhiza hallii (Druce) So Dactylorhiza hatagirea (D.Don) So Dactylorhiza hochreutinerana (K.Hellm.) So Dactylorhiza iberica (M.Bieb. ex Willd.) So Dactylorhiza incarnata (L.) So Dactylorhiza insignis (T.Stephenson & T.A.Stephenson) So Dactylorhiza insularis (Sommier) .Snchez & Herrero Dactylorhiza isculana Seiser Dactylorhiza ishorica Aver. Dactylorhiza jenensis (Brand) So Dactylorhiza jestrebiensis Businsk Dactylorhiza juennensis Perko Dactylorhiza kafiriana Renz Dactylorhiza kalopissii E.Nelson Dactylorhiza katarana H.Baumann Dactylorhiza kelleriana P.F.Hunt Dactylorhiza kerasovinensis B.Willing & E.Willing Dactylorhiza kerneriorum (So) So Dactylorhiza komiensis Aver. Dactylorhiza kopdagiana H.Baumann Dactylorhiza koutsourana B.Willing & E.Willing Dactylorhiza kulikalonica Chernyak. Dactylorhiza kuuskiae E.Breiner & R.Breiner Dactylorhiza lapponica (Laest. ex Hartm.) So Dactylorhiza latirella (P.M.Hall) So Dactylorhiza lehmannii (Klinge) So Dactylorhiza longifolia (Neuman) Aver. Dactylorhiza maculata (L.) So Dactylorhiza magna (Czerniak.) Ikonn. Dactylorhiza majalis (Rchb.) P.F.Hunt & Summerh. Dactylorhiza megapolitana (Bisse) So Dactylorhiza metsowonensis B.Baumann & H.Baumann Dactylorhiza mulignensis (Gsell) So Dactylorhiza nieschalkiorum H.Baumann & Knkele Dactylorhiza ornonensis (G.Keller & Jeanj.) So Dactylorhiza osiliensis Pikner Dactylorhiza osmanica (Klinge) P.F.Hunt & Summerh. Dactylorhiza paridaeniana Kreutz Dactylorhiza praetermissa (Druce) So Dactylorhiza prochazkana Businsk Dactylorhiza purpurella (T.Stephenson & T.A.Stephenson) So Dactylorhiza renzii H.Baumann & Knkele Dactylorhiza rizeana Renz & Taubenheim Dactylorhiza romana (Sebast.) So Dactylorhiza rombucina (Cif. & Giacom.) So Dactylorhiza ruppertii (M.Schulze) Borsos & So Dactylorhiza russowii (Klinge) Holub Dactylorhiza saccifera (Brongn.) So Dactylorhiza salictina B.Willing & E.Willing Dactylorhiza salina (Turcz. ex Lindl.) So Dactylorhiza sambucina (L.) So Dactylorhiza serbica (Fleischm.) So Dactylorhiza serreana H.Baumann Dactylorhiza sivasiana E.Baumann & Knkele ex Renz & Taubenheim Dactylorhiza sooi (Ruppert ex So) So Dactylorhiza souflikensis B.Willing & E.Willing Dactylorhiza stagni-novi D.Tyteca & Gathoye Dactylorhiza sudetica (Poech ex Rchb.f.) Aver. Dactylorhiza szaboiana (So) So Dactylorhiza transiens (Druce) So Dactylorhiza traunsteineri (Saut. ex Rchb.) So Dactylorhiza umbrosa (Kar. & Kir.) Nevski Dactylorhiza urvilleana (Steud.) H.Baumann & Knkele Dactylorhiza vallis-peenae Kmpel Dactylorhiza venusta (T.Stephenson & T.A.Stephenson) So Dactylorhiza vitosana H.Baumann Dactylorhiza vogtiana H.Baumann Dactylorhiza vorasica B.Willing & E.Willing Dactylorhiza weissenbachiana Perko Dactylorhiza wiefelspuetziana D.Tyteca

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Dactylorhiza godferyana (So) Peitz

Dactylorhiza (Orchidaceae) at The Culture Sheet - The Web's niche for plant cultivation and horticulture
Dactylorhiza nevskii H.Baumann & Knkele

Synonyms
The rather lengthy list of Dactylorhiza synonyms is maintained here
1)

Dactylorhiza (Orchidaceae) in European Russia: combined molecular and morphological analysis, Alexey B. Shipunov, Michael F. Fay, Yohan Pillo2, Richard M. Bateman and Mark W. Chase, American Journal of Botany. 2004;91:1419-1426. 2) , 5) Bateman R. M. A. M. Pridgeon M. W. Chase 1997 Phylogenetics of subtribe Orchidineae (Orchidoideae, Orchidaceae) based on nuclear ITS sequences. 2. Infrageneric relationships and reclassification to achieve monophyly of Orchis sensu stricto. Lindleyana 12: 113-141 3) An atlas of orchid pollination by Nelis A. Cingel, p167 4) , 22) The Orchid book by James Cullen, p80 6) Bateman R. M. I. Denholm 2003 The heath spotted-orchid (Dactylorhiza maculata (L.) So) in the British Isles: a cautionary case-study in delimiting infraspecific taxa and inferring their evolutionary relationships. Journal Europischer Orchideen 35: 3-36 7) Hedrn M. M. F. Fay M. W. Chase 2001 Amplified fragment length polymorphisms (AFLP) reveal details of polyploid evolution in Dactylorhiza (Orchidaceae). American Journal of Botany 88: 1868-1880 8) Hedrn M. 2002 Speciation patterns in the Dactylorhiza incarnata/maculata polyploid complex (Orchidaceae): evidence from molecular markers. Journal Europischer Orchideen 34: 707-731 9) Averyanov L. V. 1988 A review of genus Dactylorhiza Neck. ex Nevski (Orchidaceae), 1. Novosti sistematiki vysshikh rastenij 25: 48-67 10) Averyanov L. V. 1989 A review of genus Dactylorhiza Neck. ex Nevski (Orchidaceae), 2. Novosti sistematiki vysshikh rastenij 26: 47-56 11) Averyanov L. V. 1990a A review of genus Dactylorhiza Neck. ex Nevski (Orchidaceae), 3. Novosti sistematiki vysshikh rastenij 27: 32-62 12) Averyanov L. V. 1991 A review of genus Dactylorhiza Neck. ex Nevski (Orchidaceae), 4. Novosti sistematiki vysshikh rastenij 28: 33-42 13) Averyanov L. V. 1992 A review of genus Dactylorhiza Neck. ex Nevski (Orchidaceae), 5. Novosti sistematiki vysshikh rastenij 29: 14-25 14) , 32) Flora of Great Britain and Ireland: Butomaceae-Orchidaceae by Peter D. Sell and Gina Murrell, p340 15) The evolution of Dactylorhiza (Orchidaceae) allotetraploid complex : Insights from nrDNA sequences and cpDNA PCR-RFLP data, DEVOS Nicolas ; RASPE Olivier ; OH Sang-Hun ; TYTECA Daniel ; JACQUEMART Anne-Laure, Molecular phylogenetics and evolution 2006, vol. 38, no3, pp. 767-778 16) Nevski, S. A. 1935: Orchidaceae. Pp. 589-730 in: Komarov, V. L. (ed.), Flora SSSR 4. 17) pp 939-944, 1753 18) Elementa Botanica 3 p129, 1790 19) Dactylorhiza Nevski, the correct generic name of the Dactylorchids, by P.F. Hunt and V.S. Summerhayes [watsonia.org.uk] 20) Orchids of Russia and Adjacent Countries (within the borders of the former USSR) by Maria G Vakhrameeva, Koeltz Scientific Books, 2008 p114 21) http://www.myorchids.de/dactylorhiza1.htm [myorchids.de] 23) Dactylorhiza Cultivation Notes, author unknown, koolplants.co.uk [koolplants.co.uk] 24) The explorer's garden by Daniel J. Hinkley, p215 25) An encyclopedia of shade perennials by Wolfram George Schmid, p155 26) 29) , Mycorrhiza by Ajit Varma, Bertold Hock, p685 27) Terrestrial orchids: from seed to mycotrophic plant by Hanne N. Rasmussen, p295 onwards 28) Establishment of in vitro culture collection of endangered European orchids, Bulletin of Botanical Gardens, 13: 6973, 2004 30) http://www.lidaforsgarden.com/Orchids/dactylorhiza_eng.htm [lidaforsgarden.com] 31) http://web.telia.com/~u11206828/emedium.htm [web.telia.com] 33) Advances in economics and econometrics by David M. Kreps, Econometric Society. World Congress, Kenneth Frank Wallis, p971 34) Photo of hybrid in the wild: http://forum.terrorchid.org/viewtopic.php?p=9490 [forum.terrorchid.org] 35) In cultivation: http://www.heimische-orchideen-forum.de/forum/messages/1940.htm [heimische-orchideen-forum.de] 36) Field flora of the British Isles by Clive A. Stace, p685 Quick links
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The Culture Sheet tax onomy is based on the publication Vascular Plant Families and Genera compiled by R. K. Brummitt Published by the Royal Botanic Gardens, Kew in 1992 - With the permission of the Trustees of the Royal Botanic Gardens, Kew
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