Image of <i>Epipactis helleborine</i> ssp. <i>orbicularis</i> (K. Richt.) E. Klein
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Broad Leaved Helleborine

Epipactis helleborine (L.) Crantz

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A plant from Hazara, (Inayat no.23165, E) differs from typical Epipactis helleborine in having a few nearly orbicular leaves on a stout stem and a very dense inflorescence
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Flora of Pakistan Vol. 0: 15 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Epipactis helleborine is known to have been thoroughly naturalized in North America since 1879 (C. A. Luer 1975).

Extensive and striking color variations in Epipactis helleborine have led to the description of numerous varieties and forms. Most of the varieties originated in Europe, but the following forms originated in North America. With the exception of forma luteola, which has been noted only from Vermont, the other forms occur randomly throughout the range of the species: forma alba (Webster) B. Boivin, white-flowered form; forma luteola P. M. Brown, yellow-flowered; forma monotropoides (Mousley) Scoggan, albino; forma variegata (Webster) B. Boivin, variegated, the leaves with white and cream markings; forma viridens A. Gray, green-flowered.

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Flora of North America Vol. 26: 584, 585, 586, 591 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Description

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Plants slender or robust, up to 70(-100) cm tall. Rhizome short. Stem leafy throughout. Leaves spreading, ovate to ovate-lanceolate. Inflorescence normally densely many-flowered, up to 25-(35) cm long. Bracts lanceolate, lower exceeding flowers, the upper decreasing in size. Flowers from dark olive-green to yellowish-green, tinged ± intensively with red-purple. Sepals campanulate, ovate-lanceolate, up to 12 mm long; petals slightly shorter, ovate, paler green or pinkish. Labellum with cup-shaped hypochile, outside green, inside dark shining olive-brown, with nectar, 4-6 mm long and broad; epichile cordate, up to 5 mm long, the margin slightly crenulate-undulate, with 2 ± rugose bosses at base. Column short, 4-5 mm (including the anther). Ovary pedicelled, glabrous or with some hairs; ripe seed-capsule spreading.
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Flora of Pakistan Vol. 0: 15 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Description

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Plants 25–80(–100) cm, sparsely to densely pubescent. Leaves 3–10, orbiculate, elliptic to narrowly lanceolate, 4–18 × 1.5–8.5 cm. Inflorescences racemes, loose to moderately dense, commonly 1-sided; floral bracts spreading, linear to narrowly lanceolate, 10–40(–70) mm, often exceeding flowers. Flowers 15–50, small; sepals greenish, often suffused with purple; lateral sepals 10–13 × 5–6 mm, apex oblique; petals ovate, pale green, pink, purple, or yellowish, 9–11 × 4–6 mm; lip indistinctly veined, constricted at middle into 2 parts, proximal part purplish to brownish, deeply concave, not papillose, 9–12 × 8 mm, distal part recurved, pink, broadly triangular-ovate, ± flat to tip, 5 × 5 mm; calli 2, near base, brownish, not rugose; column 3–6 mm; ovary glabrous. Capsules obovoid, 9–14 mm, glabrate to densely pubescent. 2n = 36, 38, 40, 44.
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Flora of North America Vol. 26: 584, 585, 586, 591 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Distribution

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Europe, N. Africa, Himalaya (Kashmir to Bhutan), S.E. Tibet, N. Asia.
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Annotated Checklist of the Flowering Plants of Nepal Vol. 0 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Annotated Checklist of the Flowering Plants of Nepal @ eFloras.org
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Distribution

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Distribution: Northern hemisphere, Mediterranean region, S. W. Asia, Himalaya to Nepal, Sikkim, Bhutan up to 300 m. Widespread in the coniferous forests of Pakistan.
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Flora of Pakistan Vol. 0: 15 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Distribution

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introduced; B.C., N.B., Nfld. and Labr. (Nfld.), Ont., Que.; Ark., Calif., Conn., Ill., Ind., Ky., Maine, Md., Mass., Mich., Minn., Mo., Mont., N.H., N.J., N.Mex., N.Y., N.C., Ohio, Oreg., Pa., R.I., Tenn., Vt., Va., Wash., W.Va., Wis.; Eurasia; North Africa.
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Flora of North America Vol. 26: 584, 585, 586, 591 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Elevation Range

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2400-3200 m
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Annotated Checklist of the Flowering Plants of Nepal Vol. 0 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Annotated Checklist of the Flowering Plants of Nepal @ eFloras.org
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Flower/Fruit

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Fl. Per.: End of June to August.
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Flora of Pakistan Vol. 0: 15 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Flowering/Fruiting

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Flowering Jun--Oct.
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Flora of North America Vol. 26: 584, 585, 586, 591 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Habitat

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Moist to dry, rocky, shaded, deciduous to mixed woods; cedar swamps and forested stream margins; often in disturbed places such as lawns, and cracks in concrete sidewalks; 0--1300m.
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Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
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Flora of North America Vol. 26: 584, 585, 586, 591 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Synonym

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Serapias helleborine Linnaeus, Sp. Pl. 2: 949. 1753; Epipactis latifolia (Linnaeus) Allioni
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Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
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Flora of North America Vol. 26: 584, 585, 586, 591 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Flora of North America @ eFloras.org
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Cyclicity

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Flowering from June to July; fruiting in September.
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Diagnostic Description

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Epipactis helleborine var. helleborine is close relative of Epipactis helleborine var. tangutica, but differs from the latter in its flowers usually 7-40 (vs. 6-10), epichile of lip with a pair of semiorbicular, erose lamellae toward base (vs. a pair of thickened, fleshy wartlike calli toward base).
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Distribution

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Epipactis helleborine is occurring in Anhui, Gansu, Guizhou, Hebei, Hubei, Liaoning, Qinghai, Shaanxi, Shanxi, Sichuan, Xinjiang, Xizang, Yunnan of China, Afghanistan, Bhutan, Kazakhstan, Kyrgyzstan, Nepal, Pakistan, Russia, Tajikistan, Uzbekistan, N Africa, SW Asia, Europe, North America (naturalized).
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General Description

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Plants 20-70 cm tall. Rhizome short. Stem glabrous toward base, becoming pubescent above, with 2 or 3 scalelike sheaths near base. Leaves 4-7, ovate-orbicular, ovate, or elliptic-lanceolate, rarely lanceolate, upper ones narrower and lanceolate to linear-lanceolate, 3-13 cm long, 1-6 cm wide, glabrous, apex acuminate to long acuminate. Rachis 10-30 cm, pubescent, laxly to subdensely 7-40-flowered; floral bracts foliaceous, linear-lanceolate, lower ones exceeding flowers, becoming shorter above. Flowers usually nodding, resupinate, green or pale purple, out-crossing; pedicel and ovary 10-15 mm, yellow-brown tomentose. Dorsal sepal ovate-lanceolate, rarely elliptic, cymbiform, 8-13 mm long, 4-5 mm wide, apex acuminate; lateral sepals ovate-lanceolate, oblique, 9-13 mm long, ca. 4 mm wide, apex acuminate. Petals elliptic, 6.5-8 mm long, 3-4 mm wide, apex acute or obtuse; lip 6-8 mm, without a mesochile; hypochile semiglobose-saccate, 3-4 mm; epichile subtriangular or suboblate, ca. 3 mm long, 3-4 mm wide, veins sometimes thickened near apex, with a pair of semiorbicular, erose lamellae toward base of epichile, apex acute. Column 3-5 mm. Capsule obovoid-ellipsoid, ca. 10 mm, puberulent.
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Genetics

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The chromosomal number of Epipactis helleborine is 2n = 38, 40 (Averyanov et al., 1982; Stepanov and Muratova, 1992; Malakhova and Markova, 1994; Vij et al., 1995; D'Emerico et al., 1999).
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Habitat

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Growing in forests, grasslands, wooded slopes, streamsides; 200-3600 m.
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Reproduction

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Epipactis helleborine is pollinated by social wasps, and most populations show the morphological characteristics of outcrossing species. However, local predominantly selfing subspecies and varieties have been documented from drier habitats. To document geographic variation in floral morphology, ability to produce seeds through autogamy, and reproductive success in Epipactis helleborine, populations of Epipactis helleborine from three geographic regions along a latitudinal gradient of c. 1000 km from northern to southern Sweden (Ehlers et al., 2002). In the southernmost region, populations in dry and mesic habitats were compared. Supplemental hand-pollination was conducted to determine whether among-population variation in fruit set could be explained by differences in the natural level of pollination, and whether any relationship between floral morphology and fruit production could be explained by interactions with pollinators. Bagging experiments showed no evidence of autogamy in any of the study populations. Number of flowers, pollinia removal and fruit set varied significantly among populations but did not differ among regions. Pollinia removal was positively correlated with population size and both pollinia removal and fruit set were lower in dry than in mesic habitats. At the level of the individual plant, the number of pollinia removed increased more rapidly with flower number than did number of fruits produced. The hand-pollination experiment indicated that the positive relationship between number of flowers and fruit production was due to a higher degree of pollen limitation in plants with few flowers than in plants with many flowers. The experiment also showed that variation in the level of pollen limitation could only partly explain variation in fruit set among populations.
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Epipactis helleborine

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Flowers

Epipactis helleborine, the broad-leaved helleborine, is a terrestrial species of orchid with a broad distribution. It is a long lived herb which varies morphologically with ability to self-pollinate.[3]

Description

Epipactis helleborine grows to a maximum height of 92 cm (36 in) and has broad dull green leaves which are strongly ribbed and flat. [4] The flowers are arranged in long drooping racemes with dull green sepals and shorter upper petals. The lower labellum is pale red and is much shorter than the upper petals.[5]

Achlorophyllous, white Epipactis helleborine plants have been found. Achlorophyllous forms tend to be shorter, as small as 17cm. [6]

Flowering occurs June-September.[7]

Distribution

This species is widespread across much of Europe and Asia, from Portugal to China, as well as northern Africa.[1][8][9]

In North America, it is an introduced species and widely naturalized mostly in the Northeastern United States, eastern Canada and the Great Lakes Region, but also in scattered locations in other parts of the continent.[10][11][12] In the US it is sometimes referred to as the "weed orchid" or "weedy orchid" and continues to spread throughout the country to new areas including Michigan,[13] Wisconsin,[14] and the San Francisco Bay Area.[15]

Habitat

Found in woods and hedge-banks[16] and often not far from paths near human activity.[17] It is one of the most likely European orchids to be found within a city, with many sites for example in Glasgow, London and Moscow. Sometimes spotted beside car parks.[18]

Epipactis helleborine is known for its successful colonization of human-made or anthropogenic habitats such as parks, gardens or roadsides.[19] These roadside orchids exhibit special features such as large plant size and greater ability to produce flowers.[19] Pollination plays a huge role as pollinators such as Syrphidae, Culicidae, Apidae etc. possess greater species diversity and visits the flowering sites more in anthropogenic habitats as compared to native ones.[19] The visitation rates along with the reproductive success of these orchids are higher in large populations as they are more attractive to pollinators. [3]

Ecology

This species of orchid is pollinated by several species of Hymenoptera, particularly the common wasp, but also other species in the genera Vespula and Dolichovespula. Flowers release a sweet nectar to attract the wasps, which has an intoxicating effect on them.[20][21] Eight varieties of Epipactis helleborine in central Europe (Lower Silesia, Poland) had their nectar studied and they were found to contain naturally occurring oxycodone (as well as another narcotic-like opioid) in minute amounts.[22]

Epipactis helleborine requires a mycorrhizal symbiosis to germinate successfully and remains partially dependent upon the fungus when plants mature, however it is not particularly selective among fungal species. Fungi associated with the live roots include Tuber, Helotiales, Peziza, Leptodontidium, Hydnotrya and Wilcoxina.[23][24]

It has been suggested that the presence of this orchid species in a woodland is an indicator that edible truffles can be found there,[25] but this is not always the case.

Subspecies

A rather long list of names have been proposed for subspecies, varieties and forms of Epipactis helleborine, far too many to list here.[2] This is not unusual for such a widespread species. At present (June 2014) only the following are accorded international acceptance:[1]

  1. Epipactis helleborine subsp. bithynica (Robatsch) Kreutz - Turkey
  2. Epipactis helleborine subsp. degenii (Szentp. & Mónus) Kreutz - Greece
  3. Epipactis helleborine subsp. densifolia (W.Hahn, Passin & R.Wegener) Kreutz - Turkey
  4. Epipactis helleborine subsp. helleborine - widespread
  5. Epipactis helleborine subsp. latina W.Rossi & E.Klein - Italy, former Yugoslavia
  6. Epipactis helleborine subsp. leutei (Robatsch) Kreutz - Austria, Czech Republic
  7. Epipactis helleborine subsp. levantina Kreutz, Óvári & Shifman - Turkey
  8. Epipactis helleborine subsp. molochina (P.Delforge) Kreutz - Spain
  9. Epipactis helleborine subsp. neerlandica (Verm.) Buttler - Great Britain, Sweden, Belgium, Netherlands, France, Germany
  10. Epipactis helleborine subsp. orbicularis (K.Richt.) E.Klein - Austria, Czech Republic, Sweden, Germany, Switzerland, France, Spain, Italy, Baltic Republics
  11. Epipactis helleborine subsp. schubertiorum (Bartolo, Pulv. & Robatsch) Kreutz - Italy
  12. Epipactis helleborine var. tangutica (Schltr.) S.C.Chen & G.H.Zhu - China
  13. Epipactis helleborine subsp. tremolsii (Pau) E.Klein - France, Spain, Portugal, Sardinia, Italy, Algeria, Morocco

References

  1. ^ a b c Kew World Checklist of Selected Plant Species, Epipactis helleborine
  2. ^ a b Kew World Checklist of Selected Plant Families, Epipactis helleborine subsp. helleborine, synonyms
  3. ^ a b Ehlers, B. K.; Olesen, J. M.; Ågren, J. (2002). "Floral morphology and reproductive success in the orchid Epipactis helleborine: regional and local across-habitat variation". Plant Systematics and Evolution. 236 (1/2): 19–32. doi:10.1007/s00606-002-0197-x. ISSN 0378-2697. JSTOR 23644960. S2CID 9878820.
  4. ^ General Morphology and Anatomy of Chlorophyll-free and Green Forms of Epipactis helleborine
  5. ^ Webb, D.A., Parnell, J. and Doogue,D . 1996. An Irish Flora. Dundalgan Press (W. Tempest) Ltd. Dundalk.ISBN 0-85221-131-7
  6. ^ General Morphology and Anatomy of Chlorophyll-free and Green Forms of Epipactis helleborine
  7. ^ First Nature - Epipactis Helleborine
  8. ^ Altervista Flora Italiana, Elleborine comune, Epipactis helleborine (L.) Crantz
  9. ^ Flora of China v 25 p 180, 火烧兰 huo shao lan, Epipactis helleborine (Linnaeus) Crantz
  10. ^ Flora of North America v 26 p 586, Epipactis helleborine (Linnaeus) Crantz
  11. ^ Biota of North America Program, county range map
  12. ^ "World Checklist of Selected Plant Families".
  13. ^ "Broad-leaved helleborine: A weedy orchid invading lawns and flowerbeds".
  14. ^ http://www.nps.gov/plants/alien/pubs/midatlantic/plants-to-watch.htm
  15. ^ "Epipactis helleborine".
  16. ^ Clapham, A.R., Tutin, T.G. and Warburg, E.F. 1968. Excursion Flora of the British Isles. Cambridge University Press ISBN 0 521 04656 4
  17. ^ Beesley, S. and Wild, J. 1997. Urban Flora of Belfast. Institute of Irish Studies and The Queen's University of Belfast. ISBN 0-85389-695 X
  18. ^ PeerJ - Pollinator diversity and reproductive success of Epipactis helleborine (L.) Crantz (Orchidaceae) in anthropogenic and natural habitats
  19. ^ a b c Rewicz, Agnieszka; Jaskuła, Radomir; Rewicz, Tomasz; Tończyk, Grzegorz (2017-04-18). "Pollinator diversity and reproductive success of Epipactis helleborine (L.) Crantz (Orchidaceae) in anthropogenic and natural habitats". PeerJ. 5: e3159. doi:10.7717/peerj.3159. ISSN 2167-8359. PMC 5398293. PMID 28439457.
  20. ^ London Wildlife Trust - Orchid for July
  21. ^ Bioinfo - Epipactis helleborine
  22. ^ “Why do pollinators become 'sluggish'? Nectar chemical constituents from Epipactis helleborine L. Crantz Orchidaceae”. Applied Ecology & Environmental Research. 2005;3(2):29-38. Jakubska A, Przado D, Steininger M, Aniol-Kwiatkowska A, Kadej M.
  23. ^ Annals of Botany - Differences in mycorrhizal communities between Epipactis palustris, E. helleborine and its presumed sister species E. neerlandica
  24. ^ Mycorrhiza - Epipactis helleborine shows strong mycorrhizal preference towards ectomycorrhizal fungi with contrasting geographic distributions in Japan
  25. ^ Acta Biologica Szegediensis - Could orchids indicate truffle habitats? Mycorrhizal association between orchids and truffles

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Epipactis helleborine: Brief Summary

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Epipactis helleborine, the broad-leaved helleborine, is a terrestrial species of orchid with a broad distribution. It is a long lived herb which varies morphologically with ability to self-pollinate.

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