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Epipactis helleborine (L.) Crantz

Brief Summary

    Epipactis helleborine: Brief Summary
    provided by wikipedia

    Epipactis helleborine, the broad-leaved helleborine, is a terrestrial species of orchid. Its nodding flowers vary from greenish pink to purple, and it prefers shaded woodland environments.

Comprehensive Description

    Epipactis helleborine
    provided by wikipedia

    Epipactis helleborine, the broad-leaved helleborine, is a terrestrial species of orchid. Its nodding flowers vary from greenish pink to purple, and it prefers shaded woodland environments.

    Description

    Epipactis helleborine grows to a height of 80 cm and has broad dull green leaves which are strongly ribbed and flat. 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.[3]

    Distribution

    This species is widespread across much of Europe and Asia, from Portugal to China, as well as northern Africa.[1][4][5] In North America, it is 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.[6][7][8] In the US it is sometimes referred to as the "weed orchid" or "weedy orchid" and is increasingly considered invasive, in areas including Michigan,[9] Wisconsin[10] and the San Francisco Bay Area.[11]

    Habitat

    Found in woods and hedge-banks[12] and often, in Belfast, not far from paths near human activity.[13] It was even seen in Moscow, right in the middle of the city, including quite large populations. 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.[14]

    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. ^ Webb, D.A., Parnell, J. and Doogue,D . 1996. An Irish Flora. Dundalgan Press (W. Tempest) Ltd. Dundalk..mw-parser-output cite.citation{font-style:inherit}.mw-parser-output q{quotes:"""""'"'"}.mw-parser-output code.cs1-code{color:inherit;background:inherit;border:inherit;padding:inherit}.mw-parser-output .cs1-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/6/65/Lock-green.svg/9px-Lock-green.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-lock-limited a,.mw-parser-output .cs1-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Lock-gray-alt-2.svg/9px-Lock-gray-alt-2.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/a/aa/Lock-red-alt-2.svg/9px-Lock-red-alt-2.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-subscription,.mw-parser-output .cs1-registration{color:#555}.mw-parser-output .cs1-subscription span,.mw-parser-output .cs1-registration span{border-bottom:1px dotted;cursor:help}.mw-parser-output .cs1-hidden-error{display:none;font-size:100%}.mw-parser-output .cs1-visible-error{font-size:100%}.mw-parser-output .cs1-subscription,.mw-parser-output .cs1-registration,.mw-parser-output .cs1-format{font-size:95%}.mw-parser-output .cs1-kern-left,.mw-parser-output .cs1-kern-wl-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right,.mw-parser-output .cs1-kern-wl-right{padding-right:0.2em}ISBN 0-85221-131-7
    4. ^ Altervista Flora Italiana, Elleborine comune, Epipactis helleborine (L.) Crantz
    5. ^ Flora of China v 25 p 180, 火烧兰 huo shao lan, Epipactis helleborine (Linnaeus) Crantz
    6. ^ Flora of North America v 26 p 586, Epipactis helleborine (Linnaeus) Crantz
    7. ^ Biota of North America Program, county range map
    8. ^ "World Checklist of Selected Plant Families".
    9. ^ http://msue.anr.msu.edu/news/homeowners_battling_a_weedy_orchid_invading_lawns_and_flowerbeds
    10. ^ http://www.nps.gov/plants/alien/pubs/midatlantic/plants-to-watch.htm
    11. ^ http://ucjeps.berkeley.edu/cgi-bin/get_IJM.pl?tid=24407
    12. ^ 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
    13. ^ 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
    14. ^ “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.

    General Description
    provided by Plants of Tibet
    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.

Distribution

    Distribution
    provided by Plants of Tibet
    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).
    Distribution
    provided by eFloras
    Europe, N. Africa, Himalaya (Kashmir to Bhutan), S.E. Tibet, N. Asia.
    Distribution
    provided by eFloras
    Distribution: Northern hemisphere, Mediterranean region, S. W. Asia, Himalaya to Nepal, Sikkim, Bhutan up to 300 m. Widespread in the coniferous forests of Pakistan.
    Distribution
    provided by eFloras
    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.

Morphology

    Comments
    provided by eFloras
    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
    Comments
    provided by eFloras
    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.

    Description
    provided by eFloras
    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.
    Description
    provided by eFloras
    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.
    Elevation Range
    provided by eFloras
    2400-3200 m

Diagnostic Description

    Diagnostic Description
    provided by Plants of Tibet
    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).
    Synonym
    provided by eFloras
    Serapias helleborine Linnaeus, Sp. Pl. 2: 949. 1753; Epipactis latifolia (Linnaeus) Allioni

Habitat

    Habitat
    provided by Plants of Tibet
    Growing in forests, grasslands, wooded slopes, streamsides; 200-3600 m.
    Habitat
    provided by eFloras
    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.

Cyclicity

    Cyclicity
    provided by Plants of Tibet
    Flowering from June to July; fruiting in September.
    Flower/Fruit
    provided by eFloras
    Fl. Per.: End of June to August.
    Flowering/Fruiting
    provided by eFloras
    Flowering Jun--Oct.

Reproduction

    Reproduction
    provided by Plants of Tibet
    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.

Genetics

    Genetics
    provided by Plants of Tibet
    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).