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Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Eudicots
Ordo: Proteales

Familia: Proteaceae
Subfamilia: Grevilleoideae
Tribus: Roupaleae
Subtribus: Heliciinae
Genus: Helicia
Species: H. cochinchinensis – H. formosana – H. nilagirica – H. obovatifolia
Name

Helicia Lour.
Vernacular names
lietuvių: Šilkonė

Helicia is a genus of 110 species of trees and shrubs, constituting part of the plant family Proteaceae.[3] They grow naturally in rainforests throughout tropical South and Southeast Asia, including India, Sri Lanka, Indochina, Peninsular Malaysia to New Guinea and as far south as New South Wales.

Conservation

At global, national and regional government scales, many Helicia species have been threatened with extinction, as officially recognised by the International Union for Conservation of Nature (IUCN) and by continental, national and local governments. Sixteen species have official IUCN global conservation statuses of either "critically endangered", "endangered", "vulnerable" or "near threatened" (in terms of global extinction).
Naming and classification

In 1790, notable pioneer botanist João de Loureiro described this genus as Helicia in his publication Flora Cochinchinensis.[1][2] The type species for the genus was Helicia cochinchinensis, the type specimen of which was collected in Cochinchina, Vietnam.[1][2] The genus name derives from the Greek word "έλιξ" (élix), which refers to the petals, now called tepals, spirally revolving or simply rolling or coiling up on themselves, at anthesis (the flowering time when the anthers open).[1][4][5]

In 1831, botanist Nathaniel Wallich named Helicia robusta for a dried specimen of a cultivated plant in India,[6] based on the specimen's earlier 1814 name Roupala robusta by William Roxburgh.[7] Roxburgh's Calcutta botanic gardens cultivated the plant.

From the 1850s to the 1860s notable German–Australian botanist Ferdinand von Mueller formally described several new Australian species.[8][9][10] In the late 1800s and early 1900s Frederick M. Bailey concentrated further on additional Queensland species, writing descriptions of them in numerous scientific papers.[11][12][13]

In 1939, Hermann O. Sleumer described many additional Malesian species, especially in New Guinea.[14] In 1955, he published a revision of the genus.[15] In 1956, his treatment of the genus in Flora Malesiana was published.[16] From 1969 to the late 1990s botanist Don B. Foreman, who was based in Papua New Guinea and Australia, collected numerous additional species, which he formally described before he wrote the comprehensive reviews and flora treatments for the two regions;[17][18][19][20] notably in the authoritative Handbooks of the Flora of Papua New Guinea (1978–1995, to date 3 volumes), he wrote the chapters for Proteaceae and other families;[21] and in the authoritative Flora of Australia (1981–, 60 volume series) he wrote the treatment of Helicia.[4]

From the 1990s botanist Richard C. K. Chung, based in Malaysia, published new species formal descriptions and a revision of the 13 species occurring in Borneo.[22][23] In total, approximately 100 species have been formally scientifically described.[24][25][26]

Lawrie Johnson and Barbara G. Briggs grouped Helicia with Xylomelum in the subtribe Heliciinae, tribe Helicieae, and subfamily Grevilleoideae in their 1975 monograph "On the Proteaceae: the evolution and classification of a southern family".[27] However, genetics studies showed these two to be relatively unrelated, instead finding the closest genetic correlations between Hollandaea and Helicia, and therefore classifying them both in the subtribe Heliciinae within the tribe Roupaleae.[24]
Diversity and description
Helicia glabriflora from New South Wales, Australia

Helicia plants generally grow naturally as small trees, while some species grow as shrubs and some grow to medium-sized trees up to 30 m (100 ft).[16][21][28]

They grow naturally across the Malesia region with the major centre of species diversity of about fifty species in New Guinea. They grow naturally in the south west Pacific ocean region, and in north and eastern Australia. They grow naturally across southern and eastern Asia, including Indonesia, Malaysia and another centre of species diversity of about twenty species in southern China, extending to parts of the Indian subcontinent, the Philippines, Taiwan, and southern Japan.[16][21][24] The plant family Proteaceae's 1,700 species (approximate) have their greatest diversity in the southern hemisphere and smaller centres of diversity including some Helicia, in the near northern hemisphere. The species diversity of the plant family Proteaceae decreases further northwards. H. cochinchinensis has the natural distribution reaching furthest north to Japan where it grows into trees in the mountains of warmer parts and where no other species nor other Proteaceae genera occur. The same Japanese name Yama-mogashi (山もがし) for this species, also means the whole genus and the entire Proteaceae plant family.[29] In the New Guinea and southern China centres of species diversity, many species grow in forests, up to as tall as the sub-canopy, especially diverse in rainforests.[21][24] In Australia, they are generally components of rainforests, and prefer richer soils,[28] especially in the farthest south region of Helicia's global distribution, the Illawarra, New South Wales, south of Sydney, where only one species H. glabriflora occurs, preferring richer basalt soils.[30][31]
Cultivation

In India and east Asia Helicias have been cultivated in botanic gardens, from the 1800s.[7] In Australia they have rarely been cultivated, and were thought to have little horticultural value. The rusty-coloured new growth is attractive on some species.[28] In some of the better known Australian species, the flowers and fruit are generally not prominent, and plants can be slow growing. They are generally propagated by seed, the viability of which drops rapidly with time.[32]
Species

(this list may have a small number of species missing, presently it has 99, out of the approximate total stated by sources of 110)

Helicia acutifolia Sleumer[14][33] – New Guinea – Globally Vulnerable[34]
Helicia affinis Sleumer[14][33] – New Guinea
Helicia albiflora Sleumer[14][35] – New Guinea[2] – Globally Near threatened[36]
Helicia amplifolia Sleumer[14][37] – New Guinea – Globally Near threatened[38]
Helicia archboldiana Sleumer[39] – New Guinea
Helicia attenuata (Jack) Blume – Borneo[22]
Helicia australasica F.Muell.[8][13][17][40] – New Guinea, Australia[2][4][41] – Globally Vulnerable[42]
Helicia blakei Foreman[17] – Australia[2][4][43]
Helicia bullata Sleumer[44] – New Guinea
Helicia calocoma Foreman[18][45] – New Guinea[2] – Globally Vulnerable[46]
Helicia cameronii F.Muell[47] – New Guinea
Helicia carrii Sleumer[14](Syn: H. brassii, H. divaricata Sleumer (1939))[48] – New Guinea
Helicia celatus Foreman[20] – New Guinea[2]
Helicia ceylanica Gardner – Sri Lanka endemic[49]
Helicia clivicola W.W.Sm.[50] – China region
Helicia cochinchinensis Lour.[51] – China, Indochina, Taiwan, Japan[29]
Helicia coeruleopurpurea P.Royen[52] – New Guinea
Helicia commutata Sleumer[14][53] – New Guinea
Helicia dongxingensis H.S.Kiu[54] – China region
Helicia excelsa (Roxb.) Blume – Borneo[22]
Helicia falcata C.Y.Wu[55] – China region
Helicia ferruginea F.Muell.[10][13][17] – Australia[2][4]
Helicia finisterrae Lauterb.[56] – New Guinea
Helicia forbesiana F.Muell[57] – New Guinea
Helicia formosana Hemsl.[58] – China region
Helicia fragilis Foreman[20] – New Guinea[2]
Helicia fuscotomentosa Suess. – Borneo endemic[22]
Helicia glabriflora F.Muell.[9][13][17] – eastern Australia[2][4][30][31]
Helicia graciliflora Merr. – Philippines endemic[59]
Helicia grandifolia Lecomte – Vietnam endemic[60] – Globally Vulnerable[61]
Helicia grandis Hemsl.[62] – China region
Helicia grayi Foreman[17] – Australia[2][4][63]
Helicia hainanensis Hayata[64] – China region
Helicia hypoglauca Diels[65] – New Guinea, New Britain
Helicia insculpta Sleumer[14][66] – New Guinea[2]
Helicia insularis Foreman[18][67] – New Guinea[2] – Globally Endangered[68]
Helicia islandica Foreman[20] – New Guinea
Helicia kwangtungensis W.T.Wang[69] – China region
Helicia laiagamensis Foreman[18][70] – New Guinea[2]
Helicia lamingtoniana (F.M.Bailey) C.T.White ex L.S.Sm.[11][17] – Australia[2][4][71]
Helicia latifolia C.T.White[72] – New Guinea – Globally Near threatened[73]
Helicia lauterbachiana Sleumer[14](Syn: H. grandifolia Lauterb. (1910), non Lecomte (1910))[74] – New Guinea
Helicia ledermannii Diels[75] – New Guinea
Helicia lewisensis Foreman[17] – NE Queensland, Australia[2][4][76]
Helicia longespicata Sleumer[14][75] – New Guinea
Helicia longipetiolata Merr. & Chun[77] – China region
Helicia loranthoides C.Presl – Philippines endemic[78]
Helicia macrostachya Lauterb.[79] – New Guinea
Helicia maxwelliana Gibbs – Borneo: Sabah endemic, rare[22]
Helicia microneura C.T.White[80] (Syn: H. arguta Sleumer[14]) – New Guinea
Helicia microphylla Diels[81] – New Guinea
Helicia moluccana (R.Br.) Blume – Moluccas[82]
Helicia neglecta Diels ex Sleumer[14][81] – New Guinea Bismarck Archipelago endemic: New Britain, New Ireland – Globally Vulnerable[83]
Helicia nilagirica Bedd.[84] – Southeast Asia, Yunnan, India, Nepal
Helicia nortoniana (F.M.Bailey) F.M.Bailey[12][13][17] – Australia[2][4][85]
Helicia obovata Benn. – Java, Borneo[22]
Helicia obovatifolia Merr. & Chun[86] – China region

var. mixta (H.L.Li) Sleumer[87] – China region
var. obovatifolia[88] – China region

Helicia obtusata Sleumer[14](Syn: H. clemensiae Sleumer (1939))[89] – New Guinea
Helicia odorata Diels[90] – New Guinea
Helicia olivacea Sleumer[14][91] – New Guinea
Helicia oreadum Diels[92] – New Guinea
Helicia pallescens Diels[93] – New Guinea
Helicia paucinervia Merr. – Philippines endemic[94]
Helicia peekelii Lauterb.[95] – New Guinea Bismarck Archipelago: New Ireland endemic – Globally Vulnerable[96]
Helicia peltata C.T.White[97] – New Guinea – Globally Critically endangered[98]
Helicia petiolaris Benn. – Borneo[22]
Helicia platyphylla Sleumer[14][91] – New Guinea
Helicia polyosmoides Foreman[18][99] – New Guinea Bismarck Archipelago: Manus Island endemic[2] – Globally Critically endangered[100]
Helicia pterygota Sleumer – Borneo: Sabah endemic, rare[22]
Helicia pyrrhobotrya Kurz[101] – China region
Helicia recurva Foreman[19] – Queensland Australia[2][4]
Helicia rengetiensis Masam.[102] – China region
Helicia retevenia Sleumer[14][103] – New Guinea[2]
Helicia reticulata W.T.Wang[104] – China region
Helicia retusa Foreman[18][103] – New Guinea[2] – Globally Vulnerable[105]
Helicia robusta (Roxb.) R.Br. ex Wall. var. robusta – Malesia: Borneo,[22] Philippines, India[106]
Helicia rostrata Foreman[18][107] – New Guinea[2] – Globally Vulnerable[108]
Helicia rufescens Prain – Borneo, rare[22]
Helicia saruwagedica Sleumer[14][109] – New Guinea
Helicia saurauioides Sleumer[109] – New Guinea
Helicia schlechteri Lauterb.[110] – New Guinea
Helicia sellae-montis Sleumer[14][111][112] – New Guinea
Helicia serrata (R.Br.) Blume var. serrata – Borneo[22]
Helicia sessilifolia R.C.K.Chung – Borneo: Sabah & Sarawak endemic, uncommon[22][23]
Helicia shweliensis W.W.Sm.[113] – China region – Globally Endangered[114]
Helicia silvicola W.W.Sm.[115] – China region
Helicia sleumeri Foreman[20] – New Guinea
Helicia stelechantha Diels[116] – New Guinea
Helicia subcordata Foreman[18][116] – New Guinea[2] – Globally Critically endangered[117]
Helicia symplocoides R.C.K.Chung – Borneo: Sabah endemic, rare[22][23]
Helicia tibetensis H.S.Kiu[118] – China region
Helicia torricellensis Lauterb.[119] – New Guinea
Helicia tsaii W.T.Wang[120] – China region
Helicia uganensis Diels ex Sleumer[14][121] – New Guinea Bismarck Archipelago endemic: New Britain, New Ireland
Helicia varoyenii Foreman[20] – New Guinea
Helicia versteeghii Foreman[20] – New Guinea[2]
Helicia vestita W.W.Sm.[122]

var. longipes W.T.Wang[123] – China region
var. vestita[124] – China region incl. Thailand

Helicia wollastonii Ridl.[125] – New Guinea
Helicia yangchunensis H.S.Kiu[126] – China region

References

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"Helicia%". Australian Plant Name Index (APNI), Integrated Botanical Information System (IBIS) database (listing by % wildcard matching of all taxa relevant to Australia). Centre for Plant Biodiversity Research, Australian Government. Retrieved 26 Apr 2013.
"Helicia Lour. | Plants of the World Online | Kew Science". Plants of the World Online. Retrieved 3 Feb 2020.
Foreman, Don B. (1995a). "Helicia". In McCarthy, Patrick (ed.). Flora of Australia: Volume 16: Eleagnaceae, Proteaceae 1. Flora of Australia series. CSIRO Publishing / Australian Biological Resources Study. pp. 393–399. ISBN 978-0-643-05692-3.
Helicia formosana flowers closeup by mingiweng 3 July 2007 on Flickr
Helicia formosana flowers closeup by Foggy Forest 17 July 2012 on Flickr
Helicia formosana flowers closeup by mingiweng 12 June 2007 on Flickr
Wallich, Nathaniel (1831). "no. 2702 Helicia robusta". Numerical list of dried specimens of plants in the Museum of the East India Company which have been supplied by Dr. Wallich, superintendent of the [company's] botanic garden at Calcutta. London (published 1828–1849). Retrieved 21 Apr 2013 – via biodiversitylibrary.org.
Roxburgh, William; Carey, William (1814). "Roupala robusta Roxb.". Hortus Bengalensis, or a Catalogue of the Plants Growing in the Hounourable East India Company's Botanical Garden at Calcutta. (Hort. Bengal.). Serampore: Mission Press. p. 83. Retrieved 21 Apr 2013 – via wmcarey.edu/carey/hortus/.
Mueller, Ferdinand von (1857). "Nova genera et species aliquot rariores in Plagis Australiae Intratropicis nuperrime detecta: Helicia australasica". Hooker's Journal of Botany and Kew Garden Miscellany. 9: 22. Retrieved 21 Apr 2013.
Mueller, Ferdinand von (Aug 1860). "Helicia glabriflora". XIII. Fragmenta Phytographiae Australiae (fascicle). 2. Auctoritate Gubern. Coloniæ Victoriæ, Ex Officina Joannis Ferres. p. 91. Retrieved 7 Dec 2014 – via biodiversitylibrary.org.
Mueller, Ferdinand von (May 1862). "Helicia ferruginea". XVIII. Fragmenta Phytographiae Australiae (fascicle). 3. Auctoritate Gubern. Coloniæ Victoriæ, Ex Officina Joannis Ferres. p. 37. Retrieved 7 Dec 2014 – via biodiversitylibrary.org.
Bailey, F.M., (1899) Contributions to the Flora of Queensland. Queensland Agricultural Journal 5(4)
Bailey, F.M., (1901) The Queensland Flora 4
Bailey, F.M., (1913) Comprehensive Catalogue of Queensland Plants
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Sleumer, Hermann O. (1955). "Studies in old world Proteaceae, 5, A revision of the genus Helicia Lour". Blumea. 8: 7–79.
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Foreman, Don B. (1984). "A review of the genus Helicia Lour. (Proteaceae) in Australia". Brunonia. 6 (1): 59. doi:10.1071/bru9830059.
Foreman, Don B. (1985). "Seven new species of Helicia Lour. (Proteaceae) from Papua New Guinea". Muelleria. 6 (2): 79–92.
Foreman, Don B. (1986). "A new species of Helicia, new combinations and lectotypification in Triunia (Proteaceae) from Australia". Muelleria. 6 (3). pp. 193–96, fig. 1.
Foreman, Don B. (1988). "New species of Helicia Lour. (Proteaceae) from the Vogelkop Peninsula, Irian Jaya". Kew Bulletin. 53 (3): 669–681. doi:10.2307/4110485. JSTOR 4110485.
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Chung, Richard C. K. (2001). "Taxonomic notes on the Bornean Helicia and Heliciopsis (Proteaceae)" (PDF). Journal of Tropical Forest Science. 13 (3): 534–547. Retrieved 24 Apr 2013.
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Oi, Jisaburo (1965) [1953]. "Proteaceae: Helicia". In Meyer, Frederick G.; Walker, Egbert H. (eds.). Flora of Japan (in English): combined, much revised and extended translation by the author of his Flora of Japan (1953) and Flora of Japan, Pteridophyta (1957). Tokyo, Japan: National Science Museum. pp. 394–5.
Harden, Gwen J. (2001). "Helicia glabriflora – New South Wales Flora Online". PlantNET – The Plant Information Network System. 2.0. Sydney, Australia: The Royal Botanic Gardens and Domain Trust. Retrieved 25 Apr 2013.
Bofeldt, Anders (2011). "Table 1: Helicia glabriflora" (PDF). Plants at Risk in the Illawarra: Introduction to Table 1 (PDF) (Technical report). Online via LandcareIllawarra.org.au. NSW. Retrieved 25 Apr 2013. Lay summary – Tribute to Anders, Landcare Illawarra (2011).
Elliot, Rodger W.; Jones, David L.; Blake, Trevor (1992). Encyclopaedia of Australian Plants Suitable for Cultivation: Vol. 4: Eu-Go. Port Melbourne: Lothian Press. p. 285. ISBN 978-0-85091-213-5.
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"Helicia amplifolia". IUCN Red List of Threatened Species. 1998. 1998. Retrieved 28 Dec 2013.
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"Helicia australasica". Retrieved 22 Mar 2021. in Zich et al. (2020)
"Helicia australasica". IUCN Red List of Threatened Species. 1998. 1998. Retrieved 28 Dec 2013.
"Helicia blakei". Retrieved 22 Mar 2021. in Zich et al. (2020)
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"Helicia calocoma". IUCN Red List of Threatened Species. 1998. 1998. Retrieved 28 Dec 2013.
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Dassanayake, M. D.; Fosberg, F. R., eds. (1980). A Revised Handbook to the Flora of Ceylon. 2. New Delhi: Amerind Publ. Co. p. 484.
"Helicia clivicola". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
"Helicia cochinchinensis". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
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"Helicia dongxingensis". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
"Helicia falcata". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
Foreman (1995), p. 248.
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"Helicia formosana". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
"Helicia graciliflora". Archived from the original on 28 Dec 2013. Retrieved 23 Apr 2013. in LaFrankie (2011)
World Conservation Monitoring Centre (1998). "Helicia grandifolia". In: IUCN 2012. "IUCN Red List of Threatened Species. Version 2012.2". Archived from the original on 27 Jun 2014. Retrieved 23 Apr 2013.
"Helicia grandifolia". IUCN Red List of Threatened Species. 1998. 1998. Retrieved 28 Dec 2013.
"Helicia grandis". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
"Helicia grayi". Retrieved 22 Mar 2021. in Zich et al. (2020)
"Helicia hainanensis". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
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"Helicia insularis". IUCN Red List of Threatened Species. 1998. 1998. Retrieved 28 Dec 2013.
"Helicia kwangtungensis". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
Foreman (1995), pp. 251–253.
"Helicia lamingtoniana". Retrieved 22 Mar 2021. in Zich et al. (2020)
Foreman (1995), p. 253.
"Helicia latifolia". IUCN Red List of Threatened Species. 1998. 1998. Retrieved 28 Dec 2013.
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"Helicia lewisensis". Retrieved 22 Mar 2021. in Zich et al. (2020)
"Helicia longipetiolata". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
"Helicia loranthoides". Archived from the original on 28 Dec 2013. Retrieved 23 Apr 2013. in LaFrankie (2011)
Foreman (1995), pp. 254–255.
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"Helicia neglecta". IUCN Red List of Threatened Species. 1998. 1998. Retrieved 28 Dec 2013.
"Helicia nilagirica". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
"Helicia nortoniana". Retrieved 22 Mar 2021. in Zich et al. (2020)
"Helicia obovatifolia". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
"Helicia obovatifolia var. mixta". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
"Helicia obovatifolia var. obovatifolia". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
Foreman (1995), p. 257.
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"Helicia paucinervia". Archived from the original on 28 Dec 2013. Retrieved 23 Apr 2013. in LaFrankie (2011)
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"Helicia peekelii". IUCN Red List of Threatened Species. 1998. 1998. Retrieved 28 Dec 2013.
Foreman (1995), p. 260.
"Helicia peltata". IUCN Red List of Threatened Species. 1998. 1998. Retrieved 28 Dec 2013.
Foreman (1995), pp. 260–261.
"Helicia polyosmoides". IUCN Red List of Threatened Species. 1998. 1998. Retrieved 28 Dec 2013.
"Helicia pyrrhobotrya". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
"Helicia rengetiensis". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
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"Helicia reticulata". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
"Helicia retusa". IUCN Red List of Threatened Species. 1998. 1998. Retrieved 28 Dec 2013.
"Helicia robusta". Archived from the original on 28 Dec 2013. Retrieved 23 Apr 2013. in LaFrankie (2011)
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"Helicia rostrata". IUCN Red List of Threatened Species. 1998. 1998. Retrieved 28 Dec 2013.
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"Helicia shweliensis". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
"Helicia shweliensis". IUCN Red List of Threatened Species. 1998. 1998. Retrieved 28 Dec 2013.
"Helicia silvicola". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
Foreman (1995), p. 265.
"Helicia subcordata". IUCN Red List of Threatened Species. 1998. 1998. Retrieved 28 Dec 2013.
"Helicia tibetensis". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
Foreman (1995), p. 265–267.
"Helicia tsaii". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
Foreman (1995), p. 267.
"Helicia vestita". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
"Helicia vestita var. longipes". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
"Helicia vestita var. vestita". Retrieved 21 Apr 2013. in Qiu & Weston (2004)
Foreman (1995), p. 267–268.

"Helicia yangchunensis". Retrieved 21 Apr 2013. in Qiu & Weston (2004)

Cited works

Foreman, Don B. (1995). "Proteaceae". In Conn, Barry J. (ed.). Handbooks of the Flora of Papua New Guinea. 3. Melbourne: Melbourne University Press. pp. 221–270. Retrieved 22 Mar 2013 – via www.pngplants.org.
LaFrankie, J. V. (2011). "Helicia". Flora of the Philippines: Proteaceae [draft]. Flora of the Philippines online, updated 3 Sept 2012. University of the Philippines. Archived from the original on 29 Dec 2013. Retrieved 23 Apr 2013 – via philippineflora.info.
Qiu, Huaxing; Weston, Peter H. (2004). "Helicia Lour". Flora of China. Online version. eFloras. Beijing and St. Louis, MO: Science Press and Missouri Botanical Garden Press. Retrieved 21 Apr 2013.
Sleumer, Hermann O. (1956). "Helicia". Flora Malesiana Vol. 5 : Proteaceae. Series I, Spermatophyta : Flowering Plants. Leiden, The Netherlands: Rijksherbarium / Hortus Botanicus, Leiden University. pp. 164–190. Retrieved 21 Apr 2013.
Zich, F.A.; Hyland, B.P.M; Whiffen, T.; Kerrigan, R.A. (2020). "Australian Tropical Rainforest Plants Home". Australian Tropical Rainforest Plants, Edition 8. Commonwealth Scientific and Industrial Research Organisation (CSIRO). Retrieved 22 Mar 2021.

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