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15
Turczaninowia 2011, 14(2) : 15–100
УДК 582.594(597)
THE ORCHIDS OF VIETNAM ILLUSTRATED SURVEY.
Part 3. SUBFAMILY EPIDENDROIDEAE
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
EPIDENDROIDEAE
Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Summary. The article continues serial publication of illustrated critical taxonomical survey of orchids in the
ora of Vietnam. This part of the monograph includes taxonomical treatment of so-called primitive tribes (Neottieae,
Vanilleae, Gastrodieae, Nervilieae) of the largest subfamily Epidendroideae (17 genera and 41 species in Vietnam);
besides, some corrections and additions to rst and second parts of the monograph are included. Identication keys,
valid name, necessary synonyms, type material citation, short description, data on ecology and distribution, as well
as a list of studied voucher specimens for each species are provided. Mentioned species are illustrated with original line
drawings and color photographs. Two new nomenclature combinations are proposed, namely Cyrtosia faberi (Rolfe) Aver.,
and C. falconeri (Hook. f.) Aver. A new genus Miguelia Aver. with 2 species – M. somai (Hayata) Aver. and M. annamica
(Gagnep.) Aver. is established. Description of a new species – Peristylus tenuicallus Ormerod. discovered by P. Ormerod on
the base of his studies of AMES Herbarium is also presented.
Key words: Orchidaceae, ora of Vietnam, keys for identication.
Аннотация. Статья продолжает публикацию серийного издания иллюстрированного критического таксо-
номического обзора орхидных (Orchidaceae) флоры Вьетнама. Третья часть монографии включает предисло-
вие и стандартную таксономическую обработку наиболее примитивных триб (Neottieae, Vanilleae, Gastrodieae,
Nervilieae), объединяемых в подсемейство Epidendroideae и насчитывающих во флоре страны 17 родов и 41
вид. Для всех признаваемых видов приводятся законное название, наиболее важные синонимы, цитирование
аутентичного материала, краткое описание, данные по экологии и распространению, а также список изучен-
ных образцов. Все виды иллюстрированы черно-белыми рисунками и цветными фотографиями. Для 2 так-
сонов в работе предложены новые номенклатурные комбинации – Cyrtosia faberi (Rolfe) Aver. и C. falconeri
(Hook. f.) Aver. В качестве нового для науки описывается род Miguelia Aver., включающий два вида – M. somai
(Hayata) Aver. и M. annamica (Gagnep.) Aver. В публикацию также включено описание нового вида – Peristylus
tenuicallus Ormerod., открытого недавно Полом Омеродом на основании изучения гербарных коллекций Эйм-
са (AMES).
Ключевые слова: орхидные, флора Вьетнама, ключи для определения.
Ботанический институт им. В.Л. Комарова РАН, ул. Проф. Попова, 2; 197376, Санкт-Петербург, Россия;
av_leonid@yahoo.com
Russian Academy of Sciences, Komarov Botanical Institute, Prof. Popova, 2; 197376, St.-Petersburg, Russia.
Поступило в редакцию 25.11.2010 г. Submitted 25.11.2010
PREFACE
The article continues serial publication of il-
lustrated critical taxonomical survey of orchids in
the ora of Vietnam (Averyanov, 2008, 2010). The
third part of this monograph includes taxonomical
treatment of so-called primitive tribes – Neottieae,
Vanilleae, Gastrodieae and Nervilieae of the largest
subfamily – Epidendroideae with totally 17 genera
and 41 species, as well as some novelties that rep-
resent modern additions to the rst and second parts
of the monograph. As in earlier publications, illust-
rated survey is presented here in form of standard
taxonomic treatment, which includes identication
keys for all mentioned taxa and their short charac-
terization. Correct name (with standard taxonomic
reference), type, data about volume and distribution
is reported for each taxonomic group. Data for each
genus also include short description, total number of
species and number of species in the ora of Vietnam
(gures in brackets), as well as genus distribution.
Presented data for each species include:
– valid name, most signicant synonyms
and citation of most important recent monographs
and illustrations;
16 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
– available data about type;
– short description;
– available data on ecology, elevation of ob-
served habitats, phenology, frequency of occurrence
in the nature with approximate estimation of species
status according to IUCN categories for the territory
of Vietnam;
– general distribution and distribution in
Viet nam (in brackets are mentioned in alphabetical
order provinces where species was reported from);
– list of studied veried specimens and in-
dex of their host herbaria;
– when necessary short notices on species
taxonomy, biology, ecology, phenology or variation
are also provided;
– line drawings and color photographs ac-
company in the book each species reported for the
ora; collecting numbers of plants used as a model
for illustration are cited on drawings or in notices to
photographs.
Text of labels is maximally abbreviated whe-
never being cited. It usually includes only province
name, district name, collectors name and collecting
number, or if necessary date of collection. Largest
recent collections are abbreviated and designated as
series with following prexes:
CBL – Cao Bang Limestone – collections
on program of U.S.A. National Geographic Society
“Limestone Flora of Cao Bang Province of northern
Vietnam” (years 1998-1999, # 6300-98) with princi-
pal investigator Dr. Nguyen Tien Hiep;
CPC – collections made in expeditions ma-
na ged by the Center for Plant Conservation (mainly
Dr. Nguyen Tien Hiep, Prof. Leonid V. Averyanov
and Prof. Phan Ke Loc);
CPNP – collections in Cuc Phuong national
park (commonly without indication of collectors);
DDS – Prof. D.D. Soejarto – collections accor-
ding to International Cooperative Biodiversity Groups
program with this person as principal investigator;
DKH – Dr. D.K. Harder – collections ac-
cording to expeditions with this person as a princi-
pal investigator;
HAL – Dr. Nguyen Tien Hiep, Prof. Leonid
V. Averya nov, Prof. Phan Ke Loc – collections in
collaborative explorations of these persons;
HLF – Henry Luce Foundation, collections
of different collectors according to Vietnam Botani-
cal Conservation Program supported from Henry
Luce Foundation;
LX-VN – collections of Soviet (Liên Xô) –
Viet Name Expedition (commonly without exact in-
dication of collectors);
NMC – collections of staff member of Cuc
Phuong national park – Mr. Nguyen Manh Cuong
on the territory of the national park;
NTH – Dr. Nguyen Tien Hiep – collections
in expeditions with this person as principal investi-
gator;
P – Prof. Phan Ke Loc – collections in expe-
ditions with this person as principal investigator;
VA – Vietnam-American Series – collec-
tions of international group with Dr. N.T. Hiep as
principal investigator;
VH – Vietnamese Highlands – collections
on program of U.S.A. National Geographic Society
“Flora of Highlands of South Vietnam” (years 1993–
2001, # 5094-93, 5803-96, 6383-98) with principal
investigator Prof. Leonid V. Averyanov.
Modern administrative divisions of Vietnam
with names of administrative units used in the text,
as well as terminology list and terms explanation
we re presented in the rst part of this monograph
(Averya nov, 2008).
17
Turczaninowia 2011, 14(2) : 15–100
ACKNOWLEDGEMENTS
Laboratory studies, work with manuscript
and illustrations was supported by Swiss Orchid
Conservation fund of Zurich Foundation for
Orchid Conservation of Swiss Orchid Society.
Project – “Orchids of Vietnam, work on monograph
manuscript”, 2009–2010 and Russian Foundation
for Fundamental Researches (№ 09-04-90722).
We cordially thank authorities of the Center
for Plant Conservation of Vietnam Union of Sci-
ence and Technology Associations, Missouri Bo-
tanical Garden Vietnam Conservation program in
cooperation with Institute of Ecology and Biologi-
cal Resources of Vietnam Academy of Science and
Technology and Komarov Botanical Institute of the
Russian Academy of Sciences for comprehensive
help in organizations of all our investigations.
Field studies in Vietnam, the results of which
are presented in this paper, were funded by grants
from next organizations:
U.S.A. National Geographic Society –
“Flora of Highlands of the South Vietnam”, 1993–
2001 years (grants # 5094-93, 5803-96, 6383-98);
“Limestone Flora of Cao Bang Province of northern
Vietnam”, 1998–1999 (grant # 6300-98); “Botani-
cal Inventory of Unexplored Areas in Viet Nam: The
North”, 1999–2001 (grant # 6733-00); “Exploration
of rocky limestone ora and vegetation in Bac Kan
province, northern Vietnam”, 2003–2005 (# 7577-
04); “Exploration of highland ora and vegetation
in Lai Chau & Son La Provinces, north-western
Vietnam”, 2006–2007 (#8074-06); “Exploration of
primary woods along constructed highway Hanoi –
Ho Chi Minh for their sustainable conservation”,
2008–2011 (8418-08; 8800-10).
Henry Luce Foundation. Vietnam Bo-
tanical Conservation Program in Vietnam. 1999–
2008. “Preliminary updated checklist of orchids
(Orchidaceae) of Ba Be National Park”, 2002; “Pre-
liminary updated checklist of orchids (Orchidaceae)
of Bach Ma National Park”, 2003; “Preliminary
updated checklist of orchids (Orchidaceae) of Nui
Chua National Park”, 2004; “Preliminary updated
checklist of orchids (Orchidaceae) of Bu Gia Map
National Park”, 2005; “Preliminary Survey of the
Flora and Vegetation of Bi Doup – Nui Ba National
Park”, 2005; “Preliminary Survey of the Flora and
Vegetation of Da Krong Nature Reserve and allied
areas”, 2006; “Flora and vegetation of Pu Nat na-
tional park and allied areas”, March 2007; “Flora
and vegetation of Thuong Lo Municipality and allied
areas (Thua Thien – Hue Province)”, April 2007;
“Survey of the ora at Yok Don national park”,
March 2008; “Survey of the ora at Lo Go – Xa Mat
national park”, December 2008.
American Orchid Society. “Exploration
of endangered Vietnamese Paphiopedilums”, years
1996–1997; “Exploration of endangered calcium
dependent orchid ora in inaccessible rocky lime-
stone areas of the North Vietnam”, 1999–2000;
“Population studies of endemic Paphiopedilum spe-
cies in northern Vietnam”, 2001–2002; “Discovery
of endemic orchid ora in remote limestone areas
of Northern Vietnam”, 2003–2005; “Exploration of
Vietnamese orchid ora in regions allied to Laos
territory”, 2008–2009; “Assessment of orchid en-
demism in NW. Vietnam with special attention to
Paphiopedilum canhii”, 2011–2012.
Fauna & Flora International (Vietnam
Program). “The distribution of Paphiopedilum viet-
namense and its current status in the wild”, 2000;
“Community-based Conservation of the Hoang Lien
Mountain Ecosystem, Vietnam, Flora and vegeta-
tion survey of Van Ban district, Lao Cai province
of northern Vietnam”, 2002; “Preliminary survey of
orchids and gymnosperms in Trung Khanh district,
Cao Bang province northern Vietnam”, 2004.
Fauna & Flora International Vietnam
Conservation Support Program of the Ministry
of Agriculture and Rural Development, Vietnam.
“Pu Luong – Cuc Phuong Limestone Landscape
Conservation Project, Preliminary botanical survey
of primary vegetation in Pu Luong nature reserve”,
2003.
Fauna & Flora International (Vietnam
Program) & Counterpart International. “Pre-
liminary survey of Orchids (Orchidaceae) in Phong
Nha – Ke Bang National Park”, 2005.
WWF Indochina Programme. Green Cor-
ridor Project in Thua Thien – Hue Province, VN
085301. “Lowland ora and vegetation. Preliminary
survey”, 2005.
Basic Research program in Life Sciences
of Viet Nam, # 611001. “Threatened conifers and
cycads of Vietnam”, 2000–2002.
The Rufford Small Grant Foundation.
“Assessment of distribution and natural status of
Paphiopedilum canhii, Vietnam”, 2010–2011.
Chicago Zoological Society, Chicago
Board of Trade Endangered Species Fund. “As-
sessment of current natural status of critically en-
dangered species – Paphiopedilum canhii for its
conservation”, 2010–2011.
18 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Author cordially thanks T. Maisak, who was
very helpful in preparation of ink line drawings. We
also thanks Prof. Phan Ke Loc for photos used in
g. 24 e, f; Mr. Nguyen Sinh Khang for photos used
in g. 5 e, f and Mr. Pham Van The for photos used
in g. 24 i; 27 a, b. I also thank Dr. P. Emov for
permission of use his line drawing presented in g.
2 and Dr. A. Sennikov for translation of diagnosis of
Miguelia into Latin and valuable advices in nomen-
clature. Many signicant amendments and additions
were made by Paul Ormerod, Andre Schuiteman
and Eric Christenson that essentially improved pre-
sented treatment.
SUBFAMILY 5 EPIDENDROIDEAE LINDL.,
1821, Collect. Bot. App. Epidendreae. – Sub-
fam. Malaxidoideae Burnett, 1835, Outlines Bot.:
461 (sub “Malaxidae”). – Subfam. Arethusoideae
Endl., 1837, Gen. Pl.: 216 (sub “Arethuseae”).
Type: Epidendrum L.
220(~60) genera and 10 000(~480) species.
Tropical, subtropical and temperate regions of the
World, boreal zone of Northern Hemisphere.
Group of primitive tribes (Neottieae,
Vanilleae, Gastrodieae, Neottieae).
Trib. 5.1. Neottieae Lindl.,
1821, Collect. Bot. App. – Trib. Epipactieae
Endl., 1830, Fl. Poson.: 163 (sub “Epipactideae”). –
Trib. Listereae Endl., 1842, Mant. Bot. Suppl. 2: 19
(sub “Listeridae”). – Trib. Limodoreae Nees, 1845,
Gen. Pl. Monocot. 8, tab. 15 (sub “Limodorinae”). –
Subtrib. Limodorinae Benth., 1881, Journ. Linn. Soc.
London (Bot.) 18: 288 (sub “Limodoreae”). – Sub-
trib. Cephalantherinae Ptz., 1887, Entw. Nat. An-
ord. Orch.: 98 (sub “Cephalanthereae”). – Subtrib.
Epipactidinae M. Schulze, 1894, Orch. Deutsch.: 8
(sub “Epipactideae”).
Subtrib. 5.1.1 Limodorinae Benth.,
1881, Journ. Linn. Soc. Bot. 18: 288.
Type: Limodorum Boehm.
3(2) genera and 30–40(7) species. Temper-
ate, subtropical and tropical regions of the World.
Epipactis Zinn.,
1757, Cat. Pl. Gott.: 85, nom. cons.; Sei-
denf., 1978, Dansk Bot. Ark. 32, 2: 116–117; id.,
1992, Opera Bot. 114: 21-22; Aver., 1994, Ident.
Guide Vietnam. Orch.: 24; P.H. Ho, 2000, Ill. Fl.
Vietnam 3: 780; Su Horng-Jye, 2000, Fl. Taiwan 5:
858–861; Pearce et Cribb, 2002, Orch. Bhutan: 42–
47; Aver. et Averyanova, 2003, Updated Checklist
Orch. Viet.: 32; Newman et al, 2007, Checkl. Vasc.
Pl. Lao PDR: 267; Schuiteman et al., 2008, Nord.
Journ. Bot. 26: 289; Chen Sing-chi et al., 2009, Fl.
China, 25: 179–183.
Type: E. helleborine (L.) Crantz (Serapias
helleborine L.).
Terrestrial or lithophytic herbs with un-
derground rhizome, erect leafy stems and terminal
few- to many-owered, more or less secund raceme.
Leaves plicate, sessile, sheathing at the base. Flow-
ers medium-sized, resupinate, rare not resupinate.
Sepals and petals free, lip with different callosities,
divided into, concave hypochile and at elongate or
circular epichile. Column short. Pollinia 2, mealy,
normally with viscidium, without caudicles and
stipe.
25–30(2) species. N. Africa, Eurasia,
N. America.
Key to species
1. Riparian plants of swampy stream valleys, 35–120 cm tall; leaves numerous, narrowly ovate; sepals 1.2–
1.5 cm long, yellow-green with purple-brown margin; epichile narrowly obovate, straight along margin .........
........................................................................................................................................... 1. E. atromarginata
- Plants of dry rocky limestone, 15–25 cm tall; leaves 2–3, ovate; sepals less than 1 cm long, purple; epichile
circular, undulate along margin ....................................................................................................... 2. E. alata
1. E. atromarginata Seidenf.,
1992, Opera Bot. 114: 22, 23, 461, g. 5, pl.
1c; Aver., 1994, Identif. Guide. Vietnam. Orch.: 24;
P.H. Ho, 2000, Ill. Fl. Vietnam 3: 780, g. 10883;
Aver. et Averyanova, 2003, Updated Checklist Orch.
Viet.: 32. – E. ava auct. non Seidenf.: P.H. Ho,
2000, l.c.: 780, g. 10884.
Described from S. Vietnam (“Prov. Gialai-
Kontum. Kon Ha Nung”). Type (“15.05.1985 LX-
VN 1975”) – HN (holotype), LE (isotype).
Riparian or swampy herb 35–120 cm tall
with creeping stout rhizome. Stem erect, glabrous,
with numerous narrowly ovate, acuminate leaves.
Raceme secund, pubescent, usually with 5–12 re-
supinate, well opening owers 2 cm across. Sepals
and petals yellow-green with purple-brown margin,
subsimilar, ovate, acute, 12–15 mm long, densely
white pubescent outside. Lip light reddish-brown
19
Turczaninowia 2011, 14(2) : 15–100
with yellow-green base and center, 14 mm long, di-
vided into hypochile and epichile. Hypochile con-
cave, with ovate side lobes striped with dark purple-
brown nerves, disc with broad hemispheric warty
callus. Epichile narrowly obovate, nely rugose,
curved, with callus-like umbo at center. Column
light green, 6 mm tall, erect, stout, with massive
stigma and large forward protruding green anther
cap. Fig. 1; 5 a, b.
Ecology. Wet alluvial banks and riparian
rocks along streams and small rivers in shady broad-
leaved forests. 400–700 m. Fl. March-May. Very
rare (CR).
Distribution. Vietnam (Gia Lai, Quang Tri).
Laos?
Studied specimens. Gia Lai, Kon Ha Nung,
LX-VN 1975 (HN, LE); Gia Lai, Konplong, LX-VN
2275 (HN, LE); Quang Tri, Da Krong, HLF 5757
(HN), HLF 6139 (HN), HLF 6181 (HN, LE).
Notes. Critically endangered species of low-
land stream valleys.
2. E. alata
2006, Rheedea, 16, 1: 4–6, g. 3 d-g; Chen
Sing-chi et al., 2009, Fl. China, 25: 182.
Described from Vietnam (“Ha Giang Prov.,
Meo Vac Distr., Sung Tra Municipality…”). Type
(“11 December 2005 HAL 8513”) – HN (holotype),
LE (isotype).
Terrestrial herb 15–25 cm tall with short
rhizome. Stem erect, glabrous, with 2–3 distant,
ovate, leaves. Raceme, sparsely pubescent, with 2–5
distant, not resupinate, campanulate, purple-violet
owers 1 cm across. Sepals and petals subsimilar,
ovate, acute, 8–10 mm long, outside sparsely pubes-
cent. Lip purple-violet with yellow center, 10 mm
long, divided into hypochile, mesochile and epichile.
Hypochile concave, with triangular side lobes, disc
with numerous small deep brown warts. Mesochile
elongate, 3.5 mm long, 2 mm wide, with two broad
eshy low keels. Epichile circular, 3.5 mm across,
nely undulate and folded along the margin. Co-
lumn white, 5–7 mm tall, broadening from narrow
stalk into massive apex covered with forward pen-
dent anther cap; stigma 3 mm long, 4 mm wide, with
prominent rectangular, forward directed side lobes.
Fig. 2; 5 c, d; 6.
Ecology. Coniferous forests with Tsuga
chinensis on rocky limestone. 1000–1200 m. Fl.
November – December. Very rare (CR).
Distribution. Vietnam (Ha Giang). S. China
(SE. Yunnan).
Studied specimens. Type only.
Notes. Critically endangered species of pri-
mary limestone coniferous forests. Brown papillae
on honey-yellow disc of hypochile strikingly resem-
ble insect larvae that probably have certain role in
pollinators attraction.
Aphyllorchis Blume,
1825, Bijdr. 6, g. 77; id. 1849, Mus. Bot.
Lugd. 18: 30, emend.; Gagnep., 1934, Fl. Gen.
Indo-Chine 6, 5: 577–580; Seidenf., 1978, Dansk
Bot. Ark. 32, 2: 118–122; id., 1992, Opera Bot. 114:
24–25; Comber, 1990, Orch. Java: 50–51; id., Orch.
Sumatra: 118–121; Seidenf., Wood, 1992, Orch.
Malay. Sing.: 44–45; Aver., 1994, Ident. Guide Viet-
nam. Orch.: 24–25; P.H. Ho, 2000, Ill. Fl. Vietnam
3: 779; Su Horng-Jye, 2000, Fl. Taiwan 5: 749–751;
Pearce et Cribb, 2002, Orch. Bhutan: 36–37; Aver.
et Averyanova, 2003, Updated Checklist Orch. Viet.:
11; Newman et al, 2007, Checkl. Vasc. Pl. Lao PDR:
253; Schuiteman et al., 2008, Nord. Journ. Bot. 26:
364; Chen Sing-chi, Gale, 2009, Fl. China, 25: 177–
179. – Sinorchis S.C. Chen, 1978, Acta Phytotax.
Sin. 16, 4: 82.
Type: A. pallida Blume.
Achlorophyllous leaess terrestrial herbs
with short erect rhizome, few spreading thick roots,
erect stem and terminal inorescence of few- to
many resupinate owers. Sepals and petals subsimi-
lar, free, more or less spreading. Lip with distinct
short, concave, narrow hypochile to which more or
less 3-lobed epichile is joined. Column long, slen-
der, with erect apical anther. Column foot absent.
Pollinia 2, powdery.
15(5) species. Mainland tropical and sub-
tropical Asia, Indonesia, Philippines, Australia, New
Guinea.
Fig. 1. Epipactis atromarginata (HLF 5757): a –
ower, b – attened sepals and petals, c – attened lip.
20 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 2. Epipactis alata (HAL 8513, type): a – owering plant, b – inorescence, c – ower, d – attened sepals
and petals, e – attened lip, f-h – side, ventral and dorsal views of the column, i – pedicel and ovary.
21
Turczaninowia 2011, 14(2) : 15–100
Key to species
1. Petals and lip subsimilar; owers sub-actinomorphic; lip simple, oblong to elliptic, without ornamentation,
not divided into hypochile and epichile ...................................................................................... 1. A. simplex
- Flowers distinctly zygomorphic; lip very different from petals, distinctly divided into hypochile and epichile,
with spongia-like inations on its surface ...................................................................................................... 2
2. Stem rather slender, 15–50 cm tall; sepals not caudate, less than 1.5 cm long .............................................. 3
- Stem stout, up to 1.5 m tall; sepals caudate, longer than 1.5 cm .................................................................... 4
3. Stem commonly 15–30 cm tall; owers white, less than 8 mm across; sepals 4–5 mm long; epichile rough at
....................................................................................................................................................... 2. A. pallida
- Stem commonly 30–50 cm tall; owers yellow, more than 8 mm across; sepals 8–10 mm long; epichile with
fat, nely warty margin ............................................................................................................. 3. A. montana
4. Midlobe of epichile narrowly cuneate, acuminate to linear-subulate, often nely warty along margin, less
than 2 mm wide ........................................................................................................................... 4. A. evrardii
- Midlobe of epichile ovate to narrowly-ovate, 4–7 mm wide, papillose, with fat sponge-like, warty margin,
long caudate at apex ............................................................................................................. 5. A. annamensis
1. A. simplex Tang et F.T. Wang,
1951, Acta Phytotax. Sin. 1: 67. – Sinorchis
simplex (Tang et F.T. Wang) S.C. Chen, 1978, Acta
Phytotax. Sin. 16, 4: 83.
Described from S. China (“North-east
Kwang tung: Mei Hsien, Yin-na-shan, steep rocky
slope, ower white, …”). Type (“Aug. 4–31, 1932
W.T. Tsang no. 21504”) – PE?
Stem erect, slender, light yellow-brownish,
sometime with violet tint, 25–35 cm tall, with sev-
eral short sheaths and many-owered, lax inores-
cence 10–15 cm long, with 10–12 campanulate,
not widely opened owers. Floral bracts triangular-
cuneate, to 1 cm long, down reexed. Pedicel and
ovary 1.8–2.2 cm long, with sparse glandular hairs.
Sepals and petals dull yellow, with purple streaks,
subsimilar, narrowly elliptic, 8–10 mm long, acute
to obtuse, sepals eshy and keeled in apical half.
Lip light yellow, oblanceolate, elongate, simple,
in shape and size similar to petals, 9–10 mm long,
2–2.5 mm wide, thin, without ornamentation. Co-
lumn 9–11 mm long, bended at the middle, widened
to the apex, apically from both sides with 2 large
curved staminodes and erect ligulate rostellum as
tall as or taller than operculum. Stigma subterminal.
Fig. 3; 5 e, f.
Ecology. Primary evergreen broadleaved forests
with short bamboo on alluvial limestone slopes. 1100–
1200 m. Fl. October – November. Very rare (DD).
Distribution. Vietnam (Hoa Binh). S. China
(E. Guangdong).
Studied specimens. Hoa Binh, Tan Lac,
Ngoc Son – Ngo Luong nature reserve, CPC 775
(CPC Herbarium, LE).
Notes. This species supercially resembles
just a peloric sub-actinomorphic form of A. mon-
tana. Meanwhile, its column with curious large
curved staminodia and erect prominent ligulate
rostellum is very characteristic that was also men-
tioned earlier (Chen Sing-chi, Gale, 2009a). Most
probably, this plant represents fairly distinct species,
which needs further study. Unfortunately, it is very
rare and highly endangered.
2. A. pallida Blume,
1825, Bijdr., tab. 16, g. 77; Seidenf., 1978,
Dansk Bot. Ark. 32, 2: 118, 120, g. 74; Comber,
1990, Orch. Java: 51, g.; id., 2001, Orch. Sumatra:
120, g.; Seidenf., Wood, 1992, Orch. Malay. Sing.:
44, g. 12a-d.
Described from Java (“Salak”). Type – ?
Stem erect, slender, silvery-white with
sparse violet streaks, 15–30 cm tall, with several
dark brownish short sheaths and few-owered in-
orescence 2–6 cm long. Floral bracts ovate to
broadly-cuneate, 3–4 mm long. Pedicel and ovary
silvery white with dark violet streaks, 5–12 mm
long. Flowers white, not widely opening. Sepals
and petals subsimilar, ovate, obtuse, 4–5 mm long,
white, sometimes with yellowish tint, with dark vio-
let marks. Lip shorter than tepals. Hypochile with
erect, triangular side lobes. Epichile hinged at the
apex of hypochile, triangular, with 2 indistinct broad
side lobes and small eshy, obtuse midlobe. Column
white, 2.5–3 mm long. Fig. 4 a-c; 5 g, h.
Ecology. Primary mixed and coniferous for-
ests on deep silicate soils. 800–1500 m. Fl. August –
October. Very rare (VU).
Distribution. Vietnam (Dak Lak, Lam
Dong). Thailand, Malacca, Sumatra, Java, Kaliman-
tan, Philippines.
Studied specimens. Dak Lak, Chu Yang Sin
mt., HLF 5423 (HN, LE); Lam Dong, Lac Duong,
Hon Giao Ridge, HLF 5328a, HLF 5344 (HN, LE).
22 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 3. Aphyllorchis simplex (CPC 775): a – owering plant, b – ower, c – attened sepals, petals and lip, d –
column, side view, e – column apex, frontal view, f – pedicel and ovary.
23
Turczaninowia 2011, 14(2) : 15–100
3. A. montana Reichenb. f.,
1877, Linnaea 41: 57; Garay, Sweet, 1974,
Orch. Ryukyu Isl.: 45, 46, g. 2; Seidenf., 1978,
Dansk Bot. Ark. 32, 2: 121, 122, g. 75; id., 1992,
Orch. Indochina: 25; Seidenf., Wood, 1992, Orch.
Malay. Sing.: 45, g. 12 e-o; Aver., 1994, Identif.
Guide. Vietnam. Orch.: 24; P.H. Ho, 2000, Ill. Fl.
Vietnam 3: 779, g. 10882; Su Horng-Jye, 2000, Fl.
Taiwan 5: 750, g. 318; Comber, 2001, Orch. Suma-
tra: 119, g.; Pearce et Cribb, 2002, Orch. Bhutan:
37, g. 7 k-s; Aver. et Averyanova, 2003, Updated
Checklist Orch. Viet.: 11.
Described from Sri Lanka (“Ambagumowa
District”). Lectotype (“Thwaites CP 3189”) – K.
Stem erect, slender, yellow-brownish to
nearly white, 30–50 cm tall, with several short
Fig. 4. Aphyllorchis pallida (HLF 5328a): a – column and attened sepals and petals, b – lip, side view, c – at-
tened lip; A. montana (HAL 2992): d – column and attened sepals and petals, e – attened lip; A. evrardii (VH 2328):
f – median sepal, g – lateral sepal, h – column and lip, side view; i – column and attened lip, frontal view.
24 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 5. Epipactis atromarginata: a, b (HLF 6181); E. alata: c, d (HAL 8513, type); Aphyllorchis simplex: e, f
(CPC 775); A. pallida: g, h (HLF 5328a); A. montana: i (HAL 2992).
25
Turczaninowia 2011, 14(2) : 15–100
Fig. 6. Digital herbarium specimen of Epipactis alata (Averyanov L. HAL 8513, epitype).
26 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
sheaths and few- to many-owered inorescence
5–15 cm long. Floral bracts linear-cuneate, to 1.5 cm
long, reexed. Pedicel and ovary 1.5–2 cm long,
elongated after anthesis. Sepals and petals, dull yel-
low, sometimes with purple tint, subsimilar, elliptic,
8–10 mm long, sepals eshy and keeled in apical
half. Lip yellow to yellow-brownish, with white
spots. Hypochile with erect, triangular, acute lobes.
Epichile triangular-ovate, 6–8 mm long, with indis-
tinct erect roundish side lobes; midlobe eshy, ru-
gose, concave, obtuse, with fat nely warty margin.
Column 1 cm long, bended at the middle, widened
to the apex. Fig. 4 e d; 5 i; 23 a.
Ecology. Broadleaved forests on deep lime-
stone and silicate soils. 350–800 m. Fl. September –
October. Rare (VU).
Distribution. Vietnam (Kien Giang, Ninh
Tuan, Thanh Hoa, Vung Tau). Mainland tropical
Asia, Hainan, Taiwan, Kalimantan, Philippines.
Studied specimens. Cana, Phanrang, Poila-
ne s.n.; Krong Pha, Hayata 949 (P); Thanh Hoa, Ba
Thuoc, HAL 2992 (HN, LE); Kien Giang, Phu Quoc,
a. 2007 N.V. Khoi, s.n. (LE – photo); Pulo Condor,
Harmand s.n.
4. A. evrardii Gagnep.,
1931, Bull. Mus. Hist. Nat. (Paris) 2 ser. 3,
7: 680; Seidenf, 1978, Dansk Bot. Ark. 32, 2: 118,
120, g. 73; id., 1992, Orch. Indochina: 24, 25, g.
7; Aver., 1994, Identif. Guide. Vietnam. Orch.: 24;
P.H. Ho, 2000, Ill. Fl. Vietnam 3: 779, g. 10881;
Aver. et Averyanova, 2003, Updated Checklist Orch.
Viet.: 11.
Described from Vietnam (“Annam: Dalat”).
Type (“F. Evrard, N 1846”) – P.
Stem brownish-yellow, with violet stripes,
stout, erect, up to 1.2 m tall, with broad sheaths
and many-owered inorescence up to 30 cm long.
Floral bracts narrowly-cuneate, acuminate, 3–4 cm
long. Pedicel and ovary 3.5–4 cm long, elongated
after owering. Sepals and petals subsimilar, light
yellowish, with 3 violet nerves, narrowly ovate to
lanceolate, 2–4.5 cm long, attenuate to long caudate
apex. Hypochile, fused with base to column, side
lobes narrow, erect, falcate, 2–3 mm long, round-
ish at the apex. Epichile 3-lobed, 1.4–1.6 cm long;
side lobes, erect, hemicircular; midlobe narrowly-
cuneate, acuminate, 9–11 mm long, with fat, nely
papillose margin. Column slender, broadening to
the apex, 9–11 mm long. Anther large, mitre-form,
oblique. Fig. 4 f-i.
Ecology. Primary broadleaved montane
forests on deep silicate soils. 1200–1700 m. Fl. No-
vember – January. Very rare (CR).
Distribution. Vietnam (Gia Lai, Lam Dong).
Thailand, Laos.
Studied specimens. Benon Da Treu, Tixier
27 (P); Dalat, Evrard 1846 (P); Kon Tum, Dak Gley,
VH 2328 (HN, LE); Manline, Tixier drawing (P).
5. A. annamensis Aver.,
1996, Bot. Journ. (St. Petersburg), 81(10):
82, g. 6; Aver. et Averyanova, 2003, Updated
Checklist Orch. Viet.: 11.
Described from Vietnam (“Prov. Kon Tum,
Distr. Dak Gley, about 7–8 km to the S. of Dak Gley
town on Dak Poko River near Dak Tung village”).
Type (“3 December 1995 VH 2221”) – HN (holo-
type), AAU, LE, MO, P (isotypes).
Stem brownish-yellow, often with violet
marks, stout, erect, up to 1.5 m tall, with broad
sheaths and many-owered inorescence up to
30 cm long. Floral bracts linear-cuneate, 3–4 cm
long. Pedicel and ovary 3.5–4 cm long, after ower-
ing elongating. Sepals and petals subsimilar, white
to light yellowish, with 3 violet nerves, narrowly
ovate to lanceolate, 1.8–4 cm long, attenuate to long
caudate apex. Hypochile white to light yellowish,
fused with broad base to column, side lobes nar-
row, erect, falcate, 3–4 mm long. Epichile 3-lobed,
chestnut-brown with white marks, 1.4–1.6 cm long,
moveably articulated to apex of hypochile; side
lobes dull violet, erect, broadly-rounded; midlobe
triangular-ovate, 8–12 mm long, papillose, with fat
warty margin, long caudate. Column pale yellow-
green with violet tint, slender, 1–1.4 cm long. An-
ther yellow. Fig. 7; 23 b, c.
Ecology. Primary broadleaved forests on
silicate deep soils. 800–2000 m. Fl. November –
February. Rare (EN).
Distribution. Vietnam (Dak Nong, Kon
Tum, Nghe An). Endemic.
Studied specimens. Dak Nong, Dak Glong,
Ta Dung mt., HLF 5617 (HN, LE); Nghe An, Quy
Chau, Phu Lon peak HLF 3102 (HN, LE).
Subtrib. 5.1.2 Neottiinae Reichenb. f.,
in Seem., 1868, Fl. Vit.: 293.
Type: Neottia Guett.
3(1) genera and 70–100(1) species. Europe,
boreal, temperate and subtropical regions of main-
land Asia.
Listera R. Br.,
1813, in Ait. et Ait. f. Hortus Kew. ed. 2,
5: 201, nom. cons.; Gagnep., 1934, Fl. Gen. Indo-
27
Turczaninowia 2011, 14(2) : 15–100
Fig. 7. Aphyllorchis annamensis (VH 2221, type): a – owering plant, b – attened sepals and petals, c – lip,
side view, d – column, side view, e – column apex, frontal view, f – operculum, view from below.
28 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Chine 6, 4: 575-576; Seidenf., 1992, Opera Bot.
114: 26; Aver., 1994, Ident. Guide Vietnam. Orch.:
26–27; P.H. Ho, 2000, Ill. Fl. Vietnam 3: 783; Su
Horng-Jye, 2000, Fl. Taiwan 5: 952–959; Pearce et
Cribb, 2002, Orch. Bhutan: 47–53; Aver. et Averya-
nova, 2003, Updated Checklist Orch. Viet.: 43. –
Neottia Guett., 1754, Hist. Acad. Roy. Sci. Mem.
Math. Phys. (Paris, 4°) 1750: 374, nom. cons., p.p.:
Chen Sing-chi, Gale, Cribb 2009, Fl. China, 25:
184–195.
Type: L. ovata (L.) R. Br. (Ophrys ovata L.).
Terrestrial sympodial herbs with thin under-
ground rhizome, 2-leaved slender stem and terminal
few- to many-owered inorescence. Leaves sub op-
posite at the middle of stem, sessile, plicate. Flowers
small, resupinate. Sepals and petals free. Lip with-
out spur, not mobile, usually 2-lobed. Column short.
Pollinia 2, powdery, each with small viscidium.
20(1) species. Eurasia, N. America.
L. latilabra Evrard ex Gagnep.,
1931, Bull. Mus. Hist. Nat. (Paris) 2 ser. 3, 7:
683; Gagnep., 1934, Fl. Gen. Indo-Chine 6, 4: 576,
g. 15, 16; Seidenf., 1992, Orch. Indochina: 26, g.
8; Aver., 1994, Identif. Guide. Vietnam. Orch.: 26;
P.H. Ho, 2000, Ill. Fl. Vietnam 3: 783, g. 10896;
Aver. et Averyanova, 2003, Updated Checklist Orch.
Viet.: 43.
Described from S. Vietnam (“Annam: Da-
lat”). Type (“Evrard, N 1251”) – P.
Stem slender, erect, 10–25 cm tall. Leaves 2
(rarely 3), triangular-cordate, 2–4 cm long and wide,
acute, nely undulate along margin. Inorescence
few owered, 2–10 cm long. Floral bracts cuneate,
1–1.5 mm long. Pedicel and ovary 5–12 mm long,
sparsely pubescent. Flowers 3–8, dull purple-green,
6–8 mm across. Sepals and petals 3 mm long; se-
pals narrowly ovate, acute; petals narrowly lanceo-
late, obtuse. Lip 7 mm long, oblong-pandurate, with
semicircular auricles at the base, fat green midvein
and 2 triangular-falcate acute lobules at the apex.
Column very short, knob-like. Fruit ovate capsule
5–6 mm long. Fig. 8; 23 d.
Ecology. Wet mossy spring-water places
along streams in broadleaved and mixed forests on
silicate soils. 1000–1500 m. Fl. September – Octo-
ber. Very rare (CR).
Distribution. Vietnam (Lam Dong, Thua
Thien-Hue). Endemic.
Studied specimens. Dalat, Evrard 1371 (P);
Thua Thien-Hue, Bach Ma national park, HLF 1298,
HLF 1355 (HN, LE).
Trib. 5.2. Vanilleae Blume,
1835, Rumphia 1: 196.
Subtrib. 5.2.1. Galeolinae Garay,
1986, Bot. Mus. Lea. Harv. Univ. 30, 4: 233.
Type: Galeola Lour.
3(3) genera and 30(8) species. Tropical and
subtropical regions of the World.
Erythrorchis Blume,
1837, Rumphia, 1: 200, t. 70; Comber, 1990,
Orch. Java: 72–73; id., 2001, Orch. Sumatra: 127–
128; Seidenf., 1992, Opera Bot. 114: 73; Seidenf.,
Wood, 1992, Orch. Malay. Sing.: 130–132; Aver.,
1994, Ident. Guide Vietnam. Orch.: 79; P.H. Ho,
2000, Ill. Fl. Vietnam 3: 792; Su Horng-Jye, 2000,
Fl. Taiwan 5: 872–874; Aver. et Averyanova, 2003,
Updated Checklist Orch. Viet.: 35; Newman et al,
2007, Checkl. Vasc. Pl. Lao PDR: 268; Schuiteman
et al., 2008, Nord. Journ. Bot. 26: 292; Chen Sing-
chi, Gale, Cribb, 2009, Fl. China, 25: 171. – Galeola
Lour., p.p.: Gagnep., 1934, Fl. Gen. Indo-Chine 6,
5: 630–636.
Type: E. altissima (Blume) Blume (Cyrto-
sia altissima Blume).
Achlorophyllous leaess vines. Stem clim-
bing, dull reddish-brown or yellowish-brown, cy-
lind ric, exuous, much branched, glabrous, with
roots and scales at nodes. Inorescence terminal or
lateral raceme or panicle, dense, many-owered;
Fig. 8. Listera latilabra (Eberhard, 1257, type):
ower, frontal view.
29
Turczaninowia 2011, 14(2) : 15–100
rachis and ower glabrous; oral bracts small,
persistent. Flowers resupinate, not fully open-
ing. Sepals and petals often connivent; lip nearly
unlobed, broad, with stout and thick longitudinal
median ridge and apical papillose cushion. Col-
umn erect, slightly curved, with very short foot,
tapering into median ridge of lip; anther subtermi-
nal; pollinia 2, granular-mealy, without caudicle
or viscidium; stigma concave, large; rostellum
small. Fruit dry, dehiscent, long cylindric cap-
sule. Seeds with stout testa and broad surround-
ing wing.
3(1) species. Tropical Asia to islands of
W. Pacic.
E. altissima (Blume) Blume,
1837, Rumphia, 1: 200; Gagnep., 1934, Fl.
Gen. Indochine, 6, 5: 632, g. 61, 2–8; Comber,
1990, Orch. Java: 73, g.; id., 2001, Orch. Sumatra:
128, g.; Seidenf., Wood, 1992, Orch. Malay. Sing.:
132, g. 54 a, b; Su Horng-Jye, 2000, Fl. Taiwan
5: 873, g. 371. – Cyrtosia altissima Blume, 1825,
Bijdr.: 396. – Galeola altissima (Blume) Reichenb.
f., 1865, Xenia Orch. 2: 77; Seidenf., 1978, Dansk
Bot. Ark. 32, 2: 137, g. 85; Aver., 1988, Prelim.
List Vietnam. Orch. 1: 198; Gagnep., 1934, Fl. Gen.
Indo-Chine 6, 5: 635, g. 61, 2–8. – G. ochobiensis
Hayata, 1916, Icon. Pl. Formos. 6: 87. – Erythrorchis
ochobiensis (Hayata) Garay, 1986, Bot. Mus. Lea.
Harv. Univ. 30, 4: 234; Seidenf., 1992, Opera Bot.
114: 73; Seidenf., Wood, 1992, Orch. Malay. Sing.:
132, g. 54 c-e; Aver., 1994, Ident. Guide Vietnam.
Orch.: 79; P.H. Ho, 2000, Ill. Fl. Vietnam 3: 792, g.
10930; Aver., Averyanova, 2003, Updated Checkl.
Orch. Vietnam: 35.
Described from Java (“in montibus Seribu”).
Type (“Blume”) – L.
Stem yellowish-pink, reddish-brown or
olive-brown, climbing, cylindric, slender, to 10
m long, with many curved and twisted branches,
with short scales at slightly inated nodes. In-
orescence large, many-owered raceme, with
branched slender, glabrous rachis. Floral bracts
persistent, triangular, 2–3 mm long. Pedicel and
ovary erect, glabrous, 6–10 mm long. Flowers
whitish-yellow, slightly tinged with brown, of-
ten not opening widely. Sepals narrowly obovate,
10–14 mm long 3–5 mm wide, glabrous. Petals
oblanceolate, slightly narrower than sepals. Lip
slightly tinged with violet or brown, broadly obo-
vate, concave, as long as sepals, apex indistinctly
3-lobed, irregularly undulate and plicate; disc
with stout thick median hairy longitudinal ridge
from base to middle, at the apex with papillose
cushion; surface of lip side lobes with numerous
ne indistinct transversal folds. Column erect, 5–7
mm tall. Capsule dull reddish-brown to gray, nar-
rowly cylindric, 10–22 cm long, 5–10 mm wide.
Seeds with large, at, surrounding wing about 1
mm wide, wing cleft on one side. Fig. 9; 23 e-g.
Ecology. Primary and secondary broad-
leaved, evergreen, shady forests on soils derived
from silicate rocks at elev. 50–1000 m a.s.l. Fl.
April – May. Rare (EN).
Distribution. Vietnam (Danang, Dong Nai,
Hai Phong, Hanoi, Kien Giang, Ninh Thuan, Phu
Khanh, Quang Binh, Quang Nam, Quang Tri, Thua
Thien-Hue). NE. India, Myanmar, Japan, Taiwan,
Hainan, Thailand, Laos, Cambodia, Malaysia, Indo-
nesia, Philippines.
Studied specimens. Hanoi, Bavi, Balansa
2018 (LE, P); Dong Nai, Muxohay, Pierre 61 (P);
Hai Phong, Cat Ba, Nong Van Tiep NVT 3089,
(LE, UHN); Kien Giang, Phu Quoc, Godefroy,
961 (P); Phu Khanh, Nhatrang, Poilane 6247 (K,
P), Poilane 6414 (P); Quangnam-Danang, Phuoc
Son, Lang Ha, LX-VN s.n. 16.07.1986 (LE),
Quang Binh, Le Thuy, HAL 11440 (HN, LE);
Quang Binh, Quang Ninh, HAL 11571 (HN, LE);
Quang Tri, Da Krong, Da Krong Nature Reserve
HLF 6214 (HN, LE), d-EXSICCATES OF VIET-
NAMESE FLORA 0062/HLF 6214; Ninh Thuan,
Ninh Hai, Nui Chua national park HLF 4416 (HN,
LE), HLF 4603 (HN, LE).
Notes. Northern race of this species
distribu ted in Japan, Taiwan and Hainan some-
times treated as a distinct species – Erythrorchis
ochobiensis with very narrow, liform fruits.
Vietnamese specimens have more or less interme-
diate fruit morphology.
Cyrtosia Blume,
1825, Bijdr. 6, g. 6; id., 1825, ibid., 8: 396;
Comber, 1990, Orch. Java: 72; id., Orch. Sumatra:
126–127; Seidenf., 1992, Opera Bot. 114: 73; Se-
idenf., Wood, 1992, Orch. Malay. Sing.: 127–128;
Aver., 1994, Ident. Guide Vietnam. Orch.: 77; P.H.
Ho, 2000, Ill. Fl. Vietnam 3: 793; Su Horng-Jye,
2000, Fl. Taiwan 5: 835-839; Aver. et Averyanova,
2003, Updated Checklist Orch. Viet.: 25; Newman
et al, 2007, Checkl. Vasc. Pl. Lao PDR: 261; Schu-
iteman et al., 2008, Nord. Journ. Bot. 26: 279; Chen
Sing-chi, Cribb, 2009, Fl. China, 25: 168–170. –
Galeola Lour., p.p.: Gagnep., 1934, Fl. Gen. Indo-
Chine 6, 5: 630–636.
Lectotype: C. javanica Blume.
30 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 9. Erythrorchis altissima: a – owering plant (HAL 11571), b – ower, c – attened sepals, petals and lip,
d – clolumn, side view (HLF 4603), e – fruits and portion of fruiting stem (Averyanov, LX-VN s.n., 1986).
31
Turczaninowia 2011, 14(2) : 15–100
Achlorophyllous leaess terrestrial herbs
with erect stem, stout rhizome bearing woody or
eshy, tuber-like roots. Stem simple or branched,
dull pink to yellowish-brown, eshy, with scales at
nodes. Inorescence terminal or lateral raceme or
panicle, few- to many-owered, shortly hairy, o-
ral bracts persistent. Flowers commonly not widely
opening, campanulate, or subcampanulate. Sepals
and petals connivent or not; sepals often more or
less hairy outside; petals glabrous. Lip unlobed,
spurless, lip base embracing column. Column foot-
less slightly curved or straight, stout, broadening to
the apex; anther terminal; pollinia 2, granular-mealy,
without caudicle or viscidium. Fruit eshy cylindri-
cal indehiscent berry. Seeds with stout testa, wing-
less or with a narrow surrounding wing.
10(6) species. Mainland tropical and sub-
tropical Asia, Japan, Taiwan, Hainan, Indonesia.
Notes. The genus closely allies to Galeola
Lour., which species differ in climbing vine habit,
aerial roots at nodes and drying dehiscent fruit that
is rather capsule than berry. All members of the
genus have ephemeral inorescences and owers,
hence they easily overlooked in eld surveys and
collecting.
Key to species
1. Stem commonly much shorter than 1.2 m, less than 5 mm in diam. at the base; inorescence simple or hardly
branching; rachis less than 15 cm long, lateral racemes less than 2(3) cm long; owers less than 2 cm across
......................................................................................................................................................................... 2
- Stem 1.5–3.5 m tall, more than 5 mm in diam. at the base; inorescence much branching; rachis much longer
than 15 cm long, lateral racemes usually longer than 2 cm; owers larger than 2 cm across ........................ 4
2. Stem (15)20–120 cm tall; owers more or less widely opening, sepals and petals more or less thin, recurved;
lip vey eshy, with 2 large massive glabrous callosities at the center; column with large broad, ear-like, lateral
wings at apex ................................................................................................................................ 1. C. integra
- Stem to 15(20) cm tall; owers hardly opening, campanulate, sepals eshy, not recurved; lip not too much
eshy, with no callosity inside, or with 2 small nger-like erected glabrous processes at the center; column
with no large prominent wings at apex ........................................................................................................... 3
3. Inorescence branching; lip hairy inside and nely ciliate along rather thin apical margin; disc with 2 small
nger-like erected glabrous processes ............................................................................................. 2. C. nana
- Inorescence not branching; lip glabrous or slightly mealy pubescent to the apex, not ciliate along eshy
apical rim; disc with no callosities ............................................................................................. 3. C. javanica
4. Lip oblong, hardly concave, not cup-like; disc with strong nerves cowered with warty, lamellate or crenate
appendages; oral bracts outside sparsely hairy or glabrous; column 8–10 mm tall ..................... 4. C. faberi
- Lip almost round, strongly concave, cup-like; disc densely haired with long papillae; oral bracts outside
densely hairy; column 2–3 mm tall ................................................................................................................ 5
5. Inorescence narrow, usually composed with unbranched racemes less than 3.5 cm long; sepals less than
2 cm long, with distinct massive wavy keel or roughly grooved outside; lip with small erect glabrous plate
near the base, without constriction or fold at the base; anther strongly papillose .................. 5. C. lindleyana
- Inorescence broad, many branched, with branches to 35 cm long; sepals 2.5–3 cm long, outside more or less
smooth, without distinct keel; lip with small transversal fold forming a sac at the base, distinctly constricted
between the sac and lip blade; anther short papillose or nearly glabrous ................................. 6. C. falconeri
1. C. integra (Rolfe ex Downie) Garay,
1986, Bot. Mus. Lea. 30, 4: 232; Seidenf.,
1992, Orch. Indochina: 73; Aver. et Averyanova,
2003, Updated Checklist Orch. Viet.: 25; Newman
et al., 2007, Checkl. Vasc. Pl. Lao PDR: 261. – Ga-
leola integra Rolfe ex Downie, 1925, Kew Bull.
1925: 409; Seidenf., 1978, Dansk Bot. Ark. 32, 2:
130, 131, g. 80; P.H. Ho, 2000, Ill. Fl. Vietnam 3:
793, g. 10931.
Described from NW. Thailand (“Doi Suthep
900 m”). Type (“Kerr 304”) – C, K.
Stems dull red, pink-brown to yellowish-
brown, to 1(1.2) m tall, arising by 1–4 from short,
rigid vertical rhizome, clustered with eshy, cylin-
dric or clavate roots. Inorescence branching or not,
each raceme few- to many-owered. Floral bracts
triangular, acute, persistent, to 3 mm long. Pedicel
and ovary pinkish, 1–1.5 cm long, mealy-pubescent.
Flowers broadly opening, sepals and petals subsimi-
lar, 1.2–1.8 cm long, median sepal and petals brown,
brown-yellow to olive-green, lateral sepals broader,
in halves brown and yellow. Sepals mealy pubes-
cent outside; dorsal sepal, narrowly ovate, 4–5 mm
wide, concave; lateral sepals, ovate, 5–8 mm wide.
Petals narrowly ovate, 4–5 mm wide. Lip yellow,
often with whitish center, eshy, concave, subor-
32 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
bicular, hairy inside, particularly to the apex, with
2 low thick, glabrous keels at the center. Column
white, slightly curved, broadening to apex, 6–8 mm
tall, at apex with broad, ear-like, lateral wings ne-
ly denticulate along margin. Fruits cylindric, light
brown-yellow to brown-purple, to 7 cm long. Fig.
10; 23 h, i.
Ecology. Primary and secondary broadlea-
ved and mixed evergreen rather open forests (often
with bamboo) on any kind of soils at elev. 800–1500
m a.s.l. Fl. April – May (June). Very rare (VU).
Distribution. Vietnam (Bac Kan, Ha Giang,
Thua Thien-Hue). Thailand, Laos.
Studied specimens. Bac Kan, Cho Don,
HAL 4874 (HN, LE); Ha Giang, Quan Ba, HAL
1510 (HN, LE); Thua Thien-Hue, Phu Loc, Bach
Ma national park, HLF 934 (HN, LE).
2. C. nana (Rolfe ex Downie) Garay,
1986, Bot. Mus. Lea. 30: 233. – Galeola
nana Rolfe ex Downie, 1925, Kew Bull. 1925: 409;
Seidenf. 1978, Dansk Bot. Ark. 32, 2: 132, g. 81.
Described from NW. Thailand (“Doi Su-
thep”). Type (“Kerr 313”) – K.
Stem eshy, erect, white with yellow or red-
dish tint, to 15 cm tall, glabrous or apically with
sparse, mealy hairs; nodes with lanceolate scales;
internodes usually 1–3 cm. Rhizome slender, rigid,
woody, to 10 cm long, fascicled with numerous tu-
ber-like, short, eshy, clavate roots, 4–9 cm long and
0.5–1.2 cm in diam. Inorescence erect, rigid, on
stalk 2–5 cm, elongating slowly and producing ow-
ers in succession, branching or not; raceme few- to
many-owered, rusty mealy-pubescent; oral bracts
persistent, triangular, acute, 1–2 mm long, rusty pu-
bescent outside. Pedicel and ovary 8–10 mm long,
rusty haired. Flowers hardly opening, pale yellow,
lip with indistinct orange-red longitudinal stripes.
Sepals eshy, elliptic, 1–1.8 cm long, 0.9 cm wide,
concave, outside mealy pubescent. Petals, slight-
ly narrower, thin. Lip concave, cup-like, broadly
ovate, entire, 1–1.4 cm long, hairy inside, margin
sometime nely undulate and erose-ciliate, disk at
the center with 2 small pyramidal glabrous nger-
like erected glabrous processes. Column slender,
to 7 mm tall, slightly dilated to the apex, without
conspicuous wing. Fruit oblong-elliptic to cylindric
pink-purple berry, 3–6 cm long, 0.8–1 cm in diam.
Fig. 11; 24 a, b.
Ecology. Primary and secondary evergreen
broadleaved shady lowland valley forest in lime-
stone areas at elev. about 250 m a.s.l. Fl. April –
June. Very rare (VU).
Distribution. Vietnam (Nghe An, Ninh Binh,
Thanh Hoa, Quang Binh). Thailand, SW. China.
Studied collections. Ninh Binh and Thanh
Hoa, Cuc Phuong national park, Dinh kinh 181–264;
CPNP 5147 (Cuc Phuong national park Herbarium);
N.M. Cuong et al., NMC 1665 (Cuc Phuong nation-
al park Herbarium, LE); Thanh Hoa, Thach Thanh,
HAL 2896 (LE); Nghe An, Tuong Duong, HLF 6674
(HN, LE), HLF 7055 (HN, LE); Quang Binh, Minh
Hoa, VH 4677a (LE).
Notes. Very rare relictual element of prima-
ry warm-lowing lowland limestone tropical forests.
In northwestern Thailand and southwestern China it
was reported from elevations 500–1400 m a.s.l.
3. C. javanica Blume,
1825, Bijdr. 6, g. 6; id., 1825, ibid., 8: 396;
Comber, 1990, Orch. Java: 73, g.; id., 2001, Orch.
Sumatra: 126, g.; Seidenf., 1992, Orch. Indochina:
73; Seidenf., Wood, 1992, Orch. Malay. Sing.: 128,
g. 52; Aver., 1994, Ident. Guide Vietnam. Orch.:
77; P.H. Ho, 2000, Ill. Fl. Vietnam 3: 793, g.
10933; Su Horng-Jye, 2000, Fl. Taiwan 5: 838, g.
354; Aver. et Averyanova, 2003, Updated Checklist
Orch. Viet.: 25; Shih-Wen Chung, 2008, Orch. Tai-
wan, 1: 121, g. – Galeola javanica (Blume) Benth.
et Hook.f., 1883, Cen. Pl. 3: 590; J.J. Smith, 1908,
Fl. Buitenzorg 6, 2, g. 46; Seidenf., 1978, Dansk
Bot. Ark. 32, 2: 130, g. 79; Jayaweera, 1981, Fl.
Ceylon 2: 339, g. 149; Aver., 1988, Prelim. List
Vietnam. Orch. 1: 199.
Described from Java (“montium Javae in-
sulae”). Syntypes (Herb. numbers 9023222253,
9023222254, 9023222256) – L.
Stems yellowish or pink-yellow, to 20 cm
tall, often arising by 1–3 from short, rigid vertical
rhizome 6–8 cm long, covered with more or less im-
bricate yellowish-brown persistent glabrous scales.
Roots numerous, eshy, tuber-like, cylindric or clav-
ate, 5–8 cm long, 8–12 mm in diam. Inorescence
racemose, few- to many-owered. Floral bracts tri-
angular, small. Pedicel and ovary 1–1.5 cm long,
mealy pubescent. Flowers not fully opening, yel-
lowish to yellow-orange. Sepals subsimilar, eshy,
mealy pubescent outside, narrowly ovate, 1–1.4 cm
long, 4–6 mm wide, concave. Petals thin, narrowly
ovate, as long as sepals. Lip, 8–12 mm long, yel-
lowish at the base, whitish to the apex, suborbicular,
with thickened apex, glabrous or slightly mealy pu-
bescent in apical part. Column clavate, to 6 mm tall,
with erose-dentate wings at apex. Fruits cylindric,
dull pinkish-brown, to 6 cm long. Fig. 12.
33
Turczaninowia 2011, 14(2) : 15–100
Fig. 10. Cyrtosia integra: a – owering and fruiting plant (HAL 7874), b – ower, c – attened sepals and pe-
tals, d – column and reexed, attened lip, frontal view (HAL 1510).
34 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 11. Cyrtosia nana: a – fruiting plant (HAL 2896), b – attened sepals and petals, c – attened lip, d – co-
lumn, ventral and dorsal view, e – column apex without operculum (NMC 1665).
35
Turczaninowia 2011, 14(2) : 15–100
Ecology. Primary and secondary old broad-
leaved evergreen shady forests (sometimes with
bamboo) on any kind of soils at elev. 300–1500 m
a.s.l. Fl. March – May (June). Very rare (VU).
Distribution. Vietnam (Cao Bang, На Noi,
Hoa Binh, Quang Binh). Sri Lanka, NE. India, Thai-
land, Taiwan, Philippines, Malacca Peninsula, Java,
Sumatra, Kalimantan.
Studied specimens. Hanoi, Mt. Bavi,
Baslansa 2015 (P); Cao Bang, Tra Linh, NTH 2111a
(HN, LE); Hoa Binh, Mai Chau, NTH 2111 (HN);
Quang Binh, Quang Ninh, HAL 11515 (HN).
Notes. Forms with several wings at each lat-
eral side of column apex were described from Thai-
land as Cyrtosia plurialata Seidenf. (1995, Opera
Bot., 124: 13). Such plants closely ally to variable
C. javanica and may represent its marginal form.
4. C. faberi (Rolfe) Aver.,
comb. nov. – Galeola faberi Rolfe, 1896, Kew
Bull. 1896: 200; Chen Sing-chi, Tsi Zhanhuo, Luo Yibo,
1990, Nat. Orch. China: 226, g.; Jin Xiaohua, Zhao
Xiaodong, Shi Xiaochun, 2000, Nat Orch. Gaoligong-
shan Mount.: 251, g. – G. shweliensis W.W. Smith,
1921, Not. Roy. Bot. Gard. Edinb. 13: 204.
Described from S. China (“China: Sichuan:
Mt. Omei, 7000 ft.”). Type (“Faber s.n.”) – K.
Stem dull reddish-brown, to 3 m tall, in
lower part glabrous, upper part hardly sparsely rusty
hairy, with several distant ovate to lanceolate scales
2–4 cm long. Rhizome 2–3 cm in diam., covered
by broad triangular scales. Panicle composed of ter-
minal and lateral often branching racemes 5–25 cm
long, many-owered; peduncle and rachis sparsely
shortly tomentose; sterile bracts at base of racemes
narrowly ovate, 1–2 cm long, glabrous; oral bracts
ovate-triangular, 1–4 mm long, often vertical to ra-
chis, dorsally glabrous or hardly sparsely rusty to-
mentose. Pedicel and ovary 1–2 cm long, rusty to-
mentose. Flowers dull pale yellow, 2.5–3.5 cm in
diam., lip sometimes laterally with reddish stripes.
Sepals similar, narrowly elliptic to broadly lanceo-
late, 2–3 cm long, 5–8 mm wide, sparsely very short-
ly rusty tomentose outside. Petals oblong, as long as
sepals, 6–10 mm wide, nely irregularly denticulate
along margin. Lip entire, obovate to oblong, 1.6–2.2
cm long, 1–1.2 cm wide, inside with many thick
veins densely covered with warty, lamellate or cren-
ate appendages, basal part concave, loosely embracing
column, margin nely irregularly incised and undulate.
Column clavate, erect, nearly straight, 8–10 mm tall;
anther cap nely papillose. Fruit red to red-brownish,
cylindric berry. Fig. 13.
Ecology. Humid primary and secondary
broad-leaved evergreen rather open forests (some-
time with bamboo) on silicate soils commonly in
small depressions rich in humus accumulations at
elev. 1900–2000 m a.s.l. Fl. May – June. Fruits Sep-
tember – October. Very rare (VU).
Distribution. Vietnam (Lao Cai). Nepal,
Bhutan, N. India, China, Sumatra.
Studied collections. Lao Cai, Sa Pa, DKH
5992 (HN, LE, MO), NTH 2650 (HN, LE).
Notes. This species as well as C. lindleya-
na and C. falconery are spectacular giant highland
plants with unforgettable handsome appearance and
very large, banana-like, eshy and juicy, brightly
red or purple fruits. Unfortunately, owering period
of these species is fairly ephemeral, hence they are
poorly presented in botanical collections.
5. C. lindleyana Hook. f. et J. Thomson,
1855, Ill. Himal. Pl., tab. 22. – Galeola lind-
leyana (Hook. f. et J. Thomson) Rchb.f., 1865, Xe-
nia Orchid. 2: 78; King et Pantl., 1898, Ann. Roy.
Bot. Gard. Calcutta, 8: 264, tab. 352; Chen Sing-
chi, Tsi Zhanhuo, Luo Yibo, 1990, Nat. Orch. China:
227, g.; Jin Xiaohua, Zhao Xiaodong, Shi Xiaoc-
hun, 2000, Nat Orch. Gaoligongshan Mount.: 252,
g.; Comber, 2001, Orch. Sumatra: 129, g.; Pearce
et Cribb, 2002, Orch. Bhutan: 66, g. 14, a-o, pl. 4;
Shih-Wen Chung, 2008, Orch. Taiwan, 1: 168, g. –
Galeola matsudai Hayata, 1920, Icon. Pl. Formos.
9: 114. – Galeola kwangsiensis Hand.-Mazz., 1936,
Sinensia, 7: 620.
Described from NE. India (“India, Khasia”).
Type (“Hooker 357”) – K (holotype).
Stem usually not branched, dull reddish-
brown, 2–4 m tall, rusty hairy to glabrous with age,
nodes with broadly ovate spaced scales. Rhizome
horizontal, woody, 2–3 cm in diam., with sparse
broad ovate scales at nodes. Panicle composed of
terminal and lateral racemes; lateral raceme com-
monly less than 3.5 cm long, few- to 10-owered,
usually very shortly pedunculate. Sterile bracts at
base of raceme triangular to broadly ovate, 1–2.5 cm,
densely rusty pubescent. Floral bracts broadly ovate,
5–10 mm long, outside densely rusty pubescent.
Pedicel and ovary 1–2 cm long, densely rusty pu-
bescent. Flowers brightly yellow, whitish-furry out-
side, 2.5–3.5 cm in diam., lip sometimes with red
hairs. Sepals subsimilar, ovate to elliptic, 1.4–2 cm
long, 9–11 mm wide, outside densely rusty tomen-
tose, median sepal outside roughly grooved, lateral
sepals with massive keel. Petals broadly obovate to
suborbicular, as long as sepals, 1–1.4 cm wide, erose
36 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 12. Cyrtosia javanica (NTH 8111a): a – owering plant, b – attened sepals and petals, c – lip, d – column,
frontal view.
37
Turczaninowia 2011, 14(2) : 15–100
Fig. 13. Cyrtosia faberi (NTH 2650): a – owering plant, b – inorescence branch, c – attened sepals and
petals, d – attened lip, e – ovary and column, side view.
38 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
or nely toothed along margin. Lip entire, concave,
cup-shaped, broadly ovate or orbicular, about 1.2 cm
in diam., densely papillose, shortly mbriate along
margin, with small glabrous vertical plate near base.
Column yellow, stout, erect, straight, 2–3 mm tall,
at the base with 2 tufts of long papillae; anther cap
reddish-orange, haired with long papillae. Fruit pale
brown to red, cylindric to subtrigonal in section,
8–18 cm long, 1.7–2.4 cm in diam. Seeds brown,
1–1.5 mm in diam., with narrow encircled wing.
Fig. 14.
Ecology. Humid primary and secondary bro-
ad-leaved evergreen rather open forests (sometime
with bamboo) on silicate soils commonly in small
depressions rich in humus accumulations at elev.
1000–2200 m a.s.l. Fl. May – June. Very rare (VU).
Distribution. Vietnam (Ha Giang, Lao Cai).
Nepal, Bhutan, NE. India, China, Taiwan, Sumatra.
Studied collections. Ha Giang, Yen Minh,
CBL 2090 (HN, LE); Lao Cai, Than Uyen, NTH
2867 (HN, LE).
Notes. All available specimens from Viet-
nam differ from the type and other Himalayan plants
in sepals that have distinctly incise to irregularly
denticulate margin. In this connection, Vietnamese
(as well as, probably, Chinese) plants certainly rep-
resent taxonomically different eastern race of the
species.
6. C. falconeri (Hook. f.) Aver.,
comb. nov. – Galeola falconeri Hook. f.,
1890, Fl. Brit. India 6: 88; King et Pantl., 1898,
Ann. Roy. Bot. Gard. Calcutta, 8: 265, tab. 353; Su
Horng-Jye, 2000, Fl. Taiwan 5: 883, g. 376; Pearce
et Cribb, 2002, Orch. Bhutan: 64, g. 14, p-t, pl. 4;
Shih-Wen Chung, 2008, Orch. Taiwan, 1: 167, g. –
G. kuhlii auct. non (Reichenb. f.) Reichenb. f.: Liu
et Su, 1978, Fl. Taiwan, 5: 996.
Described from NE. India (“India, Garh-
wal”, “India, Sikkim”). Syntypes (“icon. Falconer
s.n.”, “Thomson s.n.”) – K.
Stem dull brownish to reddish, to 3.5 m tall,
in lower part almost glabrous, upper part sparsely
and shortly rusty hairy, with several distant ovate
or lanceolate scales 2–4 cm long. Rhizome 3–5 cm
in diam., covered by large, broad, triangular scales.
Panicle composed of terminal and lateral often
branching racemes; racemes 5–35 cm; peduncle and
rachis shortly tomentose. Floral bracts triangular,
1–3 mm, often nearly at right angle to rachis, out-
side rusty tomentose. Pedicel and ovary 1.5–3 cm
long, densely rusty tomentose. Flowers bright yel-
low, 4–5 cm in diam. Sepals elliptic-oblong, 2.2–
3 cm long, 1–1.5 cm wide, densely rusty tomen-
tose and smooth outside. Petals as long as sepals,
slightly narrower. Lip entire, concave, cup-shaped,
broadly ovate or orbicular, about 2 cm in diam.,
inside densely papillose, margin nely mbriate,
basal part loosely embracing column, near base with
transversal fold forming small sac. Column stout,
erect, slightly curved forward, 2–3 mm tall, at the
base with 2 tufts of long papillae; anther cap papil-
lose or nearly glabrous. Fruit red to purple, oblong,
cylindric, (18)20–25 cm long, to 3 cm in diam., with
nely verruculose surface. Seeds dark brown, 1.5–
2 mm in diam., with narrow encircled wing. Fig. 15;
24 c, d.
Ecology. Humid primary and secondary
broad-leaved evergreen rather open forests (some-
time with bamboo) on silicate soils commonly in
small depressions rich in humus accumulations at elev.
1500–2200 m a.s.l. Fl. May – June. Very rare (VU).
Distribution. Vietnam (Son La, Lai Chau,
Lao Cai). Bhutan, NE. India, China, Thailand.
Studied collections. Son La, Thuan Chau,
HAL 9653 (HN, LE); Lai Chau, Tam Duong, HAL
10260 (HN); Lao Cai, Van Ban, HAL 2133 (HN), HAL
2467 (HN, LE); Lao Cai, Sa Pa, s.n. (photo – LE).
Notes. Chen Sing-chi and Phillip Cribb
consider specimens from China in their treatment
for “Flora of China” (2009) as different from Hi-
malayan plants, which have slightly 3-lobed lip and
petals denticulate along margins. Vietnamese plants
may belong to the same – “Chinese” form.
Galeola Lour.,
1790, Fl. Cochinchin.: 520; Gagnep., 1934,
Fl. Gen. Indo-Chine 6, 5: 630–636; Seidenf., 1978,
Dansk Bot. Ark. 32, 2: 129–137; id., 1992, Opera
Bot. 114: 73–74; Comber, 1990, Orch. Java: 73; id.,
2001, Orch. Sumatra: 128–130; Seidenf., Wood,
1992, Orch. Malay. Sing.: 129–130; Aver., 1994,
Ident. Guide Vietnam. Orch.: 78; P.H. Ho, 2000, Ill.
Fl. Vietnam 3: 793; Su Horng-Jye, 2000, Taiwania,
45,3: 242–246; id., 2000, Fl. Taiwan 5: 882-884;
Pearce et Cribb, 2002, Orch. Bhutan: 63–67; Aver.
et Averyanova, 2003, Updated Checklist Orch.
Viet.: 36; Chen Sing-chi, Cribb, 2009, Fl. China, 25:
168–169.
Lectotype: G. nudifolia Lour.
Achlorophyllous, reddish-brown or yellow-
brown terrestrial, eshy, leaess, climbing vines
with persistent scales and aerial roots at nodes and
rigid, woody rhizome. Raceme or panicle terminal
and lateral, with many owers. Rachis pubescent,
oral bracts persistent. Flowers not widely opening,
39
Turczaninowia 2011, 14(2) : 15–100
Fig. 14. Cyrtosia lindleyana: a – owering plant, b – portion of inorescence (CBL 2090), c – ower bud, d –
attened sepals and petals, e – attened and partially dissected lip, f – column, side view (HAL 2867).
40 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 15. Cyrtosia falconeri: a – owering plant, b – branchlet of inorescence, c – attened sepals and petals,
d – attened lip, e – column, side view (China s.n.), f – fruits, and apical portion of inorescence axis (HAL 9653).
41
Turczaninowia 2011, 14(2) : 15–100
campanulate, yellow, sometimes tinged with red.
Sepals and petals free, subsimilar. Sepals outside
hairy. Petals slightly smaller than sepals, glabrous.
Lip entire, concave, cup-shaped, embracing column,
disc with longitudinal ridge or callus. Column short
and stout, strongly forward curved, without column
foot. Anther very large. Pollinia 2, cleft, mealy,
without caudicle or viscidium. Stigma large, deeply
concave; rostellum short and wide. Fruit dry, pod-
like, dehiscent capsule. Seeds with stout testa and
broad surrounding wing.
5(1) species. Tropical and subtropical Asia
to New Guinea and N. Australia.
G. nudifolia Lour.,
1790, Fl. Cochinchin.: 521; Seidenf., 1978,
Dansk Bot. Ark. 32, 2: 135, g. 83; id., 1992, Opera
Bot. 114: 74; Seidenf., Wood, 1992, Orch. Malay.
Sing.: 129, g. 53; Aver., 1994, Ident. Guide Viet-
nam. Orch.: 78; P.H. Ho, 2000, Ill. Fl. Vietnam 3:
793, g. 10932; Aver. et Averyanova, 2003, Updated
Checkl. Orch. Vietnam: 36. – Galeola hydra Rei-
chenb. f., 1865, Xen. Orch. 2: 77; Gagnep., 1934,
Fl. Gen. Indo-Chine 6, 5: 634; King et Pantl., 1898,
Ann. Bоt. Gard. Calcutta, 8, tab. 351; J.J. Smith, 1908,
Fl. Buitenzorg 6, 2, g. 45. – Erythrorchis kuhlii Rei-
chenb. f., 1865, Xen. Orch. 2: 78, tab. 119. – Galeola
kuhlii (Reichenb. f.) Reichenb. f., 1865, Xen. Orch. 2:
78; P.H. Ho, 1972, Illustr. Fl. South Vietnam 2: 1013, g.
5014; Liu et Su, 1978, Fl. Taiwan 5: 997, g. 1596.
Described from S. Vietnam (“in sylvis Co-
chinchinae”). Type (“Loureiro”) – BM.
Climbing vine to 15 m long, stem 0.5–2 cm
in diameter, rooting at nodes, with thick, very rigid,
woody rhizome and triangular reddish scales 0.5–
6 cm long. Inorescence branching, with branches
15–50 cm long, laxly bearing many owers, ow-
ering in succession, 2–3 open in a time near grow-
ing tips, all young parts rusty hairy. Pedicel and
ovary 8–20 mm long. Floral bracts eshy, triangu-
lar, acute, 9–12 mm long. Flowers not open widely,
1.5–2.5 cm across, yellow, lip with orange-red veins
inside. Sepals and petals subsimilar, elliptic, 1.2–
2.2 cm long, 0.6–1.2 cm wide, blunt to obtuse, lat-
eral sepals broader and slightly oblique; petals more
thin in texture, often undulate along upper margin.
Lip almost round, entire, 1–1.6 cm across, strongly
concave, hairy, erose-mbriate along margin, with
somewhat pointed apex and erect short callus near
base. Column 4–5 mm tall, strongly curved forwards.
Fruit cylindric, dry, dehiscent capsule, to 15 cm long
and 2.5 cm in diam., seeds shortly winged, 2 mm in
diam. Fig. 16; 24 e, f.
Ecology. Primary and secondary broad-
leaved evergreen forests, often on steep slopes in
places of humus accumulations on any kind of soils,
but preferably in region with silicate subsoil rocks,
at elev. 100–800(1000) m a.s.l. Fl. March – July.
Rare (VU).
Distribution. Vietnam (Ha Tinh, Hoa Binh,
Ninh Binh, Quang Binh, Quang Tri, Thanh Hoa,
Thua Thien-Hue). Bhutan, Myanmar, NE. India, S.
China, Taiwan, Hainan, Thailand, Laos, Cambodia,
Vietnam, Malacca Peninsula, Indonesia, Philip-
pines.
Studied specimens. Ha Tinh, Huong Son,
HAL 5000 (HN, LE); Hoa Binh, Ha Son Binh, Lu-
ong Son, Lam Son, P-6193 (HNU, LE); Ninh Binh
and Thanh Hoa, Cuc Phuong national park, 7 June
1971, Vu Van Dung 2\48, CPNP 5115; sine date,
Dinh Kinh 181-26a; sine date, Vu van Dung 181-26
(Cuc Phuong national park Herbarium);Quang Binh,
Bo Trach, Phong Nha-Ke Bang national park, HAL
6240 (HN, LE); Quang Binh, Le Thuy, HAL 11439
(HN, LE); Quang Tri, Da Krong, Da Krong nature
reserve, HLF 5758 (HN, LE), HLF 6118 (HN, LE),
HLF 6213 (HN, LE); d-EXSICCATES OF VIET-
NAMESE FLORA 0060/HLF 6213; Quang Tri,
Huong Hoa, HLF 6534 (HN, LE); Thanh Hoa, Ba
Thuoc, HAL 954 (HN); Thua Thien-Hue, Huong
Thuy, HAL 8160 (HN, LE); Thua Thien-Hue, Nam
Dong, HAL 6792 (HN, LE).
Notes. Widespread, but rather uncommon
species of lowland, submontane and montane woods,
occurring sometime in quite open places in areas of
forest cutting, particularly in wet places among log-
ging decaying timber.
Closely related species – G. cathcartii Hook.
f. differs from G. nudifolia in narrow, lanceolate se-
pals and in cuneate base of the lip. This species re-
ported from NE. India and Thailand quite may be
found in eastern regions of Vietnam.
Subtrib. 5.2.2. Vanillinae Lindl.,
1840, Gen. Sp. Orch. Pl.: 429 (sub “Vanilleae”).
Type: Vanilla Mill.
6(2) genera and 110(5) species. Tropical re-
gions of the World.
Vanilla Plum. ex Mill.,
1754, Gard. Dict. Abr. 4, 3; Gagnep., 1934,
Fl. Gen. Indo-Chine, 6, 5: 580–583; Seidenf., 1978,
Dansk Bot. Ark. 32, 2: 138–146; id., 1992, Opera
Bot. 114: 71–72; Comber, 1990, Orch. Java: 74–76;
id., 2001, Orch. Sumatra: 130–132; Seidenf., Wood,
1992, Orch. Malay. Sing.: 124–127; Aver., 1994,
42 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 16. Galeola nudifolia (HLF 6213): a – owering plant, b – branchlet of inorescence, c – attened sepals
and petals, d – attened lip, e – column, side view.
43
Turczaninowia 2011, 14(2) : 15–100
Ident. Guide Vietnam. Orch.: 75–77; P.H. Ho, 2000,
Ill. Fl. Vietnam, 3: 791–792; Su Horng-Jye, 2000,
Fl. Taiwan 5: 1063; Aver. et Averyanova, 2003,
Updated Checklist Orch. Viet.: 60; Newman et al.,
2007, Checkl. Vasc. Pl. Lao PDR: 283; Schuiteman
et al., 2008, Nord. Journ. Bot. 26: 314–315; Chen
Sing-chi, Cribb, 2009, Fl. China, 25: 167–168; Soto,
Cribb, 2010, Lankesteriana 9, 3: 355–398.
Lectotype: V. mexicana Mill.
Climbing epiphytic or lithophytic herba-
ceous succulent vines to 20 m long. Stem eshy,
green, with succulent leaves or leaess, with aerial
roots at nodes. Leaves sessile or shortly petiolate,
sometimes reduced to dull brownish scales. Ino-
rescence – axillary raceme with few to many spi-
rally arranged owers. Scape naked or with few spi-
rally arranged, triangular or leaf-like bracts. Floral
bracts small, commonly broadly triangular, obtuse,
much shorter than ovary. Flowers resupinate, widely
opening, often with abscission layer between ovary
and perianth. Sepals and petals similar, free, spread-
ing. Lip spurless, joined to lateral sides of column,
sometimes almost to top of column, more or less
trumpet-shaped, free apical part broadening, en-
tire or 3-lobed, disk often with scarious, papillate
or hairy appendages. Column long, slender, slightly
curved. Anther apical, pointing downward; pollinia
2 or 4, mealy or granular, without caudicle or viscid-
ium; rostellum often broad. Fruit cylindric, eshy,
often indehiscent. Seeds with stout testa, brown to
black, wingless.
110(3) species. Tropical regions of the World.
Note. Taxonomy of the genus is very dif-
cult and complicated because of all its species be-
ing rarely found in owers. In this treatment, I fol-
low mainly the concept of Soto Arenas and Cribb
(2010).
Key to species
1. Stem leaess, or with small scales at nodes ................................................................................. 1. V. aphylla
- Stem with normal, broad, green leaves ........................................................................................................... 2
2. Inorescence 1–3 cm long, few-owered; lip glabrous or with few small scattered papillae at apex; sepals and
lip 4–5 cm long ........................................................................................................................ 2. V. yersiniana
- Inorescence 3–14 cm long, many-owered; lip with mbriate papillae near apex; sepals and lip 3.8–4.5 cm
long ........................................................................................................................................... 3. V. siamensis
1. V. aphylla Blume,
1825, Bijdr. Fl. Ned. Ind. 8: 422; J.J. Smith,
1908, Fl. Buitenzorg, 6, 2, g. 43; Seidenf., 1978,
Dansk Bot. Ark. 32, 2: 142, g. 90; id., 1992, Opera
Bot. 114: 72; Comber, 1990, Orch. Java: 75, g.;
Sei denf., Wood, 1992, Orch. Malay. Sing.: 127, g.
51 f; Aver., 1994, Identif. Guide. Vietnam. Orch.:
77; P.H. Ho, 2000, Ill. Fl. Vietnam, 3: 791, g.
10926; Aver. et Averyanova, 2003, Updated Check-
list Orch. Viet.: 60; Soto, Cribb, 2010, Lankesteri-
ana 9, 3: 368. – V. parishii Reichenb. f., 1866, Otia
Bot. Hamburg 1: 39.
Described from Java (“Sadjram Provin-
ciae Bantam... Nusae Kambangae insulae”). Type
(“Blume, 1719”) – L.
Epiphytic or lithophytic creeping vine to
12 m long. Stem succulent, aphyllous, with inter-
nodes 5–8 cm long. Leaves reduced to small trian-
gular green scales 5–10 mm long. Inorescence very
short, 1–3 owered. Floral bracts small, broadly-tri-
angular, obtuse. Tepals pale greenish, narrowly ovate
to ovate, to 3 cm long, 8–10 mm wide, obtuse. Lip
joined to column almost to the anther, 2.2–2.6 cm
long, apically 3-lobed; side lobes erect, rounded,
with crisped reexed edges, pale green; median lobe
1 cm long, rounded, with reexed slightly toothed
edges, almost entirely covered with pale pinkish
hairs about 2 mm long. Fig. 17; 24 g, h.
Ecology. Primary and secondary broad-
leaved evergreen and deciduous dry forests, open
dry secondary shrubs, rocky outcrops in semiarid
areas on any kind of soils at elevation from sea level
to 400 m a.s.l. Fl. May – July. Rare (VU).
Distribution. Vietnam (Dac Lac, Dong Nai,
Kien Giang, Lam Dong, Ninh Thuan, Phu Khanh.
Tay Ninh). Myanmar, Thailand, Laos, Malacca Pen-
insula, Java.
Studied specimens. Dac Lac, Buon Don,
Yok Don national park, HLF 7175 (HN, LE); Kien
Giang, Phu Quoc, photo (LE); Ninh Thuan, Ninh
Hai, Nui Chua national park, HLF 3223 (HN, photo
LE); Tay Ninh, Tan Bien, Lo Go-Xa Mat national
park, HLF 7450 (HN), HLF 7592 (HN), HLF 7677
(HN), HLF 7736 (HN); Ninh Hoa, Sigaldi, 318 (P);
Suai Dan, Nhatrang, Vidal 4866 (P); Mt. Dinh, Ba-
ria, Pierre 6565 (P, K), d’Alleizette 6974 (L); Nam
Cat Tien, Tam s.n.
2. V. yersiniana Guillaumin et Sigaldi,
1964, Bull. Mus. Hist. Nat. Ser. 2, 36: 162;
44 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 17. Vanilla aphylla: a – mature plant, b – portions of mature stem (HLF 7175), c – inorescence, d –
ower, e – attened lip (Khoi, s.n.).
45
Turczaninowia 2011, 14(2) : 15–100
Soto, Cribb, 2010, Lankesteriana 9, 3: 397. – V. al-
bida auct. non Blume, 1823: Seidenf., 1978, Dansk
Bot. Ark. 32, 2: 141, p.p., g. 87; id., 1992, Opera
Bot. 114: 72, p.p.; Seidenf., Wood, 1992, Orch. Ma-
lay. Sing.: 127, p.p., g. 51, d, e, pl. 5 d; Aver., 1994,
Ident. Guide Vietnam. Orch.: 76; P.H. Ho, 2000, Ill.
Fl. Vietnam, 3: 791, g. 10927; Aver. et Averya nova,
2003, Updated Checklist Orch. Viet.: 60. – V. sumat-
rana auct. non J.J. Smith: Seidenf., 1972, Bot. Tid-
skr. 67, 1–2: 96, p.p., g. 18.
Described from southern Vietnam (“Sur la
piste Yersin du Hon Ba a Duoi Dau (Nhatrang) et
plantee a Duoi Dau pepiniere”). Syntypes (26 mars
1963, de Sigaldi 362, de Sigaldi 309) – P.
Epiphytic creeping vine. Leaves broadly lan-
ceolate to narrowly ovate, 12–14 cm long, 3.5–5 cm
wide. Inorescence 1–3 cm long, few-owered.
Flowers white with greenish tint, 6–8 cm across.
Floral bracts broadly-triangular, rounded, 2 mm
long. Sepals and petals 4–5 cm long and about 1 cm
wide, petals little smaller. Lip 4–4.5 cm long, at the
base forms narrow tube, joined to column, trumpet-
shaped, about 2 cm across the mouth, apex glabrous
or hardly sparsely papillose, disc at the center with
bundle of broad, fan-shaped, toothed scarious scales.
Column about 4 cm tall. Fruit to 8 cm long. Fig. 18.
Ecology. Primary and secondary evergreen
lowland forests on silicate soils. Fl. March – April.
Very rare (DD).
Distribution. Vietnam (Khanh Hoa).
Thailand.
Notes. Vanilla yersiniana is known from
Vietnam by alone type collection until now. This rare
species very close to Indonesian V. albida Blume
(reported from Java and Sumatra), from which it dif-
fers in more or less attenuate leaves, whitish owers
and in scarcely developed papillae on lip apex.
3. V. siamensis Rolfe ex Downie,
1925, Kew Bull. 10: 410; Seidenf., 1978,
Dansk Bot. Ark. 32, 2: 138, g. 86; Chen Sing-chi,
Tsi Zhanhuo, Luo Yibo, 1990, Nat. Orch. China:
411, g.; Soto, Cribb, 2010, Lankesteriana 9, 3:
393. – V. pierrei Gagnep., 1931, Bull. Mus. Hist.
Nat. (Paris), 2 ser. 3, 7: 686; Gagnep., 1934, Fl. Gen.
Indochine, 6, 5: 582, g. 55, 10–13 (12 – sphalm.);
Seidenf., 1992, Opera Bot. 114: 72; Aver., 1994,
Identif. Guide. Vietnam. Orch.: 76; P.H. Ho, 2000,
Ill. Fl. Vietnam, 3: 792, g. 10929; Aver. et Averya-
nova, 2003, Updated Checklist Orch. Viet.: 60.
Described from northern Thailand (“Thai-
land, Doi Sutep, 690 m”). Type (“Kerr 95”) – K
(holotype), C, P (isotypes).
Epiphytic or lithophytic creeping vine to
12 m long. Stem eshy, with long internodes and
loosely arranged, eshy leaves. Leaves elliptic, 14–
25 cm long, 6–8 cm wide, acuminate; shortly peti-
olate, petiole broad, 1.5–2.5 cm long. Inorescence
7–14 cm long, many and dense owered. Floral
bracts triangular ovate, 2–8 mm long, eshy. Pedicel
and ovary to 2 cm long. Flowers opening succes-
sively. Sepals and petals yellowish green to nearly
white, oblong or narrowly ovate, 3.8–4.5 cm long,
about 1–1.3 cm wide, obtuse. Lip rhombic-obovate,
about 4 cm long, adnate to column, trumpet-shaped,
indistinctly 3-lobed. Lateral lobes embracing co-
lumn; median lobe undulate at margin, with bunch
of fat papillae at apex; disk at center with several
subquadrate, densely imbricate, scarious scales cili-
ate at margin. Column 2–2.6 cm long, hairy at front.
Fruits odorless. Fig. 19.
Ecology. Primary and secondary evergreen
forests on silicate soils at elev. 800–1200 m a.s.l. Fl.
April – May. Fruits July – Sept. Very rare (DD).
Distribution. Vietnam (Binh Dinh). Thai-
land, SW. China, Cambodia.
Studied specimens. Mt. Binh Dinh, Pierre
s.n. (P).
Notes. In Vietnam the species is know by the
single collection until now. Its identication remains
questionable. New material is needed for additional
study that can reconrm occurrence of this rare species
in the country if it is not yet completely extinct.
Miguelia Aver.,
gen. nov. – Vanilla Plum. ex Mill., 1754,
Gard. Dict. Abr. 4, 3 p.p.: Gagnep., 1934, Fl. Gen.
Indo-Chine, 6, 5: 580–583; Seidenf., 1978, Dansk
Bot. Ark. 32, 2: 138–146; id., 1992, Opera Bot. 114:
71–72; Aver., 1994, Ident. Guide Vietnam. Orch.:
75–77; P.H. Ho, 2000, Ill. Fl. Vietnam, 3: 791–792;
Su Horng-Jye, 2000, Fl. Taiwan 5: 1063; Aver. et
Averyanova, 2003, Updated Checklist Orch. Viet.:
60–61; Newman et al, 2007, Checkl. Vasc. Pl. Lao
PDR: 283; Schuiteman et al., 2008, Nord. Journ.
Bot. 26: 314–315; Chen Sing-chi, Cribb, 2009, Fl.
China, 25: 167–168. – “Vanilla annamica group”:
Soto, Cribb, 2010, Lankesteriana 9, 3: 359.
Type: M. somai (Hayata) Aver. (Vanilla so-
mai Hayata).
Inorescentia axillaris paniculata, ramis la-
teralibus 1–2 cymis secundis bioris paucis (solita-
riis) et oribus apicalibus ferentibus, interdum ad
cymam axillarim bioram unam reducta. Inores-
centiae bracteae biformes: geminae, suboppositae,
conchoideae, apice rotundatae, aliusmodi alternan-
46 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 18. Vanilla yersiniana: a – owering plant, b – ower (Thailand s.n.), c – partially dissected lip, side view
(Sigaldi, 362, type), d – partially dissected, attened lip, e – scales at the lip center (Phusomsaeng 90).
47
Turczaninowia 2011, 14(2) : 15–100
Fig. 19. Vanilla siamensis: a – owering plant (Thailand s.n.), b – inorescence (Pierre s.n.), c – ower (Thai-
land, s.n.), d – attened lip, e – scales at the lip center, f – column, side view (Maxwel 74-471).
48 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
tes, triangulares, acutae. Bracteae geminae conchoi-
deae initio basi connatae, capsulam compressam in-
orescentiam juvenilim amplectentim formantes.
Climbing epiphytic or lithophytic herba-
ceous succulent vines to 20 m long. Stem eshy,
green, with succulent leaves and aerial roots at
nodes. Leaves shortly petiolate. Inorescence – axil-
lary panicle, with one or two secondary secund axes
bearing one to several biorous, secund cymes and
sometimes few solitary apical owers; not rare in-
orescence reduced to the single 2-owered axillary
cyme. Inorescence bracts of two different types.
First are “internodal” bracts, rigid, paired, subop-
posite, ovate, concave, rounded at apex, shell-like;
at early stage each couple joined from the base with
their margins forming attened capsule that embrac-
es juvenile inorescence spire. Such bracts placed in
leaf axil and at internodes of inorescence secondary
axis, below offshoot of each individual cyme. Bracts
of second type (“nodal” bracts) of usual appearance
are not paired, triangular, conduplicate, acute, placed
at node of axis, nearly opposite to offshoot of each
individual cyme. Floral bracts rather small, triangu-
lar or broadly triangular, acute, similar to “nodal”
inorescence bracts, but much smaller. Flowers re-
supinate, widely opening, with abscission layer be-
tween ovary and perianth. Sepals and petals similar,
free, spreading. Lip spurless, joined to lateral sides
of column almost to top of column, trumpet-shaped,
free apex broadening, entire or indistinctly 3-lobed,
densely haired with long fat clavate papillae; disk
at center with scarious, densely imbricate, recurved,
lamellate scales and 2–4 short low lamellate, lateral
keels. Column erect, slender, slightly curved. Anther
apical; pollinia mealy or granular, without caudicle
or viscidium; rostellum broad. Fruit obscurely trian-
gular in section, eshy, hardly dehiscent. Seeds with
stout black testa, wingless. Fig. 20.
2–3(2) species. SE. China, Taiwan, Laos,
Vietnam.
Etymology. Generic name is dedicated to
the memory of the genus monographer, Dr. Miguel
Angel Soto Arenas, who was tragically murdered in
Mexico at the peak of his bright botanic activity.
Note. The genus has close relation to Vanil-
la, from which strikingly differs in the inorescence
structure, inorescence bracts and the carinate disk
of the lip. The differences are shortly outlined in the
identication key presented below (see also g. 20).
Fig. 20. Schematic drawings and graphic schemes
of inorescence in species of Miguelia: a–d (M. somai),
e, f (M. annamica). Bracts on inorescence drawings
are cut in a section parallel to axis. Explanations of ab-
breviations on gures: st – stem; lf – leaf; ov – ovary;
– ower; ib – “internodal” bracts; nb – “nodal” bracts;
fb – oral bracts.
1. Scape at the base naked with no shell-like, subopposite bracts embracing axis; inorescence – raceme with
few to many spirally arranged owers; scape naked or with few triangular, or leaf-like bracts .......... Vanilla
- Scape at the base with 2 subopposite, rigid, concave, shell-like bracts embracing axis (in early stage bracts
joined with their margins forming compressed lens-like capsule that contains juvenile inorescence spire);
inorescence – panicle with 1 or 2 axes bearing one to several biorous cymes (occasionally inorescence
reduced to alone biorous axillary cyme); inorescence bracts of two different types – paired, subopposite,
shell-like, rounded at apex and alternative, triangular, acute ............................................................ Miguelia
Key to species
1. Inorescence monochasial, with 1 secondary axis, less than 1(3) cm long, with 1–2(4) biorous secund
cymes; owers in inorescence commonly open simultaneously; lip broadest at apex, inside with pink or
reddish tint .................................................................................................................................... 1. M. somai
49
Turczaninowia 2011, 14(2) : 15–100
- Inorescence commonly dichasial, with 2 secondary axes, much longer than 1 cm, each axis with several
biorous cymes and 1–3 solitary apical owers, owers in inorescence opens in succession; lip broadest at
the middle, white tinged with green ....................................................................................... 2. M. annamica
Ecology. Primary and secondary evergreen
broad-leaved, mixed and coniferous forests, as well
as secondary shrubs on any kind of soils (but more
common on rocky limestone, particularly on steep
rocky slopes and cliffs) at elev. 300–1400 m a.s.l. Fl.
April – June. Occasional (VU).
Distribution. Vietnam (Bac Kan, Cao Bang,
Ha Giang, Hoa Binh, Lang Son, Lao Cai, Ninh Binh,
Quang Binh, Son La, Thai Nguyen, Thanh Hoa).
S. China, Taiwan.
Studied specimens. Bac Kan, Cho Don,
HAL 4858 (HN, LE); Bac Kan, Na Ri, NTH 3711
(HN); Cao Bang, Bao Lac, CBL 306 (HN), CBL
1481 (HN, LE), CBL 1585 (HN, LE); Cao Bang,
Thach An, CBL 961 (HN); Ha Giang, Bac Me, HAL
6575 (HN, LE); Ha Giang, Meo Vac, CBL 1866
(HN), HAL 8499 (HN); Ha Giang, Quan Ba, CPC
156 (HN, LE); Ha Nam Ninh, Cuc Phuong, X. Bong
146 (LE); Lang Son, Huu Lien, Huu Lien protected
area, DKH 4186 (HN, LE), NTH 3257 (HN, LE);
Lao Cai, Van Ban, HAL 2296 (HN); Quang Binh,
Bo Trach, HAL 6144 (HN), HAL 6330 (HN); Quang
Binh, Minh Hoa, HAL 5880 (HN), HAL 12217 (HN,
LE); Quang Binh, Quang Ninh, HAL 11524 (HN);
Son La, Quynh Nhai, HAL 11296 (HN); Son La, Yen
Chau, HAL 9485 (HN, LE); Thai Nguyen, Dong Hy,
HAL 61 (HN, LE); Thanh Hoa, Ba Thuoc, Pu Luong
protected area, HAL 1097 (HN), HAL 3071 (HN),
HAL 3254 (HN); Thanh Hoa, Quan Hoa, HAL 3925
(HN).
Notes. Some collections cited here are pre-
sented by sterile specimens, which identication
needs conrmation. Meanwhile, it is obviously cer-
tain, that M. somai is most common species of the
genus in northern Vietnam, particularly in limestone
areas.
Vigorous plants with 4-owered inores-
cence and large owers were described from S.
China as Vanilla shenzhenica. I see no specic dif-
ference in oral morphology of this novelty.
2. M. annamica (Gagnep.) Aver.,
comb. nov. – Vanilla annamica Gagnep.,
1931, Bull. Mus. Hist. Nat. (Paris), 2 ser. 3, 7: 686;
id., 1934, Fl. Gen. Indochine, 6, 5: 584, g. 56, 1;
Lang et Tsi, 1976, Icon. Corm. Sinic. 5: 651, g.
8132; Seidenf., 1992, Opera Bot. 114: 72; Aver. et
Averyanova, 2003, Updated Checklist Orch. Viet.:
60; Soto, Cribb, 2010, Lankesteriana 9, 3: 368.
1. M. somai (Hayata) Aver.,
comb. nov. – Vanilla somai Hayata, 1916,
Icon. Pl. Formos. 6: 88, tab. 14; Soto, Cribb, 2010,
Lankesteriana 9, 3: 368. – V. grifthii Reichenb. f.
var. formosana Ito, 1911, Icon. Pl. Japan. 1, 4: 1,
tab. 1. – V. ronoensis Hayata, 1920, Icon. Pl. For-
mosan. 9: 114. – V. shenzhenica Z.J. Liu et S.C.
Chen, 2007, Acta Phytotax. Sin. 45: 301, g. 1. –
V. annamica auct. non Gagnep.: Aver., 1988, Bot.
Journ. (Le ningrad), 73, 3: 427, 429, g. 6; id., 1994,
Identif. Guide. Vietnam. Orch.: 76, g. 3; P.H. Ho,
2000, Ill. Fl. Vietnam, 3: 792, g. 10928; Aver. et
Averyanova, 2003, Updated Checklist Orch. Viet.:
60. p.p. – V. albida auct., non Blume: Su Horng-
Jye, 2000, Fl. Taiwan 5: 1065, g. 457; Shih-Wen
Chung, 2008, Orch. Taiwan 2: 203, photos.
Described from Taiwan (“Taiwan, between
Urai and Agisku …”). Type (“May 1916, Hayata
s.n.”) – TI.
Epiphytic or lithophytic creeping vine to
15 m long. Stem internodes 5–10 cm long. Leaves
shortly petiolate, narrowly ovate to elliptic-lanceo-
late, acuminate, 10–30 cm long, 4–9 cm wide, eshy.
Inorescence monochasial (very rare dichasial),
secund, 1–2(3) cm long, with 1–2 (very rare with
3–4) 2-owered secund cymes, owers simultane-
ously opening in each cyme. Inorescence “inter-
nodal” bracts oval, rigid, rounded at apex, concave,
0.8–2 cm long, 0.5–1 cm wide; “nodal” bracts trian-
gular, acute. Floral bracts triangular, 0.7–1 cm long.
Pedicel and ovary 2–3 cm long. Flowers 4–5 cm in
diam., white or white with yellowish or greenish tint,
lip pink or reddish inside. Sepals and petals simi lar,
spreading, slightly recurved, broadly oblanceolate,
2–3 cm long, 5–8 mm wide, obtuse. Lip 2.5–3 cm
long, connate for about 4/5 its length with sides of
column forming a trumpet-shaped tube. Lip apex
entire or indistinctly 3-lobed; lateral lobes large,
broad, margins crisped and irregularly incised; me-
dian lobe suborbicular, eshy, with many fat papil-
lae 1.5–2 mm long; disk at center with many dense,
scarious, incumbent, imbricate, overlapping scales
and with 2 short low lateral keels. Column slender,
glabrous, straight, 2–2.5 cm long, at the apex with
2 small dent-like wings. Anther hemispheric, about
3 mm in diam. Stigma transversely slit-like; rostel-
lum square, curved downwards and covering stigma.
Fruit cylindric, 7–10 cm long, obscurely 3-ridged,
dehiscent. Fig. 20 a-d; 21; 24 a; 27 a, b.
50 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 21. Miguelia somai: a – owering plant, b – inorescence with ower buds (HAL 6575), c – inorescence
with open owers, d – ower (CPC 156), e – partially dissected and attened lip (Bong 146).
51
Turczaninowia 2011, 14(2) : 15–100
Fig. 22. Miguelia annamica (VH 3519): a – portion of owering stem, b – inorescence, c – attened sepals
and petals, d – partially dissected and attened lip, e – column, frontal view, f – pedicel and ovary.
52 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 23. Aphyllorchis montana: a (HAL 2992); A. annamensis: b, c (HLF 5617); Listera latilabra: d (HLF
1355); Erythrorchis altissima: e (HLF 6214), f (HAL 11571), g (HLF 6214); Cyrtosia integra: h (HAL 4874), i (HAL
1510).
53
Turczaninowia 2011, 14(2) : 15–100
Fig. 24. Cyrtosia nana: a (HAL 2896), b (HLF 6674); Cyrtosia falconeri: c, d (HAL 9653); Galeola nudifolia:
e, f (HLF 6213); Vanilla aphylla: g (HLF 3223), h (HLF 7175); Miguelia somai: i (CPC 156).
54 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Described from southern Vietnam (“An-
nam: col de Mangiang, prov. Quinhon, …; Lang-
bian, entre B’dle et Dankia; …; Lang-bian, …; prov.
de Binh-Thuan, Djiring, …”). Syntypes (“Poilane
17973, 18635; Jacquel 622; Magnein 82”) – P.
Epiphytic or lithophytic creeping vine to
20 m long. Stem about 1 cm in diam., with internodes
6–15 cm long and slightly swollen nodes. Leaves
shortly petiolate or subsessile, elliptic, 15–30 cm
long, 5–10 cm wide, eshy. Inorescence dicha-
sial or occasionally monochasial, each axis secund,
(4)5–20 cm long, with (3)4–10 2-owered secund
cymes and 1–3 solitary apical owers; owers open-
ing in succession. Inorescence “internodal” bracts
oval, rigid, rounded at apex, concave, 1–1.5 cm
long, 5–7 mm wide; “nodal” bracts triangular, acute.
Floral bracts broadly triangular, 1–5 mm long, ob-
tuse to acute. Pedicel and ovary 2.5–3 cm long. Flo-
wers white, tinged with greenish. Sepals and petals
broadly oblanceolate, 2–3 cm long, 5–7 mm wide.
Lip 2–2.5 cm long, 1–1.5 cm wide, connate with
column on 3/4 of its length forming trumpet-shaped
tube. Lip apex entire or indistinctly 3-lobed; lateral
lobes broad, margins crisped and irregularly incised;
apex of median lobe somewhat obtuse, eshy, with
many fat papillae 1.5–2 mm long; disk at center with
many dense, scarious, incumbent, imbricate, over-
lapping scales and with 2–4 short lateral lamellate,
shallowly incised keels. Column slender, glabrous,
straight, 1–2 cm long. Fig. 20 e; 22.
Ecology. Primary and secondary broad-leaved
and mixed evergreen forests, commonly on steep
slopes and cliffs composed with silicate rocks at elev.
1000–1400 m a.s.l. Fl. February – April. Rare (EN).
Distribution. Vietnam (Binh Dinh, Binh
Thuan, Lam Dong, Ninh Thuan, Thua Thien Hue).
S. China, Laos.
Studied specimens. Ninh Thuan, Ninh Son,
VH 3519 (HN, LE); Thua Thien Hue, Phu Loc, Bach
Ma national park, HLF 1333 (HN, LE).
Notes. Basic structure of M. annamica o-
wers is similar to the previous species. Main differ-
ences are indicated in the key to species. Dichasial
inorescence with 2 axes bearing many biorous
cymes is characteristic for this rare species. Our
previous record of M. annamica (Aver., 1988, Bot.
Journ. (Leningrad), 73, 3: 427, 429, g. 6; id., 1994,
Identif. Guide. Vietnam. Orch.: 76, g. 3) belongs
to M. somai.
Subtrib. 5.2.3. Lecanorchidinae Dressler,
1979, Selbyana 5, 2: 205.
Type: Lecanorchis Blume.
2(1) genera and 30–40(2) species. Tropical
regions of the world with largest diversity in SE.
Asia and Australia.
Lecanorchis Blume,
1858, Mus. Bot. 2: 188; Seidenf., 1978,
Dansk Bot. Ark. 32, 2: 125–128; Hashimoto, 1990,
Ann. Tsukuba Bot. Gard. 9: 1–40; Comber, 1990,
Orch. Java: 76–77; id., 2001, Orch. Sumatra: 133–
134; Seidenf., Wood, 1992, Orch. Malay. Sing.: 132–
134; Su Horng-Jye, 2000, Fl. Taiwan, 5: 932–936;
Pearce et Cribb, 2002, Orch. Bhutan: 67–68; Aver. et
Averyanova, 2003, Updated Checklist Orch. Viet.: 41;
Chen Sing-chi, Cribb, 2009, Fl. China, 25: 171–172.
Lectotype: L. japonica Blume.
Terrestrial achlorophyllous leaess herbs
with slender, erect or ascending, rigid, branching
rhizome. Stems erect, usually black or dark brown,
slender, usually simple, with sparse small scale-
like sheaths. Inorescence terminal, branching or
not, with partial few-owered spikes. Floral bracts
small, triangular, membranous. Flowers resupinate,
yellowish to dull brownish, with small toothed ca-
lyculus below perianth. Sepals and petals free, sub-
similar. Lip base with canaliculate claw connate to
lateral sides of column forming a short tube, rarely
free, not spurred; upper part of lip entire or 3-lobed;
disk often hairy or papillose. Column footless, more
or less slender, slightly broadening toward apex,
slightly curved. Anther terminal; pollinia 2, granu-
lar-mealy, without caudicle and viscidium; rostel-
lum short. Fruit erect, cylindric capsule.
20(2) species. Tropical and subtropical Asia
to islands of Pacic.
Note: All species of this genus are unat-
tractive plants with bleak ephemeral owers easily
overlooked in botanical collecting and very poorly
represented in available collections. It is highly
probable that in Vietnam may be found more spe-
cies than yet reported.
Key to species
1. Stem 12–30 cm tall; rhizome 2–3 mm thick; sepals less than 12 mm long; lip side lobes obtuse to acute; fruit
1.8–2.4 cm long ...................................................................................................................... 1. L. vietnamica
- Stem 20–50 cm tall; rhizome 4–6 mm thick; sepals longer than 12 mm; lip side lobes round at apex; fruit
2.5–4.5 cm long .................................................................................................................... 2. L. malaccensis
55
Turczaninowia 2011, 14(2) : 15–100
1. L. vietnamica Aver.,
2005, Rheedea, 15, 2: 92, g. 7, 8, f-i.
Described from central Vietnam (“Thua
Thien-Hue Prov., A Luoi Distr., A Roang Municipal-
ity, Tra Lenh Forestry Department station, around
point 16º04’38’’N, 107º29’10’’E., at elev. 700-
800 m, on tops of ridge”). Type (“20 April 2005,
L. Averya nov, P.K. Loc, N.T. Vinh etc., HAL 7247”) –
HN (holo), LE (iso).
Rhizome vertical, gray-brown, 2–3 mm
thick. Stem slender, dark brown-purple to nearly
black, 12–30 cm tall, 0.5–1 mm thick, with many
small distant bracts and few thick gray-brown roots
at the base. Inorescence simple or branching, with
lax terminal few-owered spikes, rachis zigzag.
Floral bracts very small, ovate, obtuse, glabrous.
Ovary cylindrical, 8–12 mm long, yellowish or
pinkish. Calyculus short, campanulate, 0.5–1 mm
long, 1.5 mm across, with irregular, erose dentate
margin. Flowers sessile, hardly opening, campanu-
late. Sepals and petals subsimilar, light brown, yel-
lowish to nearly white, oblong-spatulate, with 3 in-
distinct veins, 8–11 mm long, 1.8–3 mm wide. Lip
8–11 mm long, 5–8 mm wide, adnate at the base to
lateral sides of column and forming inated sac-like
nectary; lip blade 3-lobed; side lobes erect, triangu-
lar-cuneate to broadly-falcate, acute, 1.5–2.5 mm
long, 1–1.8 mm wide, median lobe turned down at
right angle, oblong or obovate, 2–3 mm long, 2.2–
3 mm wide, usually longer than wide, rounded at
apex, densely co ve red with long, exuose, white
to lilac-pinkish hairs. Column straight, 5–6 mm
tall, 1–1.5 mm wide, stigma in form of eshy ovate
nely papillose lamella reexed forward; anther cup
hemispheric, about 1 mm across. Fruit cylindrical to
narrowly-ellipsoid black capsule, 1.8–2.4 cm long,
about 3 mm wide. Fig. 25 a-f; 27 c-e; 28.
Ecology. Primary closed, broad-leaved, ev-
ergreen, forests on silicate soils, commonly along
ridge edges and on mountain tops, at elev. 300–1600
m a.s.l. Fl. February – May. Rare (VU).
Distribution. Vietnam (Khanh Hoa, Kon
Tum, Lam Dong, Lao Cai, Nghe An, Quang Nam,
Thua Thien-Hue). Endemic.
Studied specimens. Khanh Hoa, Khanh
Son, VH 4156a (HN; LE); Kon Tum, Dak Gley,
VH 2237 (HN; LE); Lam Dong, Lac Duong, VH
3780 (HN; LE); Lao Cai, Van Ban, HAL 2715 (HN;
LE); Nghe An, Tuong Duong, HLF 6670 (HN; LE);
Quang Nam, Phuoc Son, HAL 11988 (HN; LE);
Thua Thien-Hue, Nam Dong, HAL 6982 (HN; LE);
d-EXSICCATES OF VIETNAMESE FLORA 0014/
HAL 7247.
Note: The species differs from the most
similar L. malaccensis Ridl. and L. japonica Blume
in much smaller owers, inated sac-like base of the
lip and its broadly cuneate, acute side lobes.
2. L. malaccensis Ridl.,
1893, Trans. Linn. Soc. London, Bot. 3, 9:
371; Seidenf., 1978, Dansk Bot. Ark. 32, 2: 126, g.
77; Comber, 2001, Orch. Sumatra: 133; Seidenf.,
Wood, 1992, Orch. Malay. Sing.: 134, g. 55 f, g. –
L. javanica auct. non Blume: Aver. et Averyanova,
2003, Updated Checklist Orch. Viet.: 41.
Described from central Malacca Peninsula
(“Tahan Woods”). Type (“Ridley s.n.”) – ?
Rhizome vertical or ascending, dull brown,
4–6 mm thick. Stem slender, dull brownish, later
nearly black, 20–50 cm tall, 1–2 mm thick, with few
distant bracts and many thick brownish roots at the
base. Inorescence simple, with lax terminal few-
owered spikes, rachis rather straight. Floral bracts
small, triangular, acute, glabrous. Ovary cylindrical,
2–4 cm long, nearly black. Calyculus short, cam-
panulate, about 1 mm long, 2–3 mm across, with
irregular, erose dentate margin. Flowers odorless,
sessile, hardly opening, campanulate. Sepals and
petals subsimilar, light pale purple-brown or pin-
kish, broadly lanceolate, with 3–5 veins, 15–17 mm
long, 2–3.5 mm wide, petals slightly broader. Lip as
long as sepals, the base joined to sides of column on
2–3 mm forming narrow tube, widening gradually
to 3-lobed apex 5–7 mm wide from tip to tip of side
lobes when attened, side lobes brownish, triangu-
lar, almost rounded at apex, median lobe rounded,
3–4 mm long, yellow, densely haired with white ex-
uose hairs. Column pure white, straight, 10–11 mm
tall, 1–1.5 mm wide, stigma concave, round, nely
papillose; anther cup subquadrate, about 1 mm tall.
Fruit cylindrical to narrowly-ellipsoid black or dark
brown capsule, 2.5–4.5 cm long, about 4 mm wide.
Fig. 25 g, h; 27 f, g.
Ecology. Primary and secondary, rather
dry, broad-leaved and mixed, evergreen forests on
shale and granite, commonly along ridge edges and
on mountain tops, at elev. 700–2000 m a.s.l. Fl.
March – April. Very rare (EN).
Distribution. Vietnam (Dak Nong, Kon
Tum). Thailand, Malacca Peninsula, Sumatra.
Studied specimens. Dak Nong, Dak Glong,
Ta Dung Nature Reserve, HLF 5521 (HN, LE); Kon-
tum, W slope of Ngoc Linh mt, VH 1300 (HN; LE).
Trib. 5.3. Gastrodieae Lindl.,
1821, Collect. Bot. App.
56 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 25. Lecanorchis vietnamica (HAL 7247, type): a – owering and fruiting plant, b – attened sepals and
petals, c – attened lip, d – column, side and frontal view, e – pedicel and ovary, f – apex of ovary with cuniculus;
L. malaccensis (VH 1300): g – ower with attened sepals and petals, with removed lip, h – attened lip.
57
Turczaninowia 2011, 14(2) : 15–100
Subtrib. 5.3.1. Gastrodiinae Lindl.,
1840, Gen. Sp. Orch. Pl.: 383.
Type: Gastrodia R. Br.
7(5) genera and 70–90(11) species. Tropi-
cal regions of the world with largest diversity in SE.
Asia and Australia.
Didymoplexiella Garay,
1954, Arch. Jard. Bot. Rio de Janeiro 13:
33; Seidenf., 1978, Dansk Bot. Ark. 32, 2: 175–176;
Seidenf., Wood, 1992, Orch. Malay. Sing.: 137–
139; Su Horng-Jye, 2000, Fl. Taiwan 5: 849–851;
Comber, 2001, Orch. Sumatra: 111–112; Aver. et
Averyanova, 2003, Updated Checklist Orch. Viet.:
31; Chen Sing-chi, Gale, Cribb, 2009, Fl. China, 25:
206. – Leucolena Ridley, 1891, Journ. Linn. Soc.,
Bot. 28: 340, non (DC.) Benth., 1837.
Type: Didymoplexiella ornata (Ridl.) Garay
(Laucolena ornata Ridl.).
Terrestrial achlorophyllous leaess tuberif-
erous herbs. Tubers horizontal or inclined, cylindri-
cal-fusiform, with few small sparse adpressed scales
at the base. Stem simple, erect, slender, with few
thin roots at the base, small distant bracts and termi-
nal simple racemose inorescence of 1 to many o-
wers. Inorescence rachis elongated after owe ring.
Flowers resupinate, widely opening to sub-campan-
ulate, 2-labiate, ephemeral. Sepals and petals all
joined near their base, dorsal sepal connate to petals
for 1/2–2/3 of their length forming a porrect hood;
lateral sepals united basally for half way, deexed or
weakly spreading. Lip adnate with column at base,
free from sepals and petals, not spurred, apex en-
tire or 3-lobed; disk papillose, with a 2-lobed gland.
Column erect, elongate, footless, widening to the
apex, with 2 slender, downwards curved, hook-like
subulate stelidia more than half of column length on
either side of stigma; anther terminal, with 4 mealy
pollinia in 2 pairs, each pair with viscidium, with no
caudicles, operculum with long lament; rostellum
erect, transverse, small; stigma broad, placed direct-
ly below rostellum. Capsule erect, fusiform.
10(1) species. Mainland tropical SE. Asia,
Ryukyu Islands, Malacca Peninsula, Indonesia.
D. denticulata Aver.,
2010, Taiwania, 55, 2: 92, g. 1, 3; 4. –
D. sia mensis auct. non (Rolfe) Seidenf.: Aver. et
Averya nova, 2003, Updated Checklist Orch. Viet.: 31.
Described from central Vietnam (“Quang
Binh province, Le Thuy district, Kim Thuy mu-
nicipality, around Lang Ho village at 16º59’31’’N
106º38’31’’E, … slopes of Khe Nuoc Trong Ri-
ver …”). Type (9 April 2008, L. Averyanov, P.K. Loc,
N.T. Vinh et al., HAL 11443) – HN (holotype), LE
(isotype). Epitype – d-EXSICCATES OF VIET-
NAMESE FLORA 0129/HAL 11443.
Terrestrial herb. Stem (5)10–25(35) cm
tall, 1–1.5 mm thick, olive-brownish to dull gray-
brownish or nearly black, with 1–3 small distant
triangular bracts and many-owered inorescence.
Tubers longitudinally wrinkled, 2–12 cm long, to
0.8 cm in diameter. Inorescence with many buds
densely arranged at the apex. Floral bracts triangu-
lar, 1–1.2 mm long and broad. Flowers 1 day las-
ting, opening in succession one by one. Inores-
cence rachis elongate after owering to 12 cm long.
Pedicel and ovary at owering, suberect, 6–9 mm
long. Flowers very fragrant, widely opening to sub-
campanulate, 10–15 mm across. Sepals and petals
pink, lilac, violet or bluish with brownish tint, oc-
casionally almost pure white, sub-similar in shape,
obovate, 6–9 mm long, 2.5–4 mm wide, with ob-
tuse to roundish apex; median sepal slightly lon-
ger. Lip light violet with white apex, 5–7 mm long,
2.5–3 mm wide, narrowly obovate, widening from
narrow base, 3-lobed, with eshy triangular, slightly
emarginate callus near the apex; side-lobes broadly
rounded, with down deexed irregularly denticulate
margins; median lobe bluntly rectangular or sub-
quadrate, slightly emarginate and usually irregularly
denticulate. Column 3–5 mm tall, footless, opercu-
lum hemispheric, slightly attened, nely papillose.
Fruit unknown. Fig. 26; 27 h, i; 35.
Ecology. Terrestrial ephemeroid herb. Shady
broad-leaved evergreen primary and secondary low-
land and submontane forests (often with bamboo)
on any kind of soils at elev. 150–1000 m a.s.l. Fl.
March – April. Rare (LR).
Distribution. Vietnam (Nghe An, Quang
Binh, Quang Tri, Tuyen Quang). Laos (Bolikhamxai,
Khammouan).
Studied specimens. Nghe An, Tuong
Duong, HLF 6672 (HN, LE); Quang Binh, Le Thuy,
HAL 11443 (HN, LE); Quang Binh, Minh Hoa, HAL
11676 (HN, LE); Quang Binh, Quang Ninh, HAL
11521 (HN, LE); Quang Tri, Da Krong, HLF 6216
(HN, LE), d-EXSICCATES OF VIETNAMESE
FLO RA 0061a/HLF 6216; Quang Tri, Huong Hoa
HLF 6263 (HN, LE), d-EXSICCATES OF VIET-
NAMESE FLORA 0061b/HLF 6263; Tuyen Quang,
Na Hang, HAL 218 (HN, LE).
Notes. Species name refers nely dentate
margin of the lip. It may be close to D. ornata and
D. siamensis found in western Malesia and Thai-
land, but distinctly differ from both these species
58 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 26. Didymoplexiella denticulata (HAL 11443, type): a – owering plant and old inorescence, b – apex
of inorescence, c – ower, half side view, d – column and attened sepals and petals, e – attened lip, f – lip, side
view, g – tuber.
59
Turczaninowia 2011, 14(2) : 15–100
in low, broad, denticulate lip side lobes. I think all
specimens that I have seen from Vietnam belong to
this species. It’s owers, however, are very delicate
hence lip may be actually analyzed morphology only
in fresh material. In dry and even liquid xed o-
wers, very thin lip margins usually decurved down
and adjoin densely to lower lip surface. As a result,
lip sides look straight (not denticulate) and even un-
lobed exactly resembling lip of D. siamensis. It is
quite possible that plants from Hainan, Taiwan and
Japan reported under this name actually belong to
D. denticulata.
Didymoplexiella denticulata in some areas
of its distribution was observed growing just along
Vietnam-Laotian border, in fact on eastern border of
Bolikhamxai and Khammouan provinces, why Laos
is included here in species distribution area.
Didymoplexiopsis Seidenf.,
1997, Contrib. Orch. Fl. Thailand 13: 13; Aver.
et Averyanova, 2003, Updated Checklist Orch. Viet.: 31;
Chen Sing-chi, Gale, Cribb, 2009, Fl. China, 25: 207.
Type: D. khiriwongensis Seidenf.
Terrestrial achlorophyllous leaess tuberif-
erous herbs. Tubers horizontal or inclined, cylindri-
cal-fusiform, verrucose. Stem simple, erect, slender,
with few thin roots and few small distant bracts at the
base. Inorescence terminal, simple, racemose, few-
owered, with small triangular acute oral bracts.
Flowers not resupinate, widely opening, opening
in succession, white, 2-labiate, ephemeral. Sepals
and petals subsimilar, elliptic-ovate, all spreading;
dorsal sepal and petals joined together at the base
on 1/3, free of lateral sepals. Lateral sepals united
for about 1/2 of their length. Lip free from sepals
and petals, not spurred, moveably articulated with
column foot apex, broadly rectangular, with broad
incurved wing-like side lobes and truncate, slightly
emarginate to erose-dentate apex. Lip ventral sur-
face with massive prismatic curved warty callus at
the base and with transversal obscurely dentate fold
at the middle; dorsal surface with rounded nely
verrucose yellow callus. Column with long column
foot, erect, widening to apex, with 2 slender, curved
downwards, hook-like subulate stelidia as long as
column on either side of stigma; anther terminal, broad,
with 4 mealy pollinia in 2 pairs, operculum with short
lament; rostellum erect, massive, eshy, transverse;
stigma broad, placed directly below rostellum.
Monotypic genus endemic to Indochina and
SE. China.
D. khiriwongensis Seidenf.,
1997, Contrib. Orch. Fl. Thailand 13: 13,
g. 2; Aver. et Averyanova, 2003, Updated Check-
list Orch. Viet.: 31. – Didymoplexiella hainanensis
X.H. Jin et S.C. Chen, 2004, Novon, 14: 176, g. 1.
Described from southern Thailand (“Khao
Luang, Nakorn Sritammarat 800–1000 m”). Type
(“Suksathan 1001”) – C (holo, iso).
Stem dark grey to dull yellowish-brown,
5–15 cm tall, with 2 or 3 dull brownish bracts. Tu-
bers fusiform, 5–6 cm long, 3–10 mm wide. Rachis
5–10 mm long, with few owers; oral bracts trian-
gular, acute, 1.5–2 mm long. Flowers white, nely
fragrant, lip below with yellow-orange spot. Pedicel
and ovary 0.5–1.5 cm long, dark brown. Dorsal se-
pal and petals ovate to narrowly ovate, 7–13 mm
long, 3–5 mm wide, obtuse; lateral sepals ovate,
10–13 mm long, 4 mm wide, obtuse. Lip moveable
joined with column foot apex, broadly rectangular,
5–6 mm long, 11–12 mm wide, indistinctly 3-lobu-
late, margin of side lobes erose-denticulate, midlobe
slightly emarginate; massive callus at the base of lip
covered by numerous small olive-grey warts, center
of lip often with few brous cirri. Column 2.5–3 mm
tall, as long as incurved hook-like stelidia; column
foot incurved, 4–5 mm long. Pollinia light pinkish.
Fig. 29; 36 a-c.
Ecology. Primary and secondary closed
broad-leaved and mixed evergreen forests on any
kind of soils at elev. Fl. 100–1600 m a.s.l. March –
May (July). Rare (VU).
Distribution. Vietnam (Ha Tinh, Khanh
Hoa, Quang Binh, Quang Tri, Thua Thien-Hue).
Thailand, SE. China (Hainan).
Studied specimens. Ha Tinh, Huong Son,
HAL 5040 (HN, LE), HAL 5153 (HN, LE), HAL
5262 (HN, LE), HAL 5341 (HN, LE), Khanh Hoa,
NE slope of Bi Dup mountain, VH 1441 (HN, LE),
Quang Binh, Minh Hoa, VH 4650 (HN, LE), Quang
Binh, Quang Ninh, HAL 11885 (HN, LE), Quang
Tri, Huong Hoa, HLF 5875 (HN, LE), Thua Thien-
Hue, A Luoi, HAL 7246 (HN, LE), d-EXSICCATES
OF VIETNAMESE FLORA 0015/HAL 7246 (HN,
LE), HAL 7890 (HN, LE), HAL 7944 (HN, LE),
Thua Thien-Hue, Huong Thuy, HAL 8177 (HN, LE),
Thua Thien Hue, Bach Ma HLF 1397 (HN, LE), Thua
Thien Hue, Xuan Loc, HLF 1829 (HN, LE).
Notes. Each ower in inorescence com-
monly lasts only during few morning hours, hence
this species is easily overlooked in eld studies.
Didymoplexis Griff.,
1844, Calcutta Journal Nat. Hist., 4: 383;
Seidenf., 1978, Dansk Bot. Ark. 32, 2: 177–178;
Comber, 1990, Orch. Java: 81–83; id., 2001, Orch.
60 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 27. Miguelia somai: a, b (CPC 156); Lecanorchis vietnamica: c, e (HAL 7247, type), d (HAL 6982);
L. malaccensis: f, g (VH 1300); Didymoplexiella denticulata: h (HAL 11443, type), i – (HAL 11521).
61
Turczaninowia 2011, 14(2) : 15–100
Fig. 28. Digital herbarium specimen of Lecanorchis vietnamica (Averyanov L. HAL 7247, epitype).
62 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 29. Didymoplexiopsis khiriwongensis: a – owering plant, b – ower at the beginning of anthesis (HAL
7246), c – ower with lip up-turned during anthesis, d – attened sepals and petals, e – lip from below (dorsal surface),
f – lip from above (ventral surface), g – column half-side view and sagittal lip section, h, i – column, frontal and half-
side views. (HAL 5040).
63
Turczaninowia 2011, 14(2) : 15–100
Sumatra: 112–113; Seidenf., Wood, 1992, Orch. Ma-
lay. Sing.: 139; Su Horng-Jye, 2000, Fl. Taiwan 5:
851–854; Pearce et Cribb, 2002, Orch. Bhutan: 28-
30; Aver. et Averyanova, 2003, Updated Checklist
Orch. Viet.: 31; Chen Sing-chi, Gale, Cribb, 2009,
Fl. China, 25: 205–206. – Leucorchis Blume, 1849,
Mus. Bot. Lugd. 1, 2: 31, non E. Meyer, 1839.
Type: D. pallens Griff.
Terrestrial achlorophyllous leaess tuberif-
erous herbs. Tubers horizontal or inclined, cylindri-
cal-fusiform. Stem simple, erect, with few thin roots
and 1–3 small sheaths near base, glabrous. Ino-
rescence racemose with 1 to many owers, rachis
with small broadly triangular, acute persistent oral
bracts. Flowers resupinate, subcampanulate, 2-labi-
ate, ephemeral, white to pale yellowish-brown. Se-
pals and petals all joined near their base, dorsal se-
pal connate to petals for 1/2 or more of their length
forming a porrect hood; lateral sepals united basally
for more than 1/2 of their length, deexed distally.
Lip adnate with column foot at base, free from se-
pals and petals, not spurred, entire or indistinctly
3-lobed; disk commonly papillose, with 1 or more
basal glands. Column erect, elongate, with distinct
column foot, widening to the apex, with 2 wings at
apex, forming short, tooth-like stelidia; anther ter-
minal with 4 mealy pollinia in 2 pairs, each pair with
viscidium, with no caudicles, operculum with short
lament; rostellum erect, eshy, transverse; stigma
broad, placed directly below rostellum. Capsule
erect, fusiform; pedicel usually erect, elongates after
anthesis to 20–30 cm, dull pinkish, hollow.
20(3) species: SW. Africa, Madagascar, Tro-
pical and subtropical mainland Asia, Ryukyu Is-
lands, Malay Archipelago to New Guinea, tropical
Australia, islands of SW Pacic Ocean.
All species of the genus have very short
owering period due to wich they are easily over-
looked in eld studies. Study of delicate ower struc-
ture and identication may be correctly fullled actually
only on the base of fresh and liquid preserved material.
Key to species
1. Lip moveably articulated with column foot apex, at the base with large prismatic callus covered by numerous
small olive-brown warts, at the center with yellow-orange spot; owers widely opening; pedicels in fruits
pendent, not much elongated ................................................................................................. 3. D. vietnamica
- Lip adnate to column foot, not moveable; without massive callus at the base but sometimes with papillose
rugosity along midvein; without distinct yellow-orange spot at the center; owers not widely opening,
campanulate; pedicels in fruits erect, much elongated, to 10–30 cm ............................................................. 2
2. Flowers mainly 1.5–2.2 cm across, tepals (6)8–12 mm long; lip broadly triangular-orbiculate, more than
6 mm wide, straight or slightly undulate along apical margin; column foot about 2 mm long ... 1. D. pallens
- Flowers mainly less than 1.5 cm across, tepals 6–8 mm long; lip obovate or obdeltoid, less than 5 mm wide,
nely denticulate along apical margin; column foot less than 1 mm long ............................ 2. D. micradenia
1. D. pallens Griff.,
1844, Calcutta Journal Nat. Hist., 4: 383; id.,
1845, ibid., 5, pl. 17; King, Pantl., 1898, Ann. Roy.
Bot. Gard. Calcutta, 8, tab. 346; Garay, Sweet, 1974,
Orch. Ryukyu Isl.: 51, 62, g. 3, g, h; Seidenf., 1978,
Dansk Bot. Ark. 32, 2: 178, g. 110; Comber, 1990,
Orch. Java: 83, g.; id, 2001, Orch. Sumatra: 113,
g.; Seidenf., Wood, 1992, Orch. Malay. Sing.: 138,
g. b-e; Su Horng-Jye, 2000, Fl. Taiwan, 5: 853, g.
361; Pearce et Cribb, 2002, Orch. Bhutan: 30, g. 4;
Aver., Averyanova, 2003, Updated Checklist Orch.
Viet.: 31. – Leucorchis sylvatica Blume, 1849, Mus.
Bot. 1, 2: 31. – Apetalon minutum Wight, 1851,
Icon. Pl. Ind. Orient. 5, tab. 1758. – Didymoplexis
subcampanulata Hayata, 1912, Icon. Pl. Formos.
2: 136. – D. brevipes Ohwi, 1937, Acta Phytotax.
Geobot. 6: 238. – D. sylvatica (Blume) Garay, 1995,
Opera Bot. 124: 15.
Described from India (“Calcutta, Serampo-
re”). Type (“Grifth”) – K (CAL, BR)?
Stem pale brown to dull pinkish or reddish
brown, 5–20 cm tall, at the base with 3–5 sheaths
and few to many wiry roots, often with small glo-
bose thickenings. Rachis elongates in fruits to 3 cm
long, few to many-owered, lax to subdense; oral
bracts ovate, acute, to 2 mm long. Flowers ope ning
in succession, white, campanulate; pedicel and ova-
ry erect, brown, 0.7–1.2 cm; pedicel elongate in fruit
to 15–30 cm long. Dorsal sepal and petals 6–12 mm,
united for more than 1/2 of their length; free por-
tion ovate triangular, shallowly 3-lobed, each lobe
ovate-deltoid, obtuse. Lateral sepals to 12 mm long,
connate to half-way, and with petals for 1/3 of their
length; free portion 2-lobed, each lobe ovate, ob-
tuse, reexed. Lip broadly obovate, 5–7 mm long,
8–12 mm wide, erose-crenate, lateral margins erect
or incurved, apex truncate; disk with dense row
64 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
of warty papillae along midvein. Column slightly
curved, widened to apex, 4 mm long, with 2 oblong
wings. Column foot slightly curved, 2–3 mm long.
Anther hemispaeric. Capsule cylindric-fusiform,
2–4 cm long. Fig. 30.
Ecology. Primary and secondary closed
broad-leaved evergreen forests on any kind of soils at
elev. 30–1600 m a.s.l. Fl. April – May. Very rare (DD).
Distribution. Vietnam (Bac Kan, Dak Nong,
Hoa Binh, Lam Dong, Lao Cai, Nge An, Ninh Binh,
Thua Thien-Hue). Tropical and subtropical main-
land Asia, Ryukyu Islands, Malay Archipelago to
New Guinea.
Studied specimens. Bac Kan, Na Ri, HAL
4960 (HN, LE); Dak Nong, Dak Glong, Ta Dung
nature reserve, HLF 5606 (HN, LE); Hoa Binh, Da
Bac, HAL 517 (HN, LE); Lam Dong, Lac Duong,
VH 3781 (HN, LE); Lao Cai, Van Ban, HAL 2713
(HN, LE); Nghe An, Tuong Duong, HLF 6671 (HN,
LE); Ninh Binh, Cuc Phuong national park, 1 April
1997, Henk van der Werff et al. 14201 (HN, LE);
Thua Thien-Hue, Phu Loc, Bach Ma national park,
HLF 1391a (HN, LE).
Notes. All studied specimens were collected
in fruits, hence occurrence of this species in Viet-
nam still needs conrmation.
2. D. micradenia (Reichenb. f.) Hemsl.,
1883, Journ. Linn. Soc. Bot. 20: 311; Tien-
Chuan, Shih-Wen Chung, 2007, Taiwania, 52,4:
360, g. 2, 3. – Epiphanes micradenia Reichenb.
f., 1868, Seem. Fl. Vit.: 295. – Didymoplexis minor
J.J. Smith, 1900, Bull. Inst. Bot. Buitenzorg 7: 1;
id., 1908, Orch. Java. Atlas, g. 53; Comber, 1990,
Orch. Java: 83.
Described from Fiji (“Ovalau, Fiji”). Type
(“Seemann 610”) – W (holo), K (iso).
Stem pale brown to whitish, erect, 5–15 cm
high, 1.5–2 mm in diam., glabrous, with 3–5 scarious
scales. Tubers pale brown, fusiform or moniliform,
1–5 cm long, 4–7 mm thick. Roots few, arising from
the base of stem. Raceme 2–5 cm, with 8–15 o-
wers; oral bracts ovate, 1.5 mm long, acuminate.
Pedicel and ovary 4–6 mm long, glabrous. Flowers
whitish, tinged with pale red, campanulate. Sepals
and petals connate at base into oral tube. Dorsal
sepal 6–8 mm long, connate with petal for 1/2 to
2/3 of their length, petals shorter. Lateral sepals
5–7 mm long, connate for 2/3 to 4/5 of their length,
whole structure united with petals to about 1/3 of
its length, free apex reexed, 2-lobed, lobules sub-
orbicular. Lip white, obovate to obdeltoid, 4–6 mm
long, erect on both sides, 4–5 mm wide when at-
tened, base cunea te, apex obtuse, nely irregular
denticulate along apical margin, disc with longitu-
dinal raised keel of pale brownish warty papillae.
Column white, clava te, about 5 mm long, apex with
2 truncate rhombic wings, base with short foot, less
than 1 mm long. Anther oval or hemispheric, about
0.8 mm long, white; pollinia white, connected by
broad viscidium detached from rostellum. Capsules
cylindrical, 1.8–2.2 cm long, triangular in cross sec-
tion; pedicels elongate to 10–25 cm long in fruit,
about 2 mm in diam., eshy. Fig. 31.
Ecology. Primary and secondary shady ever-
green forests at elev. 100–300 m a.s.l. Fl. March –
May. Very rare (DD).
Distribution. Vietnam (reported from center of
the country). Taiwan, Java, New Caledonia, islands of
SW. Pacic (Samoa, Vanuatu, Fiji, Tonga, Niue, Palau).
Notes. Species was reported for central
Viet nam without citation of voucher specimens
(Tien-Chuan, Shih-Wen Chung, 2007). We have
not seen any material of this species from Vietnam.
Didymoplexis micradenia morphologically close to
D. pallens, but differs in distinctly smaller owers,
very short column foot and in obdeltoid lip that has
irregularly denticulate margin.
3. D. vietnamica Ormerod,
2000, Oasis, 1, 4: 15; Aver., Averyanova,
2003, Updated Checkl. Orch. Vietnam: 31.
Described from northern Vietnam, Ninh
Binh province (“Cuc Phuong”). Type (“3-04-1997,
B. Gray 7146”) – QRS (holo).
Stem dark grey to dull purple (in fruit)
5–25 cm tall, with 1–3 bracts and few densely ar-
ranged owers opening in succession. Scape elon-
gate during anthesis to 4–5 cm. Floral bracts ovate,
acute, 1–1.5 mm long, 1 mm wide. Pedicel and ova-
ry erect, 9–12 mm long. Flowers odorless, widely
opening, broadly campanulate, 1.5–2.2 cm across,
white, lip with yellow-orange center on both surfac-
es. Sepals and petals subsimilar, ovate, 10–12 mm
long, about 7 mm wide, with obtuse to roundish
apex, petals slightly smaller. Dorsal sepal connate
to petals for 1/2–2/3 of their length; lateral sepals
similarly united to about 2/3 from the base; petals
and lateral sepals united for 1/4 to 1/3 of their length
forming broad shallow tube. Lip moveably articu-
lated with column foot apex, subquadrate, 7–10 mm
long and broad, with broad, raised lateral sides and
broad, truncate, slightly emarginate apex. Ventral lip
surface with massive prismatic, curved, warty callus
at the base and with nely verrucose callosity at the
center of dorsal lip surface. Column erect, slightly
65
Turczaninowia 2011, 14(2) : 15–100
Fig. 30. Didymoplexis pallens: a – fruiting plant (HLF 6671), b – owering plant, c – inorescence (China
s.n.), d – ower, frontal view (Java s.n.), e, f – ower and column (copy of type Grifth drawing).
66 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 31. Didymoplexis micradenia (Taiwan, s.n.): a – owering and fruiting plant, b – owers, c – attened
median sepal and petals, d – attened sepals, e – attened lip, f – column and lip, side view.
67
Turczaninowia 2011, 14(2) : 15–100
curved and widened to apex, 4–6 mm long, with 2
large, falcate, down directed acute hook-like stelidia
3 mm long, on both sides of stigma; column foot
about 1 mm long. Fruit cylindric pendent capsule,
1.8–2.2 cm long. Fig. 32; 36 d-f.
Ecology. Primary and secondary closed
ever green forests on limestone at elev. 300–600 m
a.s.l. Fl. March – April. Very rare (EN).
Distribution. Vietnam (Nghe An, Ninh
Binh). Endemic.
Studied specimens. Nghe An, Tuong
Duong, HLF 6673 (HN, LE), d-EXSICCATES OF
VIETNAMESE FLORA 0092/HLF 6673.
Notes. Very rare endemic of Vietnam. It has
rather isolated taxonomical position in the genus
having subulate hook-like stelidia, moveable lip,
large massive, warty, prismatic callus at the base of
the lip and pendant pedicels, not elongated in fruits.
According to these features it certainly closely allied
to Didymoplexiopsis Seidenf. This highly endan-
gered species is typical relictual element of primary
warm-loving lowland tropical limestone forests.
Yoania Maxim.,
1872, Bull. Acad. Imp. Sci. Petersburg, ser.
3, 18: 68; Su Horng-Jye, 2000, Fl. Taiwan 5: 1070–
1073; Pearce et Cribb, 2002, Orch. Bhutan: 244–
246; Aver. et Averyanova, 2003, Updated Checklist
Orch. Viet.: 61; Chen Sing-chi, Gale, Cribb, 2009,
Fl. China, 25: 210–211.
Type: Y. japonica Maxim.
Terrestrial leaess achlorophyllous glabrous
herbs. Rhizome coralloid, eshy, branched, with
many small scales, lacking roots. Stem erect, stout,
eshy, with distant broad cucullate bracts, leaess,
yellowish- or pinkish-white. Inorescence termi-
nal, racemose, with 1 to few loose owers. Floral
bracts often prominent. Flowers yellowish to pink
or white, eshy, subcampanulate, pedicellate. Se-
pals and petals free, similar, connivent, ovate, con-
cave. Lip oblong, concave, adnate to short column
foot, slipper-shaped or saccate, spurred or not. Co-
lumn erect, broad, winged, with short column foot
at base. Anther terminal, acuminate. Pollinia 4, in 2
pairs, granular-farinaceous, sectile, attached without
caudicles directly to a common viscidium. Stigma
large, concave; rostellum inconspicuous.
3(1) species. Bhutan, NE. India, China, Tai-
wan, Japan.
Y. prainii King et Pantl.,
1898, Ann. Roy. Bot. Gard. Calcutta, 8: 175,
tab. 237; Pearce et Cribb, 2002, Orch. Bhutan: 246,
g. 57; Aver. et Averyanova, 2003, Updated Check-
list Orch. Viet.: 61.
Described from NE. India (“India, Sikkim,
Choongthang (Chunthang)…”, “Naga Hills …”).
Syntypes (“Pantling 469”) – BM, CAL, K, (“Prain
15”) – CAL, K.
Stem pinkish-white, 7–20 cm tall, covered
with distant broadly ovate, loose, whitish scales. In-
orescence with 1–4 lax owers. Floral bracts ovate
to broadly ovate, eshy, 7–9 mm long, 3–5 mm wide,
white to pink, concave, clasping pedicel. Flowers
odorless, erect, white with purple tint, sepals in api-
cal part pink, lip white with many small deep purple
dots. Pedicel and ovary white to pink or purple, cy-
lindric, 2–4 cm long, elongating after owering. Se-
pals and petals subsimilar, concave, ovate to broadly
ovate, 1.5–1.8 cm long, 0.8–1.4 cm wide, petals dis-
tinctly wide to almost round. Lip entire, concave,
slipper-shaped, spurless, 1.5–1.8 cm long, 0.8–1
cm wide. Column white, short, stout, 7–9 mm tall,
with lateral wings at apex, and inconspicuous foot.
Anther cap beaked, light yellowish. Fruit fusiform,
5–6 cm long. Fig. 33; 36 g, h.
Ecology. Primary evergreen broadleaved
humid forests on silicate soils at elev. 1300–1500 m
a.s.l. Fl. April – May. Very rare (EN).
Distribution. Vietnam (Ha Giang). NE. In-
dia, Bhutan.
Studied specimens. Ha Giang, Yen Minh,
CBL 2091 (HN, LE).
Gastrodia R. Br.,
1810, Prodr. Fl. Nov. Holl.: 330; Seidenf.,
1978, Dansk Bot. Ark. 32, 2: 179–181; id., 1992,
Opera Bot. 114: 75; Comber, 1990, Orch. Java:
84–85; id., 2001, Orch. Sumatra: 114–115; Sei denf.,
Wood, 1992, Orch. Malay. Sing.: 139–141; Aver.,
1994, Ident. Guide Vietnam. Orch.: 82–83; Su
Horng-Jye, 2000, Fl. Taiwan 5: 890–896; P.H. Ho,
2000, Ill. Fl. Vietnam 3: 795; Pearce et Cribb, 2002,
Orch. Bhutan: 30–33; Aver. et Averyanova, 2003,
Updated Checklist Orch. Viet.: 37; Schuiteman et
al., 2008, Nord. Journ. Bot. 26: 293; Chen Sing-chi,
Cribb, 2009, Fl. China, 25: 201–205.
Type: G. sesamoides R. Br.
Terrestrial, achlorophyllous, leaess, tuber-
iferous herbs. Tubers underground, many noded,
eshy, spherical to fusiform, sometimes tuberculate
or slightly coralloid, covered with early decaying
scales. Floriferous stem erect, with few to many tu-
bular or scale-like sheaths, glabrous, pale yellowish-
brown. Inorescence terminal, few- to many-ow-
ered lax raceme. Flowers campanulate or tubular,
68 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 32. Didymoplexis vietnamica (HLF 6673): a – owering plant, b – ower, half-side view, c – attened
sepals and petals, d – lip from above (ventral surface) and from below (dorsal surface), e – column half-side view and
sagittal lip section, f, g – column, frontal and half-side views.
69
Turczaninowia 2011, 14(2) : 15–100
Fig. 33. Yoania prainii (CBL 2091): a – owering and fruiting plant, b – attened sepals, petal and lip, c – lip,
half side view, d – column, side and frontal views.
resupinate or not, pink, yellowish, olive-greenish
to brown; lip often lighter, sometimes with orange
marks. Pedicel and ovary usually elongating in fruit.
Sepals and petals united into perianth tube, with only
apices free; sometimes with deep sinus between lat-
eral sepals. Outer surface of perianth tube often ver-
rucose. Petals much smaller than sepals. Lip usu-
ally small, enclosed within perianth tube, adnate to
apex of small column foot, entire or 3-lobed, often
clawed at base, with 2 globose calli at base. Column
straight, elongate, more or less distinctly winged to-
ward apex. Anther terminal, large; pollinia 2, granu-
lar, usually composed of friable massulae, without
caudicles; stigma convex, placed at the base of col-
umn. Capsule erect, elongating with age.
50(5) species. Tropical Africa, Madagascar,
Mascarene Islands, tropical, subtropical and temper-
ate E. Asia to Pacic islands, Australia, with highest
diversity in southeast mainland Asia and Taiwan.
70 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Key to species
1. Stem less than 5 cm tall; rachis shorter than 1 cm, owers densely arranged in almost umbellate raceme
....................................................................................................................................................... 1. G. theana
- Stem longer than 5 cm, rachis longer than 1 cm, owers distant in more or less lax raceme ........................ 2
2. Stem 6–18 cm tall; owers 7–8 mm long, opening in succession one by one, short lasting; lip obovate to
almost circular, broadening to the apex, little shorter and distinctly broader than sepals .... 2. G. tonkinensis
- Stem 16–60 cm tall; owers 8–25 mm long, opening more or less simultaneously, long lasting; lip narrowly
ovate to oblong, narrowing to the apex, much shorter and distinctly narrower than sepals ........................... 3
3. Flowers 8–14 mm long .............................................................................................................. 3. G. punctata
- Flowers 16–25 mm long ................................................................................................................................. 4
4. Inorescence 2–3 owered; owers broadly campanulate, 1.6–1.7 cm long; ovary round in section ................
...................................................................................................................................................... 4. G. taiensis
- Inorescence commonly bears more than 3 owers; owers urceolate, almost tubular, 2–2.5 cm long; ovary
triangular in section ........................................................................................................................ 5. G. major
1. G. theana Aver.,
2005, Rheedea, 15, 2: 90, g. 6, 8 c-e.
Described from central Vietnam (“Vietnam,
Thu Thien-Hue Prov., Nam Dong Distr., Huong
Son Municipality, around point 16°09’58’’N,
107°36’07’’). Type (“31 March 2005, L. Averyanov,
P.K. Loc, P.V. The, A. Averyanova, N.T. Vinh, HAL
7165”) – HN (holotype), LE (isotype).
Tubers cylindrical, nely irregularly verru-
cose, hairy, 2–4 cm long, 4–8 mm in diam. Stem
erect, eshy, straight or exuose, white to yello-
wish-brown, 2.5–5 cm tall, 2–4 mm thick, with 2–3
ovate, obtuse bracts, 3–5 mm long, 2–4 mm wide,
with many small imbricate triangular acute scales
and with few slender roots at the base. Inorescence
dense raceme with 3–8 close owers. Floral bracts
yellowish-brown, ovate, 2–5 mm long. Pedicel and
ovary 3–5 mm long, ovary 2–2.5 mm thick. Flowers
tubular, hardly opening, eshy; sepals with eshy
incurved tips, roughly verrucose outside, with irreg-
ularly warty dorsal ridge-like keel. Sepals white to
pinkish-brown, oblong-ovate, 8–15 mm long, 3.5–
5 mm wide, obtuse, connate with petals on about
9/10; lateral sepals connate each other on 1/4–1/3
from the base. Petals pink, thin, oblong, 1–1.3 cm
long, 3–4 mm wide; free parts of petals eshy, very
small, sometimes hardly visible, shorter than sepals,
narrowly-triangular, 0.4–0.8 mm long, 0.2–0.3 mm
wide. Lip green with red apex and base; 5 nerved,
3.5–4 mm long, 2.5–3 mm wide, entire, nely den-
ticulate along margin, at, cordate, attenuate to blunt of
acute apex; disc with thick short keel-like median cal-
lus near lip apex and 2 white spherical calli at suddenly
narrowing lip base. Column white, straight, 4–4.5 mm
tall, 1.4–1.6 mm wide, with long narrow stelidia, at the
base with very short column foot. Anther cap hemi-
spheric, 0.6–0.7 mm across; stigma round convex,
placed at the base of column. Fig. 34; 36 i; 43.
Ecology. Primary broadleaved evergreen
low land, shady forests on shale and sandstone at elev.
300–400 m a.s.l. Fl. March – April. Very rare (EN).
Distribution. Vietnam (Thua Thien-Hue).
Endemic.
Studied specimens. Species is known till
now by type collection only.
Notes. The species supercially similar to
G. verrucosa Blume recorded from Japan, Malac-
ca, Sumatra and Java, but distinctly differs in very
small, hardly visible narrowly-triangular free part of
petals and cordate base of lip. This rare, probably
strictly endemic plant is typical element of the ab-
original warm-lowing lowland ora of central part
of Vietnam.
2. G. tonkinensis Aver. et Averyanova,
2006, Komarovia, 4: 13, g. 7.
Described from northern Vietnam (“N
Vietnam, Ha Tinh Prov., Huang Son Distr., Son
Hong municipality, around the point 18°34’06’’N,
105°11’40’’E…”). Type (“9 May 2004, Phan Ke
Loc, L. Averyanov, Pham Van The, Nguyen Tien Vinh
HAL 5263”) – HN (holotype), LE (isotype).
Tubers irregularly verrucose, cylindrical,
2–5 cm long, 3–5 mm thick. Stem slender, straight or
slightly exuous, white to light gray-brown, 6–18 cm
tall, 0.8–1 mm thick, with 1–2 distant, ovate, obtuse
bracts, 2.5–3 mm long, at the base with few close
small brown scales and few slender roots. Rachis
with (4)6–8(10) owers. Floral bracts light brown,
ovate, acute, 1.5–2.5 mm long, 1–1.5 mm wide.
Pedicel and ovary 8–10 mm long, 0.6–0.8 mm thick.
Flowers urceolate, hardly ope ning, light brown with
darker nerves, sparsely warty outside. Sepals and
petals oblong, with 1 distinct vein and eshy obtuse
tips. Sepals subsimilar, 7–8 mm long, 1.2–1.5 m
wide; median sepal connate with petals for about
71
Turczaninowia 2011, 14(2) : 15–100
4/5; lateral sepals connate with each other for 4/5
and with petals for 1/5 of their length from the base.
Petals little shorter than sepals, 6–6.5 mm long,
1.2–1.5 mm wide. Lip 5-nerved, concave, broadly
obovate or almost circular, truncate, about 3.5 mm
long, 4.5 mm wide, slightly irregularly cre nulate
along the margin, with thickened median band end-
ing in narrowly-triangular acute eshy mid-lobe; lip
blade narrowing to the base into short at claw, 2
mm long, 0.5 mm wide, with 2 lamellate erect calli,
0.5 mm tall at the base. Column narrow, straight,
4–4.5 mm tall, 0.8–1 mm wide, with narrow lateral
wings in the apical part, with short column foot. An-
ther cap hemispheric, about 0.8 mm across. Fig. 37;
44 a, b.
Ecology. Primary broadleaved evergreen
lowland dry forests on shale at elev. 300–400 m
a.s.l. Fl. April – June. Very rare (EN).
Distribution. Vietnam (Ha Tinh, Hoa Binh).
Laos?
Studied specimens. Hoa Binh, Da Bac,
HAL 329 (HN, LE).
Fig. 34. Gastrodia theana (HAL 7165, type): a – owering plant, b-d – ower, side view and view from below,
e – articially half opened ower with partially dissected lateral sepals, f – lip, side and frontal views, g – column,
frontal view.
Notes. From all species of the genus known
in SE. Asia, G. tonkinensis differs in broadly ovate,
almost circular truncate and narrowly triangular
acute eshy median lobe, as well as in lateral se-
pals, which are connate with petals for less than
half (from the base). The species may be related to
G. abscondita J.J. Sm. and G. verrucosa Blume de-
scribed from Java. This plant is probably endemic to
low hilly area on the border of Vietnam and Laos in
limits of North Indochinese oristic province. It was
discovered on Vietnamese territory very close to the
border and may be certainly found in Laos.
3. G. punctata Aver.,
2006, Rheedea, 16, 1: 7, g. 6, 7 c-d.
Described from southern Vietnam (“VIET-
NAM, Lam Dong Prov., Lac Duong Distr., Da Chais
Municipality, territory of Bi Doup – Nui Ba Nation-
al Park, around point 12º06’44’N, 108º39’27’’E…
Western slope of Bi Doup mt.”). Type (“15 Octo-
ber 2005, L. Averyanov, P.K. Loc, J. Regalado et al.,
HLF 5237a”) – HN (holotype), LE (isotype).
72 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Tubers cylindrical, 2–3 cm long, 5–8 mm
thick, grey-brown, hairy, covered with small scales.
Stem 15–35 cm tall, 2–3 mm thick, soft, dark brown,
with 3–4 broad, ovate, obtuse, bracts, upper bract
usually much longer, to 4 cm long, at the base with
few, weak, roots. Rachis 4–8 cm long, with 4–10
laxy owers. Floral bracts black, cuneate-falcate,
acute, 2–3 mm long. Pedicel and ovary 3–5 mm
long, ovary 2–2.5 mm thick, shortly cylindric, dark
brown. Flowers urceolate, hardly opening, eshy.
Sepals brown, with numerous large white spots,
oblong-ovate, 8–14 mm long, 2.5–4 mm wide, with
eshy, obtuse tips, connate with petals for more
than 3/4 of their length; lateral sepals connate mutu-
ally for about 2/3. Petals thin, connate to sepal tube
on about 3/4 of their length, its free part 1.5–2 mm
long, 0.8–1.2 mm wide, ovate, acute, narrowing to
the base. Lip narrowly ovate, at, 5.5–6.5 mm long,
3–3.5 mm wide, entire, obtuse, with 2 spherical calli
at the base, nely undulate toward apex, disc with
2 low eshy keels at basal part and 2 similar keels
toward the apex. Column 5.5–6 mm tall, 3.5–4 mm
wide, straight, at, very thin, distinctly broadening at
the middle, at apex with narrowly triangular, acute ste-
lidia, at the base with eshy falcate stigma and ovate
thickening lying along median vein. Anther cap hemi-
spheric, about 0.8–1 mm across. Fig. 38; 44 c, d.
Ecology. Primary and secondary evergreen
broad-leaved shady forests on shale and granite at
elev. 1800–2000 m a.s.l. Fl. October – November.
Very rare (VU).
Distribution. Vietnam (Lam Dong). Ende mic.
Studied specimens. Species is known till
now by type collection only.
Note: The species is probably close to
G. dye riana King et Pantl. reported from Bhutan,
Nepal and NE. India, but differs in dark, olive-brown
silvery-white spotted owers, in 4 keels on the lip and
in prominent falcate stigma at the base of column.
4. G. taiensis Tuyama,
1941, Journ. Jap. Bot. 17, 10: 579, g. 1 a,
b; Seidenf., 1992, Opera Bot. 114: 74, g. 38; Aver.,
1994, Ident. Guide Vietnam. Orch.: 82; Aver. et
Averya nova, 2003, Updated Checklist Orch. Viet.:
37; P.H. Ho, 2000, Ill. Fl. Vietnam, 3: 795, g. 10940
(sub Gastrodia “taliense”).
Described from northwestern Vietnam (“Cha-
pa’’). Type (“Hayata s.n., June 23 1913”) – TI?
Tuber fusiform, about 1.5 cm long and 5
cm in diam., pubescent, with small scales. Stem
erect, eshy, glabrous, 16–29 cm tall, 0.7–1.2 mm
in diam., with 2–3 slender roots at the base and few
distant tabular scale-like sheaths. Rachis 1.1.5 cm
long, with 2–3 owers. Floral bracts membrana-
ceous, broadly lanceolate to narrowly obovate,
acuminate, 3.5–4 mm long, 2 mm wide. Pedicel su-
berect, 4.5–5.5 mm long. Ovary narrowly obovoid,
2 mm long, 1.2 mm in diam. Flowers campanulate,
light yellowish to almost white; sepals and petals
connate to about 4/5 of their length forming broad
tube 1.1–1.2 cm in diam., free tips broadly trian-
gular to oblong, obtuse. Sepals 1.6–1.7 cm long,
5–6.5 mm wide, free tips about 3.5 mm long, 5 mm
wide. Petals little shorter, 1.3–1.4 cm long with free
tips 2.5–3 mm long, 2–4 mm wide. Lip narrowly
ovate to oblong, slightly recurved, 5–6 mm long,
2.5–2.8 mm wide, cuspidate with narrowing apex,
in apical part along median vein with short ridge-
like papillose callus, at the base with 2 small hemi-
spheric bosses. Column clavate, 5.5–6 mm long, 1.3
mm wide. Fig. 39.
Ecology. Primary broad-leaved evergreen
shady forests on silicate soils at elev. 1500–1600 m
a.s.l. Fl. June. Very rare (EN).
Distribution. Vietnam (Lao Cai). Endemic.
Studied specimens. Species is known till
now by type collection only.
Note: Insufciently known endemic species
recorded from type collection only. Diagnosis and
gure 39 are copied from original description. Au-
thor in his description said “This species is nearest
to G. dyeriana King et Pantl., but the owers are
larger and the lip is different in shape”.
5. G. major Aver.,
2006, Rheedea, 16, 1: 7, g. 3 h, 5, 7 a-b.
Described from southern Vietnam (“VIET-
NAM, Dak Lak Prov., Krong Bong Distr., Hoa Son
Municipality, vic. of Krong Kmar river, territory of
Chu Yang Sin national park, NE slopes of Chu Yang
Sin mt. system at elev. 700–900 m., around point
12º27’’55’’N, 108º19’00’’E…”). Type (“28 Octo-
ber 2005, L. Averyanov, T.V. Thao, N.T. Vinh, HLF
5444”) – HN (holotype), LE (isotype).
Tubers cylindrical, 5–8 cm long, 1.5–3 cm
in diam., hairy, with many acuminate scales. Stem
rigid, straight, olive-brown, 25–60 cm tall, 3–6 mm
thick, with 5–8 broad, ovate, obtuse distant bracts
0.5–2 mm long, and few close, imbricate, narrowly-
triangular acute scales at the base, without roots.
Raceme 4–15 cm long, with 4–10 owers. Floral
bracts cuneate, 1–1.5 cm long. Pedicel and ovary
1.5–2 cm long, ovary 3–3.5 mm in diam., triangu-
lar in section. Flowers urceolate, eshy, with honey
frag rance; ower tube outside olive-brownish, ne-
73
Turczaninowia 2011, 14(2) : 15–100
Fig. 35. Digital herbarium specimen of Didymoplexiella denticulata (Averyanov L. HAL 11443, epitype).
74 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 36. Didymoplexiopsis khiriwongensis: a-c (HAL 7246); Didymoplexis vietnamica: d-f (HLF 6673); Yoania
prainii (CBL 2091); Gastrodia theana (HAL 7165, type).
75
Turczaninowia 2011, 14(2) : 15–100
ly rugose, with numerous small, white warts. Sepals
olive-brown outside, white or pinkish inside, similar,
oblong-ovate, 2–2.5 cm long, 5–7 mm wide, eshy,
connate from base for about 3/4 of their length, free
apices obtuse, irregularly crenulate and wavy along
margin. Petals thin, yellow-orange, connate to sepal
tube for about 4/5 of their length, free apices 4–5 mm
long, 3–3.5 mm wide, triangular-cordate, acute. Lip
white, entire, 7–11 mm long, 4–5.5 mm wide, circu-
lar to ovate with attenuate, obtuse, recurved, bright-
ly orange, eshy apex, with 2 ovate calli at the base;
disc 5-veined with 2 low eshy keels united toward
the apex into thick eshy rectangular callus. Co-
lumn 9–11 mm tall, 3.5–4 mm wide, straight, eshy,
white, light greenish at the base, with orange-brown,
wing-like, straight, narrow stelidia on lateral sides
at the apex. Anther cap white, hemispheric, 1.8–
2.2 mm across; stigma round, eshy at the base of
column. Capsule erect, cylindric, nely tuberculate,
elongating to 6 cm long. Fig. 40; 44 e, f; 50.
Ecology. Secondary evergreen broad-leaved,
shady forests with bamboo on granite at elev. 700–
900 m a.s.l. Fl. October – November. Very rare (VU).
Distribution. Vietnam (Dak Lac). Endemic.
Studied specimens. Species is known till
now by type collection only.
Subtrib. 5.3.2. Epipoginae Schltr.,
1926, Notizbl. Bot. Gart. Berlin, 9: 571 (sub
“Epipogoneae”). – Ttrib. Epipogieae Parl., 1858, Fl.
Ital. 3: 388 (sub “Epipogoneae”).
Type: Epipogium J.G. Gmel. ex Borkh.
2(2) genera and 4(2) species. Europe, tropi-
cal Africa, temperate, subtropical and tropical Asia,
NE. Australia.
Stereosandra Blume,
1856, Mus. Bot. 2, 9–12: 176; Seidenf.,
1978, Dansk Bot. Ark. 32, 2: 173–174; Comber,
1990, Orch. Java: 86–87; id., 2001, Orch. Sumatra:
118; Seidenf., Wood, 1992, Orch. Malay. Sing.: 142–
143; Su Horng-Jye, 2000, Fl. Taiwan 5: 1036–1039;
Pearce et Cribb, 2002, Orch. Bhutan: 582; Aver. et
Averyanova, 2003, Updated Checklist Orch. Viet.: 56;
Chen Sing-chi, Gale, Cribb, 2009, Fl. China, 25: 207.
Type: S. javanica Blume
Terrestrial leaess, achlorophyllous delicate
herbs with ovoid to oblong tuberculate eshy under-
ground tubers. Stem erect, glabrous, slender, with
few, distant, lanceolate scarious bracts, white with
violet tint. Inorescence terminal raceme with few
to many ephemeral owers. Floral bracts cuneate-
lanceolate, membranous. Flowers pendulous, resu-
pinate, not opening widely, white with violet tips of
tepals. Ovary ovoid. Sepals and petals free, similar,
lanceolate, connivent. Lip similar to petals, entire,
concave, with incurved margins, ecalose, with small
spherical glands at base, not spurred. Column short,
stout, eshy, without column foot. Anther cap large,
erect, eshy, on broad lament arising from dorsal
side of column; pollinia 2, granular, with slender
common caudicle and attached to small common
viscidium. Stigma at the apex of column, entire or
slightly lobed, continuous with rostellum. Capsule
pendulous, ovoid to ellipsoid, maturing rapidly.
One species. SE. Asia to Pacic Islands.
S. javanica Blume,
1856, Mus. Bot. 2, 9–12: 176; Seidenf., 1978,
Dansk Bot. Ark. 32, 2: 173, g. 107; Comber, 1990,
Orch. Java: 86, g.; Seidenf., Wood, 1992, Orch.
Malay. Sing.: 143, g. 60; Su Horng-Jye, 2000,
Fl. Taiwan 5: 1038, g. 444; Aver. et Averyanova,
2003, Updated Checklist Orch. Viet.: 56; Shih-Wen
Chung, 2008, Orch. Taiwan 2: 177, photos.
Described from Java (“Bantam”). Type
(“Has
selt s.n.”) – L.
Stem white, often with violet streaks, 15–
30 cm tall, with 2–4 distant, lanceolate to narrowly
cuneate scarious scales and several yello wish papy-
Fig. 37. Gastrodia tonkinensis (HAL 5263, type):
a – owering plant, b – dissected attened ower, c – at-
tened lip.
76 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
raceous bracts clustered at the base. Tuber spherical
to ovoid, 1–2 cm long. Rachis slender, few to many-
owered. Floral bracts erect, narrowly cuneate,
acute, 3–10 mm long, less than 1 mm wide, shorter
than ovary. Flowers not widely opening, pendulous,
white, all tepals deep violet to the apex. Pedicel and
ovary about 1 cm long. Sepals and pe tals connivent,
similar, lanceolate, 5–9 mm long, 1–2 mm wide, with
acute diverging tips. Lip lanceolate, acute, as long as
sepals, little wider, concave, with incurved margins
and 2 small wart-like calli at base. Column white,
short, about 2 mm tall. Anther on broad lament,
erect, cucullate, about 2 mm tall. Stigma terminal.
Capsule white with dirty purple-violet streaks, pen-
dulous, ellipsoid to 1.5 cm long. Fig. 41; 44 g, h.
Ecology. Primary shady broad-leaved ev-
ergreen forests on silicate soils at elev. 700–800 m
a.s.l. Very rare. Fl. April – May (EN).
Fig. 38. Gastrodia punctata (HAL 5237a, type): a – owering plant, b – ower bud, c – dissected attened
ower tube, d – attened lip, e – column and lip, f, g – section of ovary.
77
Turczaninowia 2011, 14(2) : 15–100
Distribution. Vietnam (Thua Thien-Hue).
NE. India, S. China, Taiwan, Japan, Thailand, Mal-
acca Peninsula, Indonesia, New Guinea, Philippines,
Pacic islands.
Studied collections. Thua Thien-Hue, A
Luoi, HAL 7380 (HN, LE); d-EXSICCATES OF
VIETNAMESE FLORA 0019/ HAL7380.
Notes. Widespread but very rare element of
primary forests with intact soils rich in humus.
Epipogium J.G. Gmel. ex Borkh.,
1792, Tentam. Disp. Pl. German.: 139; Sei-
denf., 1978, Dansk Bot. Ark. 32, 2: 171–172; id.,
1992, Opera Bot. 114: 75–76; Comber, 1990, Orch.
Java: 86; id., 2001, Orch. Sumatra: 116–117; Sei-
denf., Wood, 1992, Orch. Malay. Sing.: 141–142;
Aver., 1994, Ident. Guide Vietnam. Orch.: 83;
P.H. Ho, 2000, Ill. Fl. Vietnam, 3: 795; Su Horng-
Jye, 2000, Fl. Taiwan 5: 861–863; Pearce et Cribb,
2002, Orch. Bhutan: 33–36; Aver. et Averyanova,
2003, Updated Checklist Orch. Viet.: 32; Newman
et al, 2007, Checkl. Vasc. Pl. Lao PDR: 267; Schui-
teman et al., 2008, Nord. Journ. Bot. 26: 289; Chen
Sing-chi, Gale, Cribb, 2009, Fl. China, 25: 207–
208. – Epipogum Rich., 1817, Mem. Mus. Nat. Hist.
Nat. 4: 36. – Galera Blume, 1825, Bijdr.: 415. – Epi-
pogon C. Patze et al., 1850, Fl. Prov. Prenssen: 93;
Gagnep., 1934, Fl. Gen. Indo-Chine 6, 5: 585–586.
Type: E. aphyllum Sw.
Terrestrial leaess, achlorophyllous herbs
with spherical, ovoid or coralloid underground e-
shy tubers. Stem erect, glabrous, eshy, with short,
scale-like sheathing scarious bracts at nodes, white
or pale yellowish. Inorescence terminal raceme
with few to many ephemeral owers. Floral bracts
ovate-lanceolate, membranous. Flowers pendulous,
resupinate or not, white or yellowish, often with
violet or purple markings. Ovary subspherical. Se-
pals and petals free, similar, lanceolate. Lip ovate
to broadly ovate, concave, spurred at base, entire or
3-lobed; disk papillose, sometimes with longitudinal
ridges or lamellae; spur broad, obtuse, shortly sac-
cate to elongate, straight or slightly curved. Column
short, eshy, with no column foot. Anther erect;
pollinia 2, each deeply cleft, clavate, farinaceous,
each with a slender caudicle attached to viscidium.
Stigma placed at base of column; rostellum broad,
sometimes reduced and obscure. Capsule ovoid to
ellipsoid, usually maturing rapidly.
3(1). Europe, tropical Africa, temperate,
sub tropical and tropical Asia, NE. Australia.
E. roseum (D. Don) Lindl.,
1857, Journ. Linn. Soc. London (Bot.), 1:
177; Aver. 1988, Prelim. List Vietnam. Orch. 1: 166;
id., 1994, Ident. Guide Vietnam. Orch.: 83; Sei-
denf., 1978, Dansk Bot. Ark. 32, 2: 171–172, g.
106; id., 1992, Opera Bot. 114: 76; Comber, 1990,
Orch. Java: 86, g.; id., 2001, Orch. Sumatra: 116,
gs.; Seidenf., Wood, 1992, Orch. Malay. Sing.:
141, g. 59; P.H. Ho, 2000, Ill. Fl. Vietnam, 3: 795,
g. 10941; Pearce et Cribb, 2002, Orch. Bhutan: 34,
g. 6; Aver. et Averyanova, 2003, Updated Check-
list Orch. Viet.: 32. – Limodorum roseum D. Don,
1825, Prodr. Fl. Nepal.: 30. – Galera nutans Blume,
1825, Bijdr. 6, g. 3; id., 1825, ibid., 8: 416. – Po-
danthera pallida Wight, 1852, Icon. Pl. Ind. Orient.
5, tab. 1759. – Epipogon nutans Rchb., 1857, Bon-
pl., 1857: 36; Gagnep., 1934, Fl. Gen. Indo-Chine 6,
5: 585, g. 56, 2–5; King et Pantl., 1898, Ann. Roy.
Bot. Gard. Calcutta, 8: 253, tab. 335.
Described from Nepal (“in Nepalia”). Type
(“Wallich s.n.”) – BM.
Stem white to yellowish, 10–45 cm tall, with
few distant amplexicaule, white, scarious sheaths to
1.5 cm long, with terminal, few to many-owered
loose raceme. Tuber spherical or ovoid to 5 cm long
and 2 cm in diam., many nodded. Rachis to 15 cm
long, sometimes pendulous toward apex; Floral
bracts membranaceous, hyaline, ovate to lanceolate,
to 12 mm long. Flowers white to light yellowish,
resupinate, pendulous, widely opening, white, of-
ten with ne purple spots on lip. Pedicel 3–6 mm
long; ovary ovoid to almost spherical, 4–6 mm long.
Sepals and petals weakly spreading, lanceolate,
8–11 mm long, with 3 veins, acute. Lip white to yel-
lowish, often with ne purple marks, entire, ovate,
Fig. 39. Gastrodia taiensis (Hayata, s.n., type):
a – inorescence with sagittal section of owers, b – at-
tened lip.
78 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
Fig. 40. Gastrodia major (HAL 5444, type): a – owering plant, b – dissected ower with attened tepals, c –
attened lip, d – lip, side view, e, f – column, side and frontal view, g – operculum, wiev from below and from above,
h – oral bract, i – section of ovary and oral bract, j – pedicel and ovary, k – ovary cross section.
79
Turczaninowia 2011, 14(2) : 15–100
Fig. 41. Stereosandra javanica (HAL 7380): a – owering plant, b – ower with reexed sepals and petals,
c – intact ower, d – column and lip, e – column, frontal view.
80 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
often with hardly attenuate apex, concave, 8–12 mm
long, 7–8 mm wide, spurred, in apical part nely erose-
denticulate; disk with 2 longitudinal sparsely papil-
lose stripes; spur nearly parallel to ovary, cylindric,
3–4 mm long, equal or little shorter than ovary, apex
slightly bid. Column gibbose, 2.5–3.5 mm tall; anther
cap curved or subglobose, attenuate. Capsule ovoid or
obovoid, 5–7 mm in diam. Fig. 42; 44 i; 51 a.
Ecology. Primary humid shady broad-
leaved or mixed evergreen forests on any kind of
soils at elev. 300–1700 m a.s.l., commonly in small
depressions with rich humus accumulations. Rare.
Fl. March – June (VU).
Distribution. Vietnam (Bac Kan, Cao
Bang, Ha Tinh, Lai Chau, Lam Dong, Nghe An,
Quang Binh, Quang Tri, Thua Thien-Hue). Tropical
areas of Africa and Asia, NE. Australia.
Studied collections. Da Tan La and Man-
line, Dalat, Tixier s.n. (P); Bac Kan, Cho Don, HAL
4827 (HN, LE); Bac Kan, Na Ri, HAL 5404 (HN,
LE); Cao Bang, Tra Linh, VH 4861 (HN, LE); Ha
Tinh, Huong Son, HAL 5214 (HN, LE); Lai Chau,
Sin Ho, NTH 2786 (HN, LE); Nghe An, Tuong
Duong, HLF 6674a (HN, LE); Quang Binh, Minh
Hoa, HAL 11806 (HN, LE); d-EXSICCATES OF
VIETNAMESE FLORA 0139/HAL 11806; Quang
Tri, Huong Hoa, HLF 6291 (HN, LE), HLF 5760
(HN, LE); Thua Thien-Hue, Phu Loc, Bach Ma Na-
tional Park, HLF 1830 (HN, LE).
Notes. Widespread, but rather rare element
of primary forests with intact soils rich in humus.
Peloric forms of this species with subactinomorphic
owers having spurless lanceolate lip similar to pet-
als were observed in Quang Tri Province (Huong
Hoa, HLF 6291, HLF 5760).
Trib. 5.4. Nervilieae (Schltr.) Dressler,
1990, Lindleyana 5, 2: 124.
Subtrib. 5.4.1. Nerviliinae Schltr.,
1926, Notizbl. Bot. Gard. Berlin, 9: 571.
Type: Nervilia Gaudich.
1 genus and 80(5) species. Tropical areas of
Africa and Asia.
Nervilia Gaudich.,
1827, in Freycinet, Voyage Uranie Physicien.
Bot., tab. 35; id., 1829, ibid.: 421, nom. conserv.;
Seidenf., 1978, Dansk Bot. Ark. 32, 2: 147–170; id.,
1992, Opera Bot. 114: 74–75; Comber, 1990, Orch.
Java: 77–81; id., 2001, Orch. Sumatra: 121–126;
Sei denf., Wood, 1992, Orch. Malay. Sing.: 134–
137; Aver., 1994, Ident. Guide Vietnam. Orch.: 80–
82; P.H. Ho, 2000, Ill. Fl. Vietnam, 3: 793–794; Su
Horng-Jye, 2000, Fl. Taiwan 5: 976–981; Pearce et
Cribb, 2002, Orch. Bhutan: 56–63; Aver. et Averya-
nova, 2003, Updated Checklist Orch. Viet.: 45–46;
Newman et al, 2007, Checkl. Vasc. Pl. Lao PDR:
274–275; Schuiteman et al., 2008, Nord. Journ. Bot.
26: 257–316; Chen Sing-chi, Gale, 2009, Fl. China,
25: 197–201. – Pogonia Andrews, 1801, Bot. Rep.,
tab. 212, p.p.: Gagnep., 1934, Fl. Gen. Indo-Chine,
6, 5: 629–630.
Type: N. aragoana Gaudich.
80(5) species. Tropical areas of Africa, Asia,
Australia and Pacic islands.
Terrestrial tuberiferous herbs. Tubers under-
ground, globose or ovoid, white, eshy, of several
nodes, with a few short roots at nodes or rootless,
often bearing few lateral runners during vegetation.
Runners produce new tubers at apex. Floriferous
stem and leaf arise from apex of tuber in succession.
Leaf erect, petiolate, solitary, usually developing af-
ter anthesis, cordate, orbicular, reniform, or elliptic,
with cordate base, palmately veined, plicate, mem-
branous or eshy, glabrous or pubescent. Inores-
cence hysteranthous, erect, terminal, glabrous, 1- to
many-owered; peduncle with 1 to several tubular
sheaths. Floral bracts, lanceolate, acute, green to
scarious. Flowers erect or pendent, resupinate or
not, opening widely or not. Pedicel usually becom-
ing erect and elongating in fruit in 1-owered spe-
cies. Sepals and petals free, similar, narrowly ovate
to lanceolate. Lip entire or 3-lobed, base spurless or
spurred, disc often hairy. Column elongate, clavate,
slender, wingless. Anther terminal, usually ovoid.
Pollinia 2, cleft, granular, sectile; caudicle absent;
viscidium reduced or prominent; rostellum broad,
entire, sometimes absent; stigma orbicular or oblong.
Notes. Species of this genus usually develop
owers and leaves at different times hence it is not
always easy to match the leaves with the owers,
particularly in herbarium collections. As a result, tax-
onomy of this group remains complicated. Flo wers of
all species are ephemeral and often overlooked during
eld surveys. It is quite possible that in Vietnam more
species can be found than were reported till now.
Key to species
1. Leaves present during anthesis, plant blossoms having fully developed leaves; leaves more or less eshy,
dark green; owers 5–6 cm in diam., sepals white, longer than 25 mm ................................. 1. N. marutana
81
Turczaninowia 2011, 14(2) : 15–100
Fig. 42. Epipogium roseum (HAL 5214): a – owering plant, b, c – ower, half side and side views, d – attened
sepals, petals and lip, e – column and spur, frontal and side views.
82 Averyanov L.V. The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae
(primitive tribes – Neottieae, Vanilleae, Gastrodieae, Nervilieae)
- Leaves develop after owering and completely die before next ower formation, plant blossoms without
leaves; leaves thin, not eshy, usually light green, sometimes with dirty purple blotches or velvety dark green
to brown; owers less than 5 cm in diam., sepals yellowish, greenish to brown, less than 25 mm long ....... 2
2. Inorescence 1-owered; leaves shallowly crenulate-polygonal, 1–5 cm across, commonly with 5 main
veins ................................................................................................................................................................ 3
- Inorescence with 2 or many owers; leaves indistinctly irregular crenulate or more or less straight along
margin, more than 5 cm across, with numerous main veins ........................................................................... 4
3. Flowers erect, widely opening; lip apex irregularly lacerate or mbriate; leaves 2–5 cm across, slightly
eshy, sparsely setulose above .............................................................................................. 2. N. crociformis
- Flowers nodding, not widely opening; lip apex with entire, glabrous margin; leaves 1–2.5 cm across,
membranaceous, glabrous ........................................................................................................... 3. N. gracilis
4. Inorescence normally 2-owered, 5–15 cm tall; lip glabrous; leaves velvety dark green to dark purple-
brown, setulose above, particularly along veins and along leaf margin; petiole 2–5 cm tall ....... 4. N. plicata
- Inorescence normally with more than 2 owers, 15–40 cm tall; lip hairy along median vein and at apex;
leaves grassy green, sometimes with large irregular dirty purple blotches, glabrous; petiole 5–20 cm tall ........
................................................................................................................................................... 5. N. aragoana
1. N. muratana S.W. Gale et S.K. Wu,
2008, Makinoa, n. ser., 7: 81. – N. macro-
glossa auct. non (Hook. f.) Schltr., 1911: Aver. et
Averya nova, 2003, Updated Checklist Orch. Viet.: 45.
Described from S. Yunnan (“China, Yun-
nan, Hekou County, Kuzhudong, alt. 200–500m
a.s.l. …”). Type (“1 March 1994, S.K. Wu, Y.P. Yang,
J. Murata & T.Kawahara 870”) – KUN.
Tuber whitish, subglobose, 5–10 mm in
diam., 3–5-noded. Leaves dark green, slightly
eshy, cordate, 4–8 cm across, broadly indistinctly
crenulate, glabrous, with 5–7 main veins, apex acute
to shortly acuminate; leaf of single plant long li-
ving, commonly remains during owering. Petiole
erect, 3–6 cm, long, with 1–2 short, broad, tubular,
scarious sheaths near base. Inorescence 6.5–10 cm
tall, 1-owered; peduncle with 2 tubular acuminate
sheaths 1–2 cm long. Floral bracts broadly lanceo-
late, acuminate, 6–7 mm long. Flower suberect,
later nodding, resupinate, widely opening, 5–6 cm
in diam. Pedicel and ovary light yellow-brownish,
7–9 mm long. Sepals and petals similar, pure white,
tepals sometime with nest brownish marks outside,
lanceolate, acuminate, slightly recurved, 28–32 mm
long, 3–4 mm wide, petals little shorter. Lip white
with irregular pinkish purple blotches on median
lobe, narrowly obovate, 2.4–2.6 cm long, 8–12 mm
wide, spurless, 3-lobed near middle; lateral lobes
rounded, erect, loosely embracing column; median
lobe ovate to triangular-cordate or almost round,
10–12 mm long, 9–12 mm, acute or almost rounded,
slightly reexed; disk with shortly haired thickened
median band. Column clavate, erect, white, 10–
12 mm tall, nely hairy along lateral sides; stigma
round; rostellum transversely elliptic, large. Fig. 45;
51 b, c.
Ecology. Primary broad-leaved and conif-
erous humid shady forests on rocky limestone at
elev. 700–1200 m a.s.l. Fl. March – April. Very rare
(EN).
Distribution. Vietnam (Hoa Binh, Nghe An,
Quang Binh). S. China (S. Yunnan).
Studied specimens. Hoa Binh, Mai Chau,
HAL 883 (HN, LE); Nghe An, Tuong Duong, HLF
6879 (HN); Quang Binh, Bo Trach, HAL 6191 (HN);
Quang Binh, Minh Hoa, HAL 12510 (HN, LE).
Notes. Species closely related to “E. Hima-
layan” Nervilia macroglossa, from which differs in
shorter and much broader lip, distinctly haired along
median vein. Vietnamese plants commonly keep
leaves during anthesis that is rare feature among spe-
cies of the genus. On the territory of China this rare
species was reported from elevations 200–500 m
that is much lower than in Vietnam.
2. N. crociformis (Zoll. et Moritzi) Sei denf.,
1978, Dansk. Bot. Ark. 32, 2: 151, g.
92; Aver. et Averyanova, 2003, Updated Check-
list Orch. Viet.: 45; Shih-Wen Chung, 2008, Orch.
Taiwan 2: 107, photos. – Bolborchis crociformis
Zoll. et Moritzi, 1846, in Moritzi, Syst. Verz.: 89. –
Pogonia prainiana King et Pantl., 1896, Journ. As.
Soc. Bengal, 65, 2: 129; King et Pantl., 1898, Ann.
Bot. Gard. Calcutta, 8, tab. 355. – Pogonia crispata
Blume, 1849, Mus. Bot. 1, 2: 32; Gagnep., 1934,
Fl. Gen. Indo-Chine, 6, 5: 629. – Nervilia crispata
(Blume) Schltr. ex K. Schum. et Lauterb., 1900,
Fl. Schutzgeb. Sudsee: 240; P.H. Ho, 2000, Ill. Fl.
Vietnam, 3: 793, g. 10935. – N. prainiana (King et
Pantl.) Seidenf., 1978, Dansk Bot. Ark. 32, 2: 149,
g. 91; Aver., 1994, Ident. Guide Vietnam. Orch.:
82; P.H. Ho, 2000, l.c.: 794, g. 10939; Seidenf.,
1992, Opera Bot. 114: 74.