Turcz1 08
Turcz1 08
Turcz1 08
582.594 (597)
L. Averyanov ..
Part 1
PREFACE
(Dressler, 1993) with few original additions and amendments. The orchid family system
in used form is presented below as follow:
ORCHIDACEAE Juss.
Subfam. 4. Orchidoideae
Trib. 4.1. Diurideae Endlicher
Subtrib. 4.1.1. Acianthinae Schlecht.
Corybas Salisb.
Subtrib. 4.1.2. Cryptostylidinae Schlecht.
Cryptostylis R. Br.
Trib. 4.2. Orchideae
Subtrib. 4.2.1. Orchidinae
Amitostigma Schlecht., Brachycorythis Lindl., Hemipilia Lindl., Platanthera
L.C. Rich., Vietorchis Aver. et Averyanova
Subtrib. 4.2.2. Habenariinae Benth.
Diplomeris D. Don, Habenaria Willd., Herminium R. Br., Pecteilis Rafin.,
Peristylus Blume
Subfam. 6. Vandoideae
Trib. 6.1. Vandeae Lindl.
Subtrib. 6.1.1. Aeridinae Pfitz.
Acampe Lindl., Adenoncos Blume, Aerides Lour., Arachnis Blume,
Ascocentrum Schlecht., Ascocentropsis Senghas & Schildh., Biermannia King et Pantl.,
Brachypeza Garay, Chiloschista Lindl., Christensonia Haager, Cleisocentron Bruehl,
Cleisomeria G. Don, Cleisostoma Blume, Cleisostomopsis Seidenf., Cryptopylos Garay,
Diploprora Hook. f., Doritis Lindl., Eparmatostigma Garay, Esmeralda Reichenb.,
Gastrochilus D. Don, Grossourdya Reichenb., Holcoglossum Schlecht., Hygrochilus
Pfitz., Kingidium P.F. Hunt, Luisia Gaudich., Malleola J.J. Smith, Micropera Lindl.,
Microsaccus Blume, Ornithochilus Lindl., Papilionanthe Schlecht., Parapteroceras
Aver., Pelatantheria Ridl., Pennilabium J.J. Smith, Phalaenopsis Blume, Pomatocalpa
Breda, Pteroceras Hassk., Renanthera Lour., Rhynchostylis Blume, Rhynchogyna
Seidenf. et Garay, Robiquetia Gaudich., Saccolabiopsis J.J. Smith, Sarcoglyphis Garay,
Schoenorchis Blume, Smitinandia Holttum, Stereochilus Lindl., Staurochilus Pfitz.,
Taeniophyllum Blume, Thrixspermum Lour., Trichoglottis Blume, Tuberolabium
Yamamoto, Uncifera Lindl., Vanda Jones, Vandopsis Pfitz.
short description, total number of species and number of species in the flora of Vietnam
(figures in brackets), as well as the genus distribution.
For each species (subspecies or variety) the following information is reported:
valid name, most significant synonyms and citation of most important recent
monographs,
all available data about type,
short description,
available data on ecology, elevation of observed habitats, phenology, fre-
quency 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 Vietnam (provinces where species
was reported from are mentioned in alphabetical order in brackets ),
list of studied verified specimens and index of their host herbaria,
when necessary short notices on species taxonomy, biology, ecology,
phenology or variation are also provided.
line drawings or (and) color photographs accompany in the book each species
reported for the flora of Vietnam; collecting numbers of plants used as a model for
illustration are cited on drawings or in notices to photographs.
Text of labels is extremely abbreviated whenever being cited. It usually includes
only province name, district name, collectors name and collector number, or, if necessary
date of collection. Largest recent collections are abbreviated and designated as series
with following prefixes:
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 principal investigator Dr. Nguyen Tien Hiep,
CPNP collections in Cuc Phuong national park (commonly without indication
of collectors),
DDS Prof. D.D. Soejarto collections according to International Coope-
rative Biodiversity Groups program with this person as principal investigator,
DKH Dr. D. Harder collections according to expeditions with this person
as a principal investigator,
HAL Dr. Nguyen Tien Hiep, Prof. Leonid V. Averyanov, Prof. Phan Ke
Loc collections in collaborative explorations of these persons,
HLF Henry Luce Foundation, collections of different collectors according
to Vietnam Botanical Conservation Program supported from Henry Luce Foundation,
LX-VN collections of Soviet-Vietnamese Expedition (commonly without
exact indication 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 investigator,
P Prof. Phan Ke Loc collections in expeditions with this person as principal
investigator,
10 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Names of provinces, names of city areas and figures designating these units on the map:
Map 1. Modern administrative division of Vietnam territory (for 2007 year) (except Hoang Sa
and Truong Sa Archipelagos).
12 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Fig. 2. Basic orchid morphology: a, b transversal section of ovary with axial placentation in
Apostasia odorata (a) and parietal placentation in Dendrobium cariniferum (b); c-e main
types of orchid growth system with monopodial shoot (c), sympodial shoot with apical inflo-
rescence (d) and sympodial shoot with lateral inflorescence (e); rings designate flowers, filled
rings designate fruits, lines designate bracts/leaves, arrows show direction of growth, broken
line designate obsolete old part of shoot; f-i main types of orchid inflorescences according
to features of their development: heteranthous in Coelogyne cristata (f), proteranthous in
C. lentiginosa (g), synanthous in Pholidota rubra (h) and hysteranthous (i) inflorescence in
C. fimbriata; j, k types of leaf development (transversal section of developing leaves) with
convolute leaves in Liparis nervosa (j) and duplicate leaves in C. fimbriata (k); l-n types of
orchid leaves (transversal section of mature leaves) with plicate in Tainia viridifusca (l),
conduplicate in Paphiopedilum concolor (m) and laterally flattened leaves in Oberonia
ensiformis (n).
Turczaninowia 2008, 11(1) : 5168 15
Glabrous hairless.
Gland 1 an appendage, protuberance, or other structure which secretes
nectary substances. Fig. 6.
Gland 2 see viscidium.
Globose spherical.
Gynostemium see column.
Hamulus a pollinium stalk which is formed by modified bended rostellum
apex. Fig. 10.
Hastate spear-shaped, with the basal lobes turned outwards. Fig. 11, 12.
Heteranthous an apical inflorescence produced on a separate shoot which
does not develop to produce a pseudobulb and leaves. Fig. 2, f.
Hirsute hairy.
Hispid bristly.
Humus epiphytes epiphytes growing in forks of tree stem on thick
accumulations of humus. Fig. 1, e.
Hyaline glass-like, colorless, translucent.
Hygroscopic capable of absorbing moisture from the atmosphere.
Hypochile the basal portion of a complex lip. Fig. 6, 9.
Hysteranthous an apical inflorescence produced after the pseudobulbs
and leaves have developed. Fig. 2, i.
Imbricate overlapping.
Incumbent lying on or against.
Indehiscent not opening at maturity along definite lines or by pores.
Index (leaf, tepal index) the length of flat figure divided by its width.
Indumentum any covering, commonly refers to hairiness.
Inferior inserted below, like the ovary in the orchids.
Inflorescence the flowering part of a plant, which includes usually floral
axis, floral bracts and flowers. Fig. 3, 59.
Inflorescence bract more or less reduced leaf-like organ bearing an
inflorescence or partial inflorescence (or reduced inflorescence bud) in its axil. Fig. 3, 7.
Internode the section of a stem between two nodes. Fig. 3.
Isthmus a narrow portion of a lip or petal.
Keel a prominent longitudinal ridge, like the keel of a boat. Fig. 6, 8, 9.
Labellum see lip.
Lacerate cut or cleft irregularly, refers usually to the margin of leaf or
tepal. Fig. 11.
Lacinate cut into narrow, irregular lobes or segments, refers usually to the
margin of leaf or tepal. Fig. 11.
Lamella a membrane or septum.
Lamina a blade, the expanded flat portion of a leaf or tepal.
Lanceolate lance-shaped, narrow, tapering toward the apex more than
toward the base; with an index around 47. Fig. 11.
Lateral borne on or near the side of midline of a bilaterally symmetrical organ.
Turczaninowia 2008, 11(1) : 5168 25
Fig. 10. Main types of pollinaria and pollinium stalk in orchids: a, b sagittal section of column
(a) and pollinarium(b) of flower of Tropidia curculigoides, that illustrate hamulus formation;
c, d sagittal section of column (c) and pollinarium (d) of flower of Doritis pulcherrima, that
illustrate stipe (tegula) formation; e, f sagittal section of column (e) and pollinarium (f) of
flower of Dactylorhiza incarnata, that illustrate caudicle formation.
28 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Fig. 11. Main types of shapes and margins of orchid leaf and tepal.
Turczaninowia 2008, 11(1) : 5168 31
Fig. 12. Main types of apices and bases of orchid leaf and tepal.
32 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Reticulate netted.
Retrorse directed backwards or downwards.
Retuse shallowly notched at a rounded apex. Fig. 12.
Revolute rolled back from the margin.
Rhizome horizontal, underground or surface-resting stem bearing scale
leaves and adventitious roots; in sympodial orchids composed of the bases of successive
shoots. Fig. 3, 6.
Rhombic diamond-shaped. Fig. 11.
Rhomboidal quadrangular, nearly rhombic, with obtuse lateral angles.
Root ridge elongate callus on fleshy rhizome which bears tufts of root hairs
in some rootless orchids like species of Cheirostylis.
Root-stem tuberoid see tuberoid.
Rosette a densely clustered spiral of leaves, usually borne near the ground.
Rostellum highly modified, often beak-like sterile median portion of stigma
lying between the functional part of stigma (stigmas) and stamen. Fig. 610.
Rostellar arms lateral narrow wing-like rostellar extensions in some
representatives of subtribe Goodyerinae. Fig. 6.
Rostrate beaked.
Rosulate with the leaves arranged in basal rosettes on very short stem.
Rotund circular in outline.
Round see rounded.
Rounded (round, rotund) with a rounded apex or base. Fig. 12.
Rugose wrinkled.
Rugulose somewhat wrinkled.
Saccate with a conspicuous swelling hollow.
Sagittal section see longitudinal section.
Sagittate arrowhead-shaped. Fig. 12.
Saprophyte plants which obtain nutrients by absorption of organic substances
from the soil (as a rule, indirectly, by means of mycorrhizal symbiotic or mycotrophic
relationships with soil fungi). Fig. 1, b.
Saprophytic see mycotrophic.
Scale any thin, flat, scarious structure commonly of leaf origin. Fig. 3.
Scandent climbing.
Scape a leafless and flowerless lower part of an inflorescence. Fig. 3, 57, 9.
Scarious thin, dry and membranous.
Scattered arranged without apparent order.
Sectile the conditions in which soft, granular pollinia are subdivided into
small packets, which are usually connected by elastic threads.
Secund arranged on one side of the axis only.
Semicircular half-round. Fig. 11.
Sepals in orchids three segments of the outer whorl of perianth (calyx). Fig. 4.
Septum a partition, like in the spur or sac in many species of subtribe
Aeridinae.
Turczaninowia 2008, 11(1) : 5168 33
Serrate saw-like; toothed along the margin, with the sharp, regular teeth
pointing forward. Fig. 11.
Serrulate saw-like; toothed along the margin with minute, sharp, regular
forward-pointing teeth. Fig. 11.
Sessile attached directly, without stalk or petiole.
Sessile leaf leaf lacking the petiole.
Seta (pl. setae) a bristle.
Setaceous bristle-like; with bristles.
Sheath a leaf-like structure which enfolds a stem, pseudobulb, or young
inflorescence.
Sigmoid S-shaped.
Spatulate oblong and attenuated at the base, like a spatula. Fig. 11.
Spicate arranged in a spike.
Spike an unbranched inflorescence with sessile or subsessile flowers
maturing from the bottom upwards.
Spur a slender, tubular or sac-like often nectar-containing projection
commonly formed by the base of the lip. Fig. 6, 7, 9.
Stamen the male reproductive organ of a flower, consisting of an anther
and filament. Fig. 5.
Staminode (staminodium) a sterile stamen, often modified in shape and
size. Fig. 4, 5, 7.
Staminodium see staminode.
Stelidium (pl. stelidia) a wing- or arm-like lateral appendage on the top of
the column. Fig. 8, 9.
Stigma the portion of the pistil which is receptive to pollen. Fig. 46, 810.
Stigma lobes two completely separated lateral lobes of stigma. Fig. 7.
Stigmatophore an elongated process on either side of the base of the
column bearing lateral stigmatic lobe. Fig. 7.
Stipe (tegula) a non viscid band or strap of columnar tissue which connects
the pollinia to the viscidium. Fig. 9, 10.
Stipitate raised on a stalk.
Style the slender part of the pistil which connects the ovary with the stigma;
forms a part of the column. Fig. 4, 5.
Subacute slightly acute. Fig. 12.
Subulate awl-shaped. Fig. 11.
Sulcate with longitudinal grooves or furrows.
Sympodial growth growth in which each new shoot is determinate and
terminates in one season with terminal or lateral inflorescence. Fig. 2, d, e.
Synanthous an inflorescence when pseudobulb, leaf and apical inflorescence
are produced together. Fig. 2, h.
Synsepal (synsepalum) a compound organ formed by the union of two
lateral sepals, like in species of Paphiopedilum. Fig. 5.
Tegula see stipe.
34 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
ACKNOWLEDGEMENTS
Laboratory studies and work with manuscript and illustrations was supported
by Swiss Orchid Conservation fund of Zurich Foundation for Orchid Con-
servation of Swiss Orchid Society. Project Orchids of Vietnam, work on mono-
graph manuscript, 20052007.
We cordially thank authorities of the Institute of Ecology and Biological Re-
sources of the Academy of Science and Technology of Vietnam 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 the next organizations:
U.S.A. National Geographic Society Flora of Highlands of the South
Vietnam 19932001 years (grants # 5094-93, 5803-96, 6383-98); Limestone Flora
of Cao Bang Province of northern Vietnam 19981999 years (grant # 6300-98);
Botanical Inventory of Unexplored Areas in Viet Nam: The North 19992001
(grant # 6733-00); Exploration of rocky limestone flora and vegetation in Bac
Kan province, northern Vietnam 20032005 (# 7577-04); Exploration of highland
flora and vegetation in Lai Chau & Son La Provinces, north-western Vietnam
20062007 (#8074-06)
U.S.A. National Science Foundation Collaborative Research: A Multi-
Taxa Inventory of Threatened Conservation areas in Viet Nam 19992001 (grant
# DEB-9870231).
Henry Luce Foundation. Vietnam Botanical Conservation Program in
Vietnam. 19992006. Preliminary updated checklist of orchids (Orchidaceae)
of Ba Be National Park, 2002; Preliminary updated checklist of orchids (Orchi-
daceae) 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; Preli-
minary 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.
American Orchid Society. Exploration of endangered Vietnamese Pa-
phiopedilums, years 19961997; Exploration of endangered calcium depen-
dent orchid flora in inaccessible rocky limestone areas of the North Vietnam,
years 19992000; Population studies of endemic Paphiopedilum species in nor-
thern Vietnam, 20012002 years; Discovery of endemic orchid flora in remote
limestone areas of Northern Vietnam, years 20032005; Exploration of Vietna-
mese orchid flora in regions allied to Laos territory, years 20082009.
International Cooperative Biodiversity Groups (ICBG) Grant No.
1-UO1-TW01015-01, through funds from the National Institutes of Health, National
Science Foundation, and the Foreign Agriculture Research Service of the United Sta-
tes Department of Agriculture, U.S.A., 20002005.
36 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Author cordially thanks Ms. A. Averyanova and Ms. T. Maisak, who were
very helpful in preparation of line drawings, as well as thanks Prof. Phan Ke Loc for
photographs used in Fig. 24, c; 31, f; 36, d; 43, g; 44, d and Dr. H. Koopowiz for
photograph used in Fig. 26, i.
Turczaninowia 2008, 11(1) : 5168 37
Orchidaceae Juss.
1789, Gen. Pl.: 64 (sub Orchideae). Fam. Apostasiaceae Lindl., 1833,
Nixus Pl.: 22 (sub Apostasieae). Fam. Cypripediaceae Lindl., 1833, l.c.: 22 (sub
Cypripedieae). Fam. Neottiaceae Horan., 1834, Prim. Lin. Syst. Nat.: 50. Fam.
Vanillaceae Lindl., 1835, Key Bot.: 73. Fam. Limodoraceae Horan., 1847, haract.
Ess. Fam. Reg. Veg.: 44.
Typ: Orchis L.
750800(158) genera and 2000025000(1005) species. Cosmopolites, except
extra arid, salted, and polar areas.
1. Plants green, photosynthetic, with normal green leaves, green stem or green roots ....... 2
Mycotrophic (saprophytic), achlorophyllous leafless plants, without any green parts ... 151
2. Flowers with 23 fertile stamens, pollen waxy, amorphous, sticky or powdery, not
forming distinct pollinia ..................................................................................................... 3
Flowers with 1 fertile stamen, pollen mass commonly forming coherent pollinia ........ 5
3. Leaves thin, plicate; all tepals sub-similar, flowers sub-actinomorphic, with 23 stamens;
staminode if present small, inconspicuous; ovary with 3 chambers ............................. 4
Leaves coriaceous, conduplicate; tepals very unequal, lateral sepals joined, lip deeply
saccate; flowers distinctly zygomorphic, with 2 lateral stamens and large shield-like
median staminode; ovary with 1 chamber .................................................... Paphiopedilum
4. Tepals more than 1.5 cm long, ovate; flowers with 3 stamens; inflorescence simple,
erect ........................................................................................................................... Neuwiedia
Tepals less than 1.5 cm long, narrowly-ovate to lanceolate; flowers with 2 stamens;
inflorescence normally branched, curved and spreading .................................... Apostasia
5. Mainly terrestrial and lithophytic plants without pseudobulbs; leaves plicate, thin
(not succulent), not articulated at the vase; inflorescence terminal; anther wall not
caducous; pollinia soft, granular or sectile ...................................................................... 6
Terrestrial, lithophytic and epiphytic plants; shoots sympodial pseudobulbous, or
monopodial thin and rigid; leaves commonly conduplicate, coriaceous or succulent,
usually articulate at the base; inflorescence lateral, rare terminal; anther wall form
caducous cap (operculum); pollinia hard, very rare sectile ............................................ 7
6. Plants with creeping rhizome or underground tubers; anther erect or slightly
incumbent; pollinia apically attached to viscidium; rostellum elongate ....................... 8
Plants with underground root-stem tuberoids; anther erect, at the base firmly united
with column; pollinia sectile with caudiculae basally attached to viscidium or viscidia,
rare without caudicles; rostellum in form of fold protruding between thecae ............ 28
7. Sympodial plants; pollinia soft, waxy, rare cartilaginous, without stalks or caudiculae,
very rare with stipes or hamulus; anther erect in early stages of ontogeny ................ 38
Sympodial or monopodial plants; pollinia solid, cartilaginous or bony, commonly
with stipes; anther incumbent in early stages of ontogeny, often strongly deflexed at
maturity ................................................................................................................................. 85
8 (6). Stem erect, with a close, more or less dense fascicle of roots at the base .................... 9
38 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Stem decumbent, rhizome-like, rooting at the nodes, at the apex ascending and bearing
a terminal long stalked inflorescence .............................................................................. 14
9. All normal leaves radical, the stem only with rudimentary leaves and sheaths .......... 10
Normal leaves distant along all length of the stem, or stem at the middle with 2
opposite leaves ................................................................................................................... 11
10. Flowers less than 5 mm across, resupinate, in dense twisted spike; leaves 310,
narrowly-lanceolate to linear ................................................................................. Spiranthes
Flowers 1.52.5 cm across, not resupinate, in loose not twisted spike; leaves 12 (3),
ovate ....................................................................................................................... Cryptostylis
11. Stems weak, slender, 11.5 mm thick, less than 15 cm tall; at the middle with 2 broadly
cordate opposite leaves 12(3) cm wide ...................................................................... Listera
Stems robust and rigid, more than 1.5 mm thick, more than 15 cm tall; with numerous
distant leaves, rarely with 2 sub-opposite leaves 512 cm wide .................................. 12
12. Stems 1.53 m tall; inflorescence normally branching; sepals and petals 23 cm long
................................................................................................................................ Corymborkis
Stems less than 1.5 m tall; inflorescence simple; sepals and petals 515 mm long ..... 13
13. Flowers, in many flowered spike or in dense heads; sepals and petals white, 0.61 cm
long; lip simple narrowly-ovate, boat-shaped ........................................................ Tropidia
Flowers in lax few flowered raceme; sepals and petals yellow-green or pink-purple;
1.21.8 cm long; lip distinctly 3-lobed, cup-shaped .............................................. Epipactis
14 (8). Flowers not resupinate (or partially resupinate), lip commonly turned upwards ....... 15
Flowers resupinate, lip commonly tuned down ............................................................. 16
15. Leaves uniformly green; lip and column not twisted .............................................. Hetaeria
Leaves with contrast golden or reddish reticulated veins; lip and column distinctly
twisted .......................................................................................................................... Macodes
16. Spur at the base of lip containing neither glands nor hairs ........................................... 17
Spur hairy within or with papillae or glands on either internal lateral side .................. 18
17. Spur short and fat, epichile of lip broad triangular acute without any ornaments;
sepals olive-green, petals and lip white ................................................................ Erythrodes
Spur long and slender, epichile of the lip truncate to slightly bilobed; disc of lip with
two flat calli at the middle and a thick lamella extending to the tip; tepals including lip
white with pink blotches ............................................................................................ Herpysma
18. Spur densely haired within ....................................................................................... Goodyera
Spur with a single gland or group of papillae on either internal lateral side ................ 19
19. Spur with group of dense numerous, long, fat papillae on either internal lateral side;
lip strongly S-bended; mesochile with two tall conspicuous semicircular longitudinal
keels ........................................................................................................................ Orchipedum
Spur with a single gland or few papillae on either internal lateral side; lip not strongly
bended; mesochile without keels, rare with 12 rather low longitudinal keels ........... 20
20. Apex of the lip not widening into a blade, broadly acute, fleshy .................... Vrydagzynea
Apex of the lip widened to a flat thin commonly bilobed blade often dentate or fringed
along margin ........................................................................................................................ 21
21. Mesochile of the lip form distinct conspicuous claw as long as epichile or longer ... 22
Mesochile of the lip very short, appear as constriction between cup-like base of lip
and epichile .......................................................................................................................... 26
22. Claw of lip fringed or toothed along either lateral side .................................................. 23
Claw of lip straight along lateral margins ........................................................................ 24
Turczaninowia 2008, 11(1) : 5168 39
23. Lip spurred, the spur protruding between the lateral sepals; stigma lobes 2, on either
side of base of rostellum; leaves usually deep velvety green, brown or black with
silvery, goldenor reddishnet of veins............................................................ Anoectochilus
Lip with small, globular, saccate base, more or less enclosed by the bases of the
lateral sepals; stigma entire; leaves commonly uniform green, rare plant aphyllous .....
............................................................................................................................... Odontochilus
24. Tiny plants 46 cm tall with leaves about 1 cm long; inflorescence 12-flowered;
column short, not twisted; stigmas on short process ......................................... Myrmechis
Plants normally larger than 6 cm tall with leaves much longer than 1 cm; inflorescence
normally few to many-flowered; column twisted or not twisted; stigmas sessile ...... 25
25. Column and lip distinctly twisted; mesochile without longitudinal keels .............. Ludisia
Lip and column not twisted, rarely slightly twisted; mesochile with 12 more or less
conspicuous longitudinal keels ............................................................................. Rhomboda
26 (21). Leaves black; epichile 3-lobed, brightly yellow, with 4 close callosities on the disc ....
..................................................................................................................................... Zeuxinella
Leaves green to gray-green; epichile 2-lobed, flat, without any ornament ................. 27
27. Sepals free; epichile not toothed or fringed along margin; plant with creeping stem
rooting at nodes ............................................................................................................ Zeuxine
Sepals connate on half way of their length, epichile distinctly toothed or fringed
along margin; plants with succulent creeping rootless articulate rhizome ... Cheirostylis
28 (6). Tiny plants less than 6 cm tall with 1 cordate leaf and single large flower; lip with 2
short spurs; lateral sepals filiform, much narrower than large broad median sepal;
pollinia without caudicles ........................................................................................... Corybas
Plants regularly taller than 6 cm, with 1 to many leaves; inflorescence with few to
many flowers, rare with 1 flower; lip with 1 spur or sac, rarely spurless; all sepals
ovate to elliptic, subsimilar; pollinia with caudicles and basal viscidia ....................... 29
29. One or two stigmas distinctly concave ........................................................................... 30
Stigmasconvex or stalked, rarelyhardlyconcave........................................................ 32
30. Many-leaved stem with distant leaves; floral bracts larger than flowers, similar to
normal leaves; tuberoids palmate or attenuate ............................................ Brachycorythis
Plant with 12(3) radical leaves; floral bracts inconspicuous, shorter than leaves;
tuberoids attenuate to globose ........................................................................................ 31
31. Lip deeply 3-lobed; plant with 12(3) ovate to narrowly ovate, uniformly green leaves;
stigma divided into 2 lobes; rostellum very small inconspicuous; tuberoids attenuate
.................................................................................................................................. Amitostigma
Lip entire or very indistinctly trilobe; plant with 1(2) circular to broadly ovate, purple-
violet variegated leaves; stigma entire, V-shaped; rostellum large elongate, horn-
shaped, curved upwards; tuberoids globose ....................................................... Hemipilia
32 (29). Rostellum very large, horn-shaped, turned upward, as high as or higher than anther ......... 33
Rostellum much smaller, inconspicuous, shorter than half of column length, sometimes
hardly visible ....................................................................................................................... 34
33. Plants 610 cm tall with 12 narrowly-lanceolate leaves; inflorescence normally with
1(2) flowers; flowers white; lip entire or indistinctly 3-lobed ............................. Diplomeris
Plants normally taller than 10 cm with 310 broadly-lanceolate leaves; inflorescence
normally with (1)320 flowers; flowers purple, orange-red to deep yellow; lip distinctly
4-lobed ....................................................................................... Habenaria (H. rhodocheila)
34. Two stigma-lobes placed on elongated more or less long stigmaphores extending
40 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
46. Stem at the base inflated, fleshy, pseudobulbous; lip elastically hinged to the column
and connected with column base with 2 elastic flanges .................................. Plocoglottis
Stem at the base not inflated; lip movably or firmly joined to columnfoot or column
base, not connected with column with elastic flanges .................................................. 47
47. Lip spurred; inflorescence normally with 24(6) flowers; flowers purple-violet, 5
8 cm across ....................................................................................................................... Phaius
Lip without spur or sac at the base; inflorescence normally with more than 10 flowers;
flowers white to yellow-greenish, less than 3 cm across ..................... Cephalantheropsis
48 (43). Stem fleshy, creeping, rooting at nodes with ascending apex and few distant leaves ...... 49
Stem pseudobulbous, short and flesh, with few sub-radical leaves ............................ 51
49. Flowers not resupinate; lip with short spur, turned upward .................... Nephelaphyllum
Flowers resupinate; lip with long slender spur or spurless, turned down .................. 50
50. Lip with a long slender spur; inflorescence 1-flowered ..................................... Hancockia
Lip spurless; inflorescence normally many-flowered ................................. Mischobulbm
51 (48). Plants with underground tuberous rhizome ................................................................... 52
Stem pseudobulbous, green, short and flesh, sometimes covered with leaf sheaths ....... 55
52. Flowers purple-violet, not resupinate; sepals connate, forming a narrow tube held at
right angle to the ovary ...................................................................................... Anthogonium
Flowers white, pinkish to purple-violet and yellow-orange, resupinate; sepals free .... 53
53. Inflorescence terminal; leaves elliptic to ovate ............................................................ Bletia
Inflorescence lateral; leaves linear to lanceolate, rare narrowly elliptic ...................... 54
54. Inflorescence normally many-flowered, dense, secund spike; flowers not widely
opening, less than 1 cm across, light pinkish; each growth normally with 1(2) linear
leaves ...................................................................................................................... Pachystoma
Inflorescence normally few-flowered, all-faced, loose raceme; flowers widely opening,
normally more than 1 cm across, pink-violet or yellow; each growth normally with
several lanceolate to narrowly elliptic leaves .................................................. Spathoglottis
55 (51). Pseudobulbs consist of 1 internode ............................................................................... 56
Pseudobulbs consist of 2 or more internodes ................................................................ 57
56. Pseudobulbs 2-leaved; leaves sessile with broad base; inflorescence, flower bracts,
pedicel, ovary and sepals outside brown hairy ......................................................... Eriodes
Pseudobulbs 1-leaved; leaves petiolate with long distinct petiole, all plant hairless ....
............................................................................................................................................ Tainia
57. Each growth with 12 leaves; sepals connate, forming broad urn-like tube; lip mobile
on a large columnfoot .................................................................................. Acanthephippium
Each growth usually with more than 2 leaves; sepals free; lip not mobile, firmly attached
to the column base ............................................................................................................. 58
58. Column margins over nearly their whole length connate with base of lip .......... Calanthe
Lip only connate with column at or near its base ...................................................... Phaius
59 (41). Lip at base flat or slightly concave .................................................................................. 60
Lip at base saccate ............................................................................................................. 62
60. Lip at base S-curved in lateral view; sepals and petals about equal width .......... Panisea
Lip at the base more or less flat, not S-curved; petals usually narrower than sepals ..... 61
61. Lip entire to obscurely lobed; bulbs and leaves short-living annual; leaves thin,
herbaceous, deciduous; inflorescence usually 1-flowered; lip at base convolute around
base of column ............................................................................................................... Pleione
Lip usually distinctly 3-lobed; bulbs and leaves long-living perennial; leaves rigid,
42 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
normally with several to many leaves sheathing at the base; inflorescence usually
with more than 2 flowers; flowers commonly lasting more than 1 day ........... Dendrobium
72. Inflorescence appears terminal, from the apex of pseudobulb; lip not mobile, not
hinged at base; menthum rather spur-like .......................................................... Epigeneium
Inflorescence distinctly lateral, from the rhizomatous part; lip mobile, movably hinged
to columnfoot; menthum rather saccate ......................................................................... 73
73. Operculum with conspicuous horn at apex; sepals more or less equal size and form,
spreading ............................................................................................................................ Trias
Operculum without horn at apex; lateral sepals commonly different in size and shape
from median sepal, spreading or not ............................................................... Bulbophyllum
74 (69). Stipe very large and massive, as broad as long, with broad apex which occupies
whole clinandrium; inflorescence with 1 hardly opening flower ....................... Hamularia
Stipe narrow and slender, much longer than broad, placed in furrow in front of column
apex; inflorescence many-flowered, rare with 12 widely opening flowers ................ 75
75. Petals fascinated to the basal portion of distally naked columnfoot; columnfoot longer
than column; lip with large falcate erect auricles at base; pollinia with 1 common stipe
................................................................................................................................... Monomeria
Petals fascinated along the whole edge of the columnfoot; columnfoot of equal length
or shorter than column; lip without conspicuous auricles at base; pollinia with 2
stipes .............................................................................................................................. Sunipia
76 (67). Pollinia 6; lip joined to columnfoot, with tall conspicuous appendage on upper surface
toward the base; stem not pseudobulbous with numerous distichous leaves ..... Appendicula
Pollinia 8; lip not firmly joined to columnfoot, without tall appendage on upper surface;
stem pseudobulbous or not ............................................................................................. 77
77. Lip with a sac-shaped hypochile separated from epichile by transversal ridge; stem
not pseudobulbous, rod-like with numerous internodes and distichous grass-like
leaves; flowers very small, less than 2 mm across, in dense terminal head ...... Agrostophyllum
Hypochile not separated from epichile by a transversal ridge; stem pseudobulbous or
not; flowers commonly larger than 2 mm across, not in dense terminal heads ........... 78
78. Pollinia joined by a common long thin caudicle ............................................................. 79
Pollinia not joined by a common long thin caudicle ...................................................... 80
79. Plant with flat normally 2-leaved pseudobulbs, rare stem not pseudobulbous with few
sub-radical distichous leaves; flowers hardly opening, sub-cleistogamous; lip at the
base with a longitudinal thickening; operculum vertical behind column, beaked; column
without columnfoot ..................................................................................................... Thelasis
Plant with several sub-radical distichous leaves; flowers campanulate; lip without
thickening; operculum horizontal on top of column, not beaked; column with small,
but distinct columnfoot ............................................................................................. Phreatia
80. Column with 2 vertical arms or lobules bearing stigmas; stem thin not pseudobulbous
.................................................................................................................................. Ceratostylis
Column without vertical arms or lobules bearing stigmas; stem pseudobulbous or not
.................................................................................................................................................. 81
81. Sepals free; flowers widely opening or campanulate .................................................... 82
Sepals connate, forming tube; flowers hardly opening ................................................ 84
82. Young leaves densely haired ................................................................................ Trichotosia
Leaves not hairy ................................................................................................................. 83
83. Lip movably joined to a columnfoot; column long, forward curved; columnfoot
44 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
distinctly broader than column, in front with large fleshy cushion .................. Callostylis
Lip not very movable, joined to a columnfoot; column short, not strongly curved;
columnfoot not much broader than column, without conspicuous cushion .......... Eria
84 (81). Pseudobulbs elongate, 410 cm tall, normally with 35 leaves; inflorescence 315 cm
long, of many distichous flowers ....................................................................... Cryptochilus
Pseudobulbs flat, less than 1 cm tall, normally with 2 leaves; inflorescence less than
0.5 cm long, normally with 12 flowers ........................................................................ Porpax
85 (7). Sympodial plants ................................................................................................................. 86
Monopodial plants .......................................................................................................... 98
86. Lateral sepals free or partially merged into a synsepalum ......................................... 87
Lateral sepals merged completely into a synsepalum ................................... Acriopsis
87. Stem pseudobulbous; pseudobulbs 1-leaved, consist of 1 internode; column much
curved, apically with 2 forwards curved horns .............................................................. 88
Stems usually of several internodes, pseudobulbous or not, with 1 to several leaves;
column straight or slightly curved, apically without large, conspicuous horns ........ 89
88. Lip 2-lobed at apex with small insignificant median dent; column S-curved; on the
upper side of columnfoot a narrow nectary opening at base of lip; pollinia 2; operculum
semi globular ............................................................................................................. Thecostele
Lip at the apex acute, entire; column bent forwards, but not S-curved; columnfoot
hollow with an entrance near the articulate base of lip; pollinia 4; operculum conical
....................................................................................................................................... Thecopus
89. Plants develop neither pseudobulbs nor tubers, stem thin, not pseudobulbous ...... 90
Plants develop tubers, corms or pseudobulbs of several internodes, or pseudobulbous
stems often covered by close leaf sheaths ..................................................................... 92
90. Terrestrial grass-like plants with erect or climbing stem up to 1 m long; lip densely
haired along midvein ............................................................................................... Dipodium
Epiphytes with stem less than 20 cm tall, not grass-like; lip not haired ....................... 91
91. Leaves linear, grass-like, laterally compressed or strongly conduplicate; inflorescence
dense, not branching distichous spike; flowers resupinate, lasting few hours ... Bromheadia
Leaves lanceolate to ovate, hardly conduplicate or flat; inflorescence lax, often
branching all-facing raceme or panicle; flowers not resupinate, long lasting ......... Polystachya
92 (89). Column at front edge with small horizontal triangular or obtuse wings; anther with 2
pollinia ................................................................................................................................... 93
No wings at front of column; anther with 2 cleft or 4 pollinia ........................................ 95
93. Columnfoot simple, without nectary; lip without claw ...................................... Diglyphosa
Columnfoot with a saccate nectary or a spur-like nectary formed with connate bases
of lateral sepals; lip at base narrowing into a claw ......................................................... 94
94. Columnfoot short, fat, with a saccate nectary opening at base of column between
vertical oblong cushions, lateral sepals not connate at base .................... Chrysoglossum
Columnfoot long forming a slender spur-like menthum together with connate bases
of lateral sepals .......................................................................................................... Collabium
95 (92). Stem inflated of many internodes, more or less pseudobulbous, usually covered by
dense leaf sheaths; anther with 2 cleft pollinia, or pollinia 4 in pairs; pollinarium
without stipes, viscidium large, broad with truncate front edge ..................... Cymbidium
Plants with tubers, corms or epigeous corm-like pseudobulbs of a few internodes;
anther with 2 cleft pollinia or pollinia 4 in pairs; pollinarium with short strap-shaped
stipes, viscidium rather large, sometimes forming a single structure with the stipe ......... 96
Turczaninowia 2008, 11(1) : 5168 45
111. Lip with transversal ridge or rugose callosities at the base of epichile ...................... 112
Lip without transversal ridge or callosities on epichile ............................................... 113
112. Large plants with stem up to 1 m tall and leaves 2030 cm long; hypochile sac-like,
hairy inside ..................................................................................................................... Acampe
Small plants with stem up to 20 cm tall and leaves 510 cm long; hypochile spur-like,
not hairy inside ..................................................................................................... Smitinandia
113. Large plants with climbing stem normally longer than 50 cm; leaves coriaceous, flat,
oblong; flowers red or yellow, 25 cm across; lip much shorter than median sepal ........
................................................................................................................................... Renanthera
Small plant with pendant or creeping stem normally less than 20 cm long; leaves
fleshy, lanceolate to sub-terete; flowers white to purple, less than 5 mm across; lip as
long as or longer than median sepal ................................................................. Schoenorchis
114 (109). Leaves terete; spur back-wall bears inside Y-shaped callus .................... Cleisostomopsis
Leaves dorsiventral; spur back-wall inside bears a rising tongue ............................. 115
115. Rising tongue on the lip usually hairy, placed close to entrance to spur .................. 116
Back-wall rising tongue glabrous, placed deep in spur ................................ Pomatocalpa
116. Inflorescence very short, less than 1 cm long, commonly 12-flowered; flowers less
than 1.5 cm across ................................................................................................ Trichoglottis
Inflorescence normally 1025 cm long; commonly bears more than 2 flowers; flowers
more than 1.5 cm across ...................................................................................... Staurochilus
117 (108). Vine-like plant with climbing or hanging stem and numerous ovate leaves usually
less than 3 cm long; column with 2 long thin glabrous stelidia incurved above apex;
inflorescence very short, less than 1 cm long, 12(3)-flowered ................... Pelatantheria
Stem erect, rare vine-like and ascending, leaves usually longer than 3 cm; column
without large stelidia; inflorescence longer than 1 cm with few to many flowers ..... 118
118. Floral bracts large, longer than flowers; bracts, ovary and flowers densely pubescent
.................................................................................................................................. Cleisomeria
Floral bracts small insignificant, shorter than flowers; bracts, ovary and flowers
glabrous, rarely papillose .................................................................................................. 119
119. Rostellum projection narrow, very long, twice longer than broad of operculum, supports
a thin linear stipe, to 9 times as long as diameter of pollinia ........................................ 120
Rostellum projection shorter; stipe usually shorter than 9 diameters of pollinia ..... 121
120. Vine-like plant with climbing or hanging stem commonly 1530 cm long, rooting along
all its length; leaves distant, coriaceous; rostellum projection distally upwards turned,
stipe linear on all its length ...................................................................................... Micropera
Plants with short erect stem, usually up to 10 cm tall, rooting at the base, leaves
fleshy, distinctly distichous, arranged into 2 dense rows; rostellum projection distally
curved down, stipe linear broadening to the apex ........................................... Stereochilus
121. Clinandrium top with a rising dorsal longitudinal ridge with a furrow along edge
bearing long linear stipe and dorsally placed pollinia .................................... Sarcoglyphis
Column without conspicuous dorsal ridge at the top; pollinia not placed dorsally;
stipe very variable in form .................................................................................... Cleisostoma
122 (105). Sepals and petals broadly ovate; stem erect, straight, semi-woody; leaves usually 3
5 cm broad; lip broadly spatulate at the apex; epichile with high median keel ...... Hygrochilus
Sepals and petals narrowly-oblong, spatulate toward the apex; stem slender, often
climbing, vine-like; leaves less than 3 cm broad; lip acute at the apex; epichile with
low median keel ............................................................................................................ Arachnis
Turczaninowia 2008, 11(1) : 5168 47
123 (104). Lip without distinct spur or sac, but hypochile often more or less concave; leaves
dorsiventral, usually thin, or lacking .............................................................................. 124
Lip with distinct spur or sac; leaves dorsiventral, usually fleshy or terete ............... 125
124. Stipe short and broad, shorter than diameter of pollinia; plants with normal leaves;
stem usually longer than 1 cm, rooting along all its length ............................ Thrixspermum
Stipe linear to strap-shaped, about twice as long as diameter of pollinia; leaves usually
lacking; stem usually less than 1 cm long with densely clustering numerous roots ......
................................................................................................................................. Chiloschista
125. Spur of lip with longitudinal septum .................................................................. Cleisostoma
Spur of lip without longitudinal septum ........................................................................ 126
126. Leaves terete, usually purple ........................................................................... Rhynchogyna
Leaves dorsiventral, usually green, sometimes with purple tint ................................ 127
127. Stem short, normally less than 2 cm long; leaves thin, ovate, 36 cm wide; flowers
more than 1 cm across, in lax few flowered inflorescence normally longer than 5 cm .....
..................................................................................................................................... Kingidium
Stem 420 cm long; leaves lanceolate-oblong, fleshy, less than 1 cm wide; flowers less
than 1 cm across in dense inflorescence 35 cm long .................................. Cleisocentron
128 (103). Anther with 2 pollinia, more or less but not completely cleft or split ........................ 129
Pollen-masses more integrated ......................................................................................... 139
129. Flowers without columnfoot, or columnfoot very indistinct ...................................... 130
Flowers with distinct, but sometimes short columnfoot ............................................. 135
130. Inflorescence lax raceme, commonly with 25 flowers, flowers commonly more than
1.5 cm across ....................................................................................................................... 131
Inflorescence dense spike with numerous flowers; flowers commonly less than 1.5 cm
across .................................................................................................................................... 132
131. Epichile 2-lobed, truncate or obtuse, more or less flesh or rigid, with entire margin;
stipe short and broad ....................................................................................................... Vanda
Epichile entire, broad, semicircular, thin, denticulate along margin; stipe narrowly-
lanceolate ............................................................................................................ Christensonia
132. Flowers 11.5 cm across; robust plants with erect semi-woody stem; leaves large,
very rigid, normally 2040 cm long often with many light-colored veins .... Rhynchostylis
Flowers much less than 1 cm across; plants smaller with erect or pendent stem; leaves
not particularly rigid, normally less than 15 cm long, uniformly green ...................... 133
133. Tinycanopy epiphytes; leaves normally25 cm long; flowers 23 mm across ... Saccolabiopsis
Medium size epiphytes; leaves normally longer than 5 cm; flowers more than 4 mm
across .................................................................................................................................... 134
134. Long stemmed plants with distant leaves; stipe lanceolate to spathulate, slightly
dilated and curved at apex; spur distinctly inflated at the apex ......................... Robiquetia
Rather short stemmed plants with few, more or less close leaves; stipe S-shaped,
rising above and behind pollinia, which are separated by a vertical longitudinal stipe
lamella; spur not inflated at the apex .......................................................................... Uncifera
135 (129). Leaves terete; stem vine-like, slender, climbing ............................................ Papilionanthe
Leaves dorsiventral; stem not vine-like, more or less erect ........................................ 136
136. Mid-lobe of epichile well developed, large, flat, cordate, triangular or oblong;
hypochile flat, concave or form conspicuous forward curved spur .......................... 137
Mid-lobe of epichile reduced in form of small fleshy dent, spur or sac placed centrally
on the lip .............................................................................................................................. 138
48 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
137. Hypochile flat or slightly concave; epichile at the base with high keels, teeth or
thread-like bifurcate appendages; inflorescence with few distichous flowers ... Phalaenopsis
Hypochile form conspicuous forward curved spur; epichile flat without ornaments;
inflorescence with numerous all-faced flowers .......................................................... Aerides
138. Columnfoot as long as or longer than column; rachis of inflorescence not swollen
along whole its length; stem longer than 2 cm, bearing normally several leaves ......... Pteroceras
Column much shorter than column; rachis of inflorescence distinctly swollen toward
the apex; stem very short, less than 2 cm long, bearing normally 13 leaves .... Brachypeza
139 (128). Anther with 2 porate pollinia ........................................................................................ 140
Anther with 2 whole pollinia, not cleft, split or porate ................................................. 147
140. Lip movable; columnfoot as long as or longer than column; lateral sepals attached
with their bases to the basal portion of distally naked columnfoot; spur entrance
partially closed by a horizontal plate arising from the back-wall of the lip .... Cryptopylos
Lip not movable; columnfoot shorter, rarely as long as or longer than column; lateral
sepals attached to columnfoot along whole its length; spur entrance free .............. 141
141. Leaves terete, veryrigid; inflorescence veryshort, commonlyless than 1 cm long ................ Luisia
Leaves dorsiventral, sometimes fleshy conduplicate to sub-terete, but not particularly
rigid; inflorescence commonly longer than 1 cm ......................................................... 142
142. Small canopy epiphytes with leaves normally less than 6 cm; mid-lobe of the lip large,
terete, conical at the apex, fleshy, furrowed and concave toward the base; flowers
distichous, opening in succession and lasting few hours ............................... Biermannia
Medium size epiphytes with leaves commonly longer than 6 cm; mid-lobe of the lip
more or less flat, not terete or fleshy; flowers all-way faced, lasting few days ......... 143
143. No distinct rising side-lobes on the sac-like hypochile, usually just a fleshy rim joining
in front of sac entrance ..................................................................................................... 144
Distinct, but sometimes small, side-lobes rising from lateral sides of hypochile which
forms a saccate to cylindrical spur ................................................................................. 145
144. Inflorescence umbel-like, less than 3 cm long; epichile broadly fastened in front of the
rim connecting edges of the hypochile, usually crescent-shaped, commonly shorter
than broad; the edges of the stigma not swelling ............................................. Gastrochilus
Inflorescence dense raceme much longer than 3 cm; epichile narrow, triangular,
concave to the base connecting by lateral sides the edges of the hypochile; the
edges of the stigma concavity much swelling into protruding ring-like structure .............
.......................................................................................................................... Eparmatostigma
145. Epichile thin, much broadening from a narrow base; leaves strongly conduplicate,
semi-terete; inflorescence lax few-flowered raceme, normally longer than 15 cm. ...............
.............................................................................................................................. Holcoglossum
Epichile more or less fleshy, ligulate-oblong or broadly cuneate, not narrowing toward
the base; leaves distinctly dorsiventral, very rare sub-terete; inflorescence dense,
many-flowered spike, normally shorter than 15 cm ..................................................... 146
146. Epichile ligulate-oblong, more or less flat; lip side-lobes rather straight, no transversal
callus at the base of epichile; stipe broad and rather short, twice longer than diameter
of pollinia .............................................................................................................. Ascocentrum
Epichile broadly-cuneate, concave; lip side-lobes strongly incurved, closing the
entrance of the spur, at the base of epichile large fleshy transverse callus connecting
the front-edges of side-lobes, curving inwards; stipe narrow, longer than twice diameter
of pollinia .......................................................................................................... Ascocentropsis
Turczaninowia 2008, 11(1) : 5168 49
147 (139). Column without distinct columnfoot; stipe linear, much broadening at apex, 3-5 times
as long as diameter of pollinia ........................................................................................ 148
Column with distinct columnfoot; stipe ovate to broadly-lanceolate, not much
broadening at apex, as long as or few longer as diameter of pollinia .......................... 149
148. Lip side-lobes very large, flat, ciliate or finely fringed along front margin .... Pennilabium
Lip side-lobes small in form of insignificant dents, entire along margin ............. Malleola
149. Lip movable, 5-lobed; side-lobes and lateral lobes of epichile long, conspicuous,
erect, longer than broad, mid-lobe very small in form of short fleshy dent ........ Grossourdya
Lip not movable; epichile short, obtuse, entire, very fleshy, lateral lobes insignificant
or broadly oblong to semi-circular as long as broad ................................................... 150
150. Stem usually 48 cm long, erect to suberect, with more or less distant leaves;
inflorescence lax, 58 cm long; lip with broadly-oblong or semi-circular side-lobes;
mid-lobe fleshy, broadly conical .................................................................. Parapteroceras
Stem usually less than 4 cm long, pendulous, with close placed leaves; inflorescence
short, head-like, commonly less than 3 cm long with densely placed flowers opening
in succession; lip very fleshy, with broadly obtuse indistinct lobes .......... Tuberolabium
151 (1). Climbing vines commonly 210 m long ......................................................................... 152
Terrestrial erect herbs commonly less than 2 m tall ...................................................... 153
152. Inflorescence rachis and sepals outside densely pubescent; flowers brightly yellow,
23 cm across, column stout ....................................................................................... Galeola
Inflorescence rachis and sepals outside glabrous; flowers light olive green to dull
pink-whitish, less than 2 cm across, column slender ...................................... Erythrorchis
153. Lateral sepals joined each together for a large part of their length ............................ 154
Lateral sepals free from one another or only joined at the base ................................. 157
154. Sepals and petals joined into 5-lobed tube, split between lateral sepals; flowers hardly
opening; sepals fleshy, light brownish to brownish-pink; stigma at the base of column;
pollinia 2 ..................................................................................................................... Gastrodia
Sepals and petals free or joined less than half of their length, newer form tube; flowers
widely opening; sepals and petals very thin, white or blue-violet; stigma near top of
column; pollinia 2 or 4 ....................................................................................................... 155
155. Column without stelidia; flowers white ........................................................... Didymoplexis
Column near apex with conspicuous long decurved hook-like stelidia; flowers white
or blue-violet ........................................................................................................................ 156
156. Median sepal and petals joined to about half of their length, fused with joined lateral
sepals near their base; columnfoot absent; flowers blue-violet .............. Didymoplexiella
Median sepal and petals free; columnfoot present; flowers white .... Didymoplexiopsis
157 (153). Lip entire, not lobed ....................................................................................................... 158
Lip distinctly divided into 23 lobes .............................................................................. 160
158. Sepals pubescent or mealy outside; sepals and petals yellow to yellow-brown, lip
pale yellow; plants more or less robust, stem reddish-brown to pinkish-brown, rigid;
fruits juicy, cylindrical, berry-like, dull red-purple; tubers numerous, cylindrical, finger-
like .................................................................................................................................. Cyrtosia
Sepals glabrous; sepals and petals white, sometimes with black-violet tips, lip white
with purple tint or purple nerves, sometime with black-violet tip; plants slender, stem
white, soft and weak; fruits dry yellowish-white broadly ovate capsule; tuber one,
ovate to cylindrical-ovate .................................................................................................. 159
159. Sepals and petals white, rare with light pink nerves; lip ovate, white or with light pink
50 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
center or with light purple spots, at the base with short broad spur ................ Epipogium
Sepals, petals and lip white, with black-violet tips; lip narrowly ovate to broadly-
lanceolate, not spurred ...................................................................................... Stereosandra
160 (157). Flowers with a distinct calyculus beneath perianth; lip apex densely long haired;
stem very brittle, dark brown to black, less than 1 mm thick ........................... Lecanorchis
Calyculus beneath perianth absent; lip at the apex glabrous or papillose; stem not
particularly brittle, white, yellowish, olive to yellowish-brown, more than 1 mm thick
.................................................................................................................................................. 161
161. Inflorescence yellowish-brown to olive-brown, normally with more than 5 flowers; lip
distinctly divided into hypochile, mesochile and epichile ......................................... 162
Inflorescence white, normally with 13 (5) or more flowers; lip not divided into
hypochile, mesochile and epichile ................................................................................... 163
162. Plants more than 25 cm tall; apex of the lip acute to broadly acute ................ Aphyllorchis
Plants less than 25 cm tall; apex of the lip distinctly bilobed ........................ Odontochilus
163. Flowers brightly yellow, widely opening; sepals and petals more or less thin, lip side-
lobes flat, reflexed, mid-lobe at the apex broadly truncate; column without stelidia ......
...................................................................................................................................... Vietorchis
Flowers white with purple tint, hardly opening to campanulate; sepals and petals
fleshy, lip side-lobes upward incurved, mid-lobe broadly acute; column at the apex
with large lateral stelidia .................................................................................................. Yoania
Key to varieties
1. Inflorescence glabrous or with very few scattered hairs ....... 1. N. zollingeri var. annamensis
Inflorescence densely hairy .............................................................................................. 2.
2. Flowers brightly yellow; ripe fruit white ................................. 2. N. zollingeri var. javanica
Flowers pure white with yellow central spot on lip; ripe fruit orange to red ........................
...................................................................................................... 3. N. zollingeri var. balansae
1b. N. zollingeri var. javanica (J.J. Smith) de Vogel, 1969, Blumea, 17, 2:
329. N. javanica J.J. Smith. 1914, Bull. Jard. Bot. Buitenzorg 2, 14: 5.
Described from Java ( Artana). Lectotype (J.J. Smith s.n. a. 1900) BO.
Plants up to 40 cm tall, all parts more or less densely haired. Flowers bright yellow.
Dorsal keel on petals and lip densely hairy. Ripe fruit white juicy berry. Fig. 14, a-d.
Ecology. Shady broad-leaved evergreen submontane forests on granite and
sandstone. 600950 m. Fl. March. Rare (LR).
Distribution. Vietnam (Dak Lak, Ninh Thuan). Java, Sumatra, Bali.
Studied specimens. Ninh Thuan, Nui Chua mt HLF 4415 (HN, LE); Dak Lak, Chu
Yang Sin mt, HLF 5479 (HN, LE).
Notes. Ripe berries of this variety spicy-fragrant, sweet and edible.
Gagnep. 1933, Bull. Soc. Bot. France, 80: 350. N. griffithii auct. non Reichenb. f.:
Seidenf. 1992, l.c.: 11; Aver. 1994, Identif. Guide Viet. Orch.: 12; Aver. et Averyanova,
2003, Updated Checklist Orch. Viet.: 46. N. inae auct. non de Vogel: Seidenf. 1992,
l.c.: 13; Aver. 1994, l.c.: 12; Aver. et Averyanova, 2003, l.c.: 46.
Described from N Vietnam (Tonkin: mt. Bavi, ). Lectotype (n 2000
(Balansa)) P.
Plants up to 40 cm tall, all parts more or less densely. Flowers pure white, lip
with yellow spot in center. Dorsal keel on petals and lip densely hairy. Ripe fruit
orange to red juicy berry. Fig. 13, a; 14, e-g.
Ecology. Shady broad-leaved, evergreen lowland and submontane forests of
sandstone, granite and rocky limestone. 4501250 m. Fl. May June. Rare (LR).
Distribution. Vietnam (Dak Nong, Khanh Hoa, Kon Tum; Ninh Binh, Phu
Tho, Quang Ninh).
Studied specimens. Cam Pha, Cong Hoa, Quang Ninh, Tran s.n. (LE, UHN); Ninh
Binh, Cuc Phuong, Duc B 356 (HN), CPNP 5116 (HN), N.M. Cuong, H.L. Quyen, NMC 309
(HN), DDS 11060 (HN), Averyanov et al., HAL 1686 (HN, LE), Duc B 356 (HN), N.T. Hiep, NTH
3056 (HN); Dak Nong Prov., Ta Dung mt, HLF 5567 (HN, LE); Mt. Bavi, Ha Son Binh 800 m,
Balansa, 2000; Song Cao near Nhatrang 800 m, Evrard, 508 (AMES) & Poilane, 3408 (P);
Dakto, Gialai-Kontum, Dao, 230 (HN).
Notes. Closely allied to N. zollingeri var. singapureana (Backer) de Vogel,
from which differs in large pure white flowers. Specimens from Cuc Phuong national
park occasionally have flowers with 4 fertile stamens.
Apostasia Blume, 1825, Bijdr. 6, fig. 39; id. 1825, ibid. 8: 423; De Vogel, 1969,
Blumea, 17, 2: 313350; Chen et Lang, 1986, Acta Phytotax. Sin. 24, 5: 348351.
Type: A. odorata Blume
Sympodial terrestrial herbs with erect semi-woody commonly unbranched stem,
with few aerial roots at the base. Leaves few to numerous, spirally arranged, plicate,
with acute to filiform tip. Inflorescence sub-terminal, erect to arching and pendent,
usually branched, many-flowered. Flowers, sessile, not resupinate, opening widely,
hardly zygomorphic. Tepals free, sub-similar, broadly lanceolate, more or less recurved,
with dorsal keel extending apically into acute beak, the lip a little larger and wider.
Column straight or curved, usually with 2 fertile stamens, often with 1 staminode,
filaments short, basally jointed to the style; anthers free; stigma 3-lobed or suborbicular.
Fruit dry cylindrical capsule. Seeds with black hard alveolate testa.
7(4) species. Tropical and subtropical Asia to N Australia and New Guinea.
Key to species
1. Plants 512 cm tall; leaves petiolate, leaf blade ovate to elliptic, 1.52.5 cm long, 0.5
1 cm wide ........................................................................................................... 3. A. nipponica
Plants 20100 cm tall; leaves sessile, linear to lanceolate, 735 cm long, 0.32.2 cm
wide ........................................................................................................................................ 2
2. Inflorescence at the base with bunch of numerous sterile bracts; flowers white or
yellow; tepals strongly reflexed right back; staminode absent ...................... 4. A. nuda
Turczaninowia 2008, 11(1) : 5168 53
Inflorescence without bunch of sterile bracts at the base; flowers yellow; tepals
reflexed back or straight; staminode present, joined most of the way to style ............ 3
3. Tepals less than 6 mm long; capsule less than 1.8 cm long; column terete, without
wings below staminode; leaves linear to narrowly lanceolate, commonly to 1.5 cm
wide ....................................................................................................................... 1. A. wallichii
Tepals more than 6 mm long; capsule longer than 1.8 cm; column more or less flattened,
with 2 wings below staminode; leaves narrowly lanceolate to narrowly elliptic,
commonly 1.52 cm wide ..................................................................................... 2. A. odorata
1. A. wallichii R. Br. 1830, in Wall. Pl. As. Rar. 1: 75; Seidenf. 1992, Orch.
Indochina: 13; Aver. 1994, Identif. Guide Viet. Orch.: 13; Aver. et Averyanova, 2003,
Updated Checklist Orch. Viet.: 11.
Described from Nepal (in valle Nagaliae minora, Noakote dicta). Lectotype
(Wallich, 4448) K (holo), E (iso).
Plant 15100 cm tall. Leaves numerous, close, linear to narrowly lanceolate,
1235 by 315 mm. Inflorescence, simple to many branched, many flowered. Flowers
pale yellow. Tepals 45 by 0.61.2 mm. Column straight to strongly curved at base,
cylindrical, without wings. Stamens 2.34 mm long. Staminode entirely adnate to column
or with free obtuse tip. Ovary 11.7 cm long, 0.81 mm in diam. Capsule 11.8 cm
long, 1.42 mm wide. Fig. 15, a, b.
Ecology. Shady broad-leaved evergreen and coniferous lowland and sub-
montane forests on shale, sandstone, granite, gneiss and basalt. 2001200 m. Fl. March,
April. Occasional (LR).
Distribution. Vietnam (Dong Nai, Gia Lai, Khanh Hoa, Kon Tum, Lam Dong,
Ninh Thuan, Phu Tho, Quang Binh, Quang Nam, Tay Ninh, Thua Thien Hue).
Tropical Asia from Sri Lanka and Nepal to Philippines, N Australia and New Guinea.
Studied specimens. Prenh, Dalat, Evrard, 1469, 2314 (P); Lo Go, Tay Ninh,
Mueller, 1470 (P); Gian Lau me, Dong Nai, Bienhoa, Pierre, 6583 (P); Mt Tamire, Pierre, 6583
(P); Nhatrang, Poilane, 3808 (P); Ba Na, Danang, Poilane, 7277 (P); Go O to Mo O, Quang
Nam 1000 m, Poilane, 31671 (P); Gialai Kontum, Kbang Kon Ha Nung, LX-VN 2739 (LE);
Khanh Hoa, Khanh Son, Hon Giao mt., VH 4322 (HN, LE); Kon Tum, Kon Plong, DKH 4690
(HN, LE); Kontum, Kon Plong, Mang La, VH 5185 (HN, LE); Kontum, Kon Plong, Mang La, VH
5327 (HN, LE); Ninh Thuan, Ninh Hai, Nui Chua mt., HLF 4321 (HN, LE), HLF 4362 (HN, LE);
Ninh Thuan, Ninh Hai, Nui Chua mt., HLF 4362 (HN, LE); Thua Thien Hue, A Luoi, HAL 7968
(HN, LE); Quang Binh, Minh Hoa, HAL 11708 (HN).
2. A. odorata Blume, 1825, Bijdr. 6, fig. 39; id. 1825, ibid. 8: 423; Seidenf.
1992, Orch. Indochina: 13; Aver. 1994, Identif. Guide Viet. Orch.: 13; Aver. et
Averyanova, 2003, Updated Checklist Orch. Viet.: 11. A. thorelii Gagnep. 1933,
Bull. Soc. Bot. France, 70: 350.
Described from Java (montis Salak). Lectotype (Blume s.n.) L.
Plant 3080 cm tall. Leaves numerous, close, narrowly lanceolate to lanceolate,
1525 by 1.52.2 cm. Inflorescence simple to few branched, many flowered. Flowers
bright yellow. Tepals 912 by 1.52 mm. Column straight to strongly curved at base,
54 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Fig. 13. Neuwiedia zollingeri var. balansae: a flower. Apostasia odorata: b flowering
plant, c flower, d column; A. nipponica: e flowering and fruiting plant, f flower. A. nuda:
g flowering plant.
Turczaninowia 2008, 11(1) : 5168 55
Fig. 14. Neuwiedia zollingeri var. javanica: a, b (HLF 4415), c, d (HLF 5479). N. zollingeri
var. balansae: e, f (HAL 1686), g (HLF 5567).
56 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Fig. 15. Apostasia wallichii: a, b (VH 5185). A. odorata: c-e (HAL 5337). A. nipponica: f (HLF
1406), g, h (HAL 7175).
Turczaninowia 2008, 11(1) : 5168 57
usually laterally flattened, with 2 short longitudinal low to tall quadrangular wings below
staminode. Stamens 2, 56 mm long. Staminode adnate to column with free obtuse
tip. Ovary 1018 mm long, 11.2 mm wide. Capsule 1.82.5 cm long, 1.52.5 mm
wide. Fig. 13, b-d; 15, c-e.
Ecology. Shady broad-leaved evergreen and coniferous lowland, submontane
and montane forests on shale, sandstone, granite, gneiss, rhyolite and quartzite. 300
1600 m. Fl. March June. Occasional (LR).
Distribution. Vietnam (Dak Lak, Dong Nai, Ha Tinh, Khanh Hoa, Lam Dong,
Lao Cai, Phu Tho, Quang Binh, Vinh Phuc). Mainland tropical Asia, Hainan, Java,
Sumatra, Kalimantan.
Studied specimens. Bao Loc 10001200, Poilane, 22045 (P); Bavi, 1200 m,
Chevalier, 37777 (P); Dak Lak, Chu Yang Sin mt., HLF 5481 (HN, LE); Ha Tinh, Huong Son,
HAL 1281 (HN, LE), HAL 5337 (HN, LE); Khanh Hoa, Khanh Son, Hon Giao mt., VH 4174 (HN,
LE), VH 4378 (HN, LE); Lao Cai, Van Ban, DKH 6626 (HN, LE), DKH 6905 (HN, LE); Lao Cai,
Van Ban, HAL 2354 (HN, LE); Vinh Phu, Tam Dao 3, LX-VN 2368 (LE); Quang Binh, Quang
Ninh distr., HAL 11519 (HN); Quang Binh, Minh Hoa, HAL 11624 (HN).
Notes. The name A. thorelii is attributed to plants with tall quadrangular
column wings and rather broad leaves. Taxonomical status of such marginal plants is
still uncertain.
4. A. nuda R. Br. 1830, in Wall. Pl. As. Rar. 1: 76, tab. 85; Seidenf. 1992,
Orch. Indochina: 15; Aver. 1994, Identif. Guide Viet. Orch.: 13; Aver. et Averyanova,
2003, Updated Checklist Orch. Viet.: 11.
Described from Malacca Peninsular (in montibus insulae Penang).
Lectotype (Porter s.n. Wallich 4449) K.
Plant 1240 cm tall. Leaves numerous, close, linear to linear lanceolate, 720
58 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
by 38 mm. Inflorescence simple to few branched, many flowered with tuft of small
narrow sterile bracts at base. Flowers white to yellow. Tepals 3.54.5 by 0.60.8 mm.
Column strongly curved. Stamens 2, 3.23.5 mm long. Staminode absent. Ovary 8
10 mm long, 0.81 mm wide. Capsule 1.21.4 cm long, 1 mm wide. Fig. 13, g.
Ecology. Shady coniferous and broad-leaved evergreen lowland and mountain
forests on shale, 3201400 m. Fl. January, February. Very rare (DD).
Distribution. Vietnam (Dak Lak, Thua Thien Hue). Myanmar, Thailand,
Malacca Peninsular, Java, Sumatra, Kalimantan.
Studied specimens. Dak Lak, Chu Yang Sin mt., HLF 5438 (HN, LE); Thua Thien
Hue, Nam Dong, HAL 6983 (HN, LE).
Notes. Flowers of this species varies from yellow to white. Flower color of
Vietnamese samples is unknown.
Key to species
1. Lip inflated, sub-spherical to ovoid, calceolate with involuted upper apical margin;
dorsal sepal and synsepal more or less similar to petals in color and shape, white,
yellowish, dull yellow-greenish or pink, more or less of one color; petals broadly-
ovate to circular, less than twice as long as broad, very rarely petals narrowly-ovate
(in this case petals and lip hooked at apex) ....................................................................... 2
Lip goblet-like, cylindrical, with incurved side-lobes and apical margin of the pouch
Turczaninowia 2008, 11(1) : 5168 59
straight, not involuted; dorsal sepal and synsepal distinctly differ from petals in color
and shape, petals much narrower than dorsal sepal, dorsal sepal and petals commonly
variously colored, often contrastly striped and spotted; petals tapering, narrowly-
oblong or spathulate, more than twice as long as broad, newer hooked at the apex ......... 8
2. Dorsal sepal and synsepal distinctly smaller than lateral petals; lip inflated, more or
less spherical, thin-textured, lightly grooved along the veins; staminode large, broad
at apex, flat or longitudinally conduplicate, sometimes with involute side margins;
flowers white, pale yellowish-green or pink, sometimes with brown or purple stripes
along nerves; leaves tessellated or uniformly green; plant of calcareous or acidic
silicate substrates ................................................................................................................ 3
Dorsal sepal and synsepal more or less similar with petals size; lip is ovoid, rather
thick in texture, plain on surface; staminode middle sized, distinctly 3- or 5-dentate at
the apex, with a large narrow central tooth; flowers pale yellowish-white to yellow,
sepals and petals usually with small purple spots; leaves markedly tessellated; plant
of calcareous substrates ...................................................................................... 7. P. concolor
3. Leaves distinctly tessellated above and heavily spotted or uniformly tinged with
dirty-purple-violet beneath, 1015(20) cm long, 1.54 cm wide; flowers pink, pink-
purple or pale green; lip longer or hardly shorter than lateral petals; underground
stolons may be present or not; plants of calcareous or acidic silicate substrates .... 4
Leaves uniformly green on both sides, rare with few small purple-violet marks beneath
near the base, (15)2035 cm long, 46 cm wide; flowers white or pale yellowish-green;
lip distinctly smaller than lateral petals; plants of calcareous substrates, never
developing stolons .............................................................................................................. 7
4. Staminode flat, with a broad apex; sepals and petals uniformly light pink to pink-
purple; plants of calcareous or acidic silicate substrates, without stolons .................. 5
Staminode longitudinally conduplicate or with deep fold along midvein at the apex;
sepals apple-green or yellowish-pink with purple stripes along veins; plants of
calcareous substrates, with more or less elongate underground stolons ..................... 6
5. Flowers 7.510 cm across; sepals less than 4.5 cm long, subcircular to broadly-ovate,
spreading; staminode with bright yellow center and central purple mark; leaves
narrowly-oblong to oblong, 811 cm long, 33.5(4) cm wide, tessellated more or less
contrastingly deep-green/whitish-green; plant of acidic silicate substrates ..... 1. P. delenatii
Flowers 1012 cm across; sepals longer than 4.5 cm, broadly-oblong to broadly-
ovate, down directed; staminode with dull yellow or light olive-green center; leaves
ovate to oblong-elliptic, (8)1120 cm long, (2.5)47 cm wide, tessellated glossy deep-
green/silver-gray; plant of calcareous substrates ..................................... 2. P. vietnamense
6. Flowers (8)912.2 cm across, pale yellowish-green to apple-green, dorsal sepal and
petals with brown-purple stripes; dorsal sepal cuspidate-acuminate at apex, more
than 4 cm long; petals ovate to narrowly-ovate, subacute at apex, longer than 4 cm;
staminode more than 12.5 mm long, broadly ovate, convex at the base, apically with a
deep fold along the midvein and involute side margins ............................. 3. P. malipoense
Flowers 68(8.7) cm across, pink to pink-purple dorsal sepal and petals with purple
striped along veins; dorsal sepal obtuse or shortly acute at apex, less than 4 cm long;
petals broadly obovate to subcircular, rounded at apex, less than 4 cm long; staminode
less than 12.5 mm long, elliptic, longitudinally conduplicate ................... 4. P. micranthum
7(3). Flowers commonly subcampanulate, sepals and petals light yellowish-green with
purple-brown flushing at the base; lip yellowish-white; staminode broadly triangular,
60 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Dorsal sepal broadly-ovate to ovate, more or less flat, with flat margins at the base,
light dull yellowish-green, heavily spotted with large maroon spots; petals broadly-
oblong to narrowly-obovate, 1.41.6 cm wide, dull yellowish, flushed with pink-purple,
spotted with numerous large maroon spots; lip pink-purple ................... 12. P. henryanum
15(11). Dorsal sepal 4.85.2 cm long, white, sometimes with light brownish or green at the
base, more or less heavily spotted with brown-purple; petals spathulate, usually less
than 5.5 cm long, 1.82.5 cm wide; peduncle commonly erect, shortly densely hairy
with short purple-violet hairs; lithophytic plant .................................... 13. P. gratrixianum
Dorsal sepal (4.5) 57 cm long, greenish with a white margin and glossy maroon
central area, rare maroon spotted; petals obovate-spathulate, usually more than 5.5 cm
long, 2.54.6 cm wide; peduncle commonly arching, densely villosely hairy with long
whitish hairs; epiphytic plant, very rare a lithophyte .................................... 14. P. villosum
16(9). Dorsal sepal pale green with darker venation, forming a hood arching over the lip;
staminode broadly cordate with 13 obtuse apical teeth or lunate with broad falcate
side-lobes, 3- or 5-denticulate at the apex; petals spathulate, broadening toward apex;
lip often with 26 obtuse teeth on front margin ................................. 15. P. appletonianum
Dorsal sepal erect, more or less flat, with reflexed margins at the base, white with
purple or green veins; staminode lunate with narrow falcate side-lobes and 13 small
median teeth; petals oblong, sigmoid or elliptic; lip front margin without obtuse teeth ... 17
17. Petals more or less sigmoid, down-curved, green with a pink-purple apex and prominent
deep maroon hairy warts along the upper margin; dorsal sepal subcircular to broadly-
elliptic, white, green/purple striped; lip pink-brown to dull purple; leaves commonly
elliptic to obovate, 1020 cm long ................................................................... 16. P. callosum
Petals elliptic, horizontally spreading, deep-purple, finely purple-violet spotted in
lower part, without hairy warts along the upper margin; dorsal sepal broadly-ovate,
white, deep-purple striped; lip brown-purple to deep purple-violet; leaves commonly
oblong to elliptic, 714 cm long ................................................................... 17. P. purpuratum
1. P. delenatii Guillaum. 1924, Bull. Soc. Bot. France, 71: 554, 558; Seidenf.
1992, Orch. Indochina: 16; Aver. 1994, Identif. Guide Vietnam. Orch.: 16; Aver. et
Averyanova, 2003, Updated Checklist Orch. Viet.: 48; Aver. et al. 2003, Slipper Orch.
Vietnam: 89. Cypripedium delenatii (Guillaum.) C.H. Curtis, 1931, Gard. Chron.
Ser. 3, 89: 208.
Described from Vietnam (Tonkin: Sans localite precise. ...; Annam: Nha trang).
Lectotype (Tonkin, Delenat, cult. Mornay, Fleuriste de la ville de Paris) P.
Lithophytic or terrestrial herb with 57 oblong-elliptic leaves up to 11 by 33.9
cm, distinctly tessellated, heavily purple-spotted below. Peduncle with 12(3) flowers,
purple-spotted, up to 22 cm long; pedicel and ovary up to 5.5 cm long, spotted with
purple, white hairy. Flower 7.58 cm across, pale pink with pink or purple-pink lip.
Median sepal ovate, 1.73.5 by 1.82.5 cm. Synsepal similar, 1.93 by 1.42.9 cm.
Petals broadly elliptic, obtuse, 34.2 by 2.43.8 cm. Lip ellipsoidal to subglobose, 2.5
3.8 by 2.53 cm. Staminode convex, ovate, 1417 by 1316 mm with red and yellow
markings. Fig. 16, a-c; 23, a.
Ecology. Shady, broad-leaved, evergreen lowland and submontane forests on
granite and gneiss 7501300 m. Fl. November, December. Very rare (CR).
62 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
3. P. malipoense S.C. Chen et Z.H. Tsi, 1984, Acta Phytotax. Sin. 22, 2: 119;
Aver. 1994, Identif. Guide Vietnam. Orch.: 16; Aver. et Averyanova, 2003, Updated
Checklist Orch. Viet.: 49; Aver. et al. 2003, Slipper Orch. Vietnam: 104.
Described from S China (Yunnan: Malipo, Hwang Jin In, 13001600 m).
Type (Nov. 11 1947, K.M. Feng 13161) PE.
Distribution. N Vietnam, S China, NE Laos.
Key to varieties
1. Petals broadly obovate to obovate; lip wider than petals, 3.85.4 cm wide; staminode
Turczaninowia 2008, 11(1) : 5168 63
with deep brown-purple or maroon apex; leaves heavily spotted with purple-violet
beneath .............................................................................. 3a. P. malipoense var. malipoense
Petals obovate to narrowly obovate or lanceolate; lip of equal width or narrower than
petals, usually less than 3.8 cm wide; staminode with white, light yellowish or light
greenish apex marked with finest brown-purple marks; leaves sparsely spotted with
purple-violet beneath .......................................................................................................... 2
2. Petals obovate to narrowly obovate; lip subglobose, more than 1.8 cm wide, truncate
at the apex .................................................................................. 3b. P. malipoense var. jackii
Petals narrowly obovate to lanceolate; lip narrowly ovate, slightly bilaterally
compressed, hooked at apex ................................................... 3c. P. malipoense var. hiepii
3b. P. malipoense var. jackii (H.S. Hua) Aver. 1997, Orchids, 66, 2: 153.
P. jackii H.S. Hua, 1996, Die Orchidee, 46, 3: U4.
Described from S China (Yunnan, Wenshan) questionable. Type (12
64 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Fig. 16. Paphiopedilum delenatii: a flowering plant and flower, b, c staminode; P. viet-
namense: d flowering plant, e staminode. P. malipoense var. malipoense: f flowering
plant, g, h staminode (HAL 6207). P. malipoense var. hiepii: i flower, j lip, k, l staminode.
Turczaninowia 2008, 11(1) : 5168 65
3c. P. malipoense var. hiepii (Aver.) P.J. Cribb, 1998, Genus Paphiopedilum:
88. P. hiepii Aver. 1998, Orchids, 67, 3: 261. P. jackii H.S. Hua var. hiepii (Aver.)
Koop. 2000, Orchid Digest, 64, 4: 168. P. angustatum Z.J. Liu et S.C. Chen, 2000,
Acta Phytotax. Sin. 38, 5: 464.
Described from N. Vietnam (Prov. Tuyen Quang, Distr. Tuyen Quang).
Type (1415 June 1995, N.T. Hiep s.n.) LE.
Terrestrial herb. Leaves nearly white with dark green venation above and
sparse purple-violet spotting beneath. Flowers differ from the typical variety in narrow
ovate or lanceolate petals hooked at the apex, a small narrow laterally compressed
yellowish-green lip and a white staminode with few fine purple marks at apex. Fig. 16,
i-l; 23, e.
Ecology. Shady broad-leaved evergreen dry forests on soft slate limestone
and schist. 500600 m. Fl. March, April. Extinct in nature (EW).
Distribution. Vietnam (Son La?, Tuyen Quang). S China?
Notes. Species is known only by type collection. Report from S China may be
based on imported plants originated from locus classicus area. Doubtful plants with
similar coloration of leaves were occasionally observed in Son La province (HAL 9555).
purple. Median sepal ovate, 1.53.6 by 1.73 cm. Synsepal elliptic, 1.43.3 by 1.1
2.5 cm. Petals broadly obovate, 1.94.3 by 2.34.4 cm, ciliate with long white hairs.
Lip white to pink-purple, deeply inflated, 510.4 by 3.45.6 cm. Staminode white,
yellow above, spotted with red, conduplicate, 1012.5 mm long. Pedicel and ovary
3.54.6 cm long, brown-purple spotted, white hairy. Fig. 17, a-d; 23, f.
Ecology. Evergreen and semi-deciduous, mixed and coniferous forest on rocky
limestone. 6001600 m. Fl. March, April. Rare (EN).
Distribution. Vietnam (Bac Kan, Cao Bang, Ha Giang, Tuyen Quang), S China.
Studied specimens. Bac Kan, Cho Don, HAL 4771 (HN, LE), HAL 4794 (HN, LE),
HAL 4864 (HN, LE), HLF 796 (HN, LE); Cao Bang, Bao Lac, CBL 1477 (HN, LE), CBL 332
(HN); Cao Bang, Ha Lang, CBL 627 (HN, LE), CBL 767 (HN, LE); Cao Bang, Nguyen Binh,
CBL 555 (HN, LE), CBL 1277 (HN, LE), CBL 1577 (HN, LE); Cao Bang, Tra Linh, CBL 1079
(HN, LE), CBL 1222/2 (LE); Cao Bang, Trung Khanh, HAL 5457 (HN), HAL 5643 (HN, LE),
HAL 5685 (HN), HAL 5745 (HN); Ha Giang, Dong Van, CBL 1776 (HN, LE), NTH 3346 (HN,
LE), HAL 8594 (HN), HAL 8615 (HN), HAL 8653 (HN), HAL 8689 (HN); Ha Giang, Meo Vac,
CBL 1822 (HN, LE), NTH 3432 (HN, LE), HAL 8504 (HN), HAL 8564 (HN); Ha Giang, Quan
Ba, NTH 3576 (HN, K, LE), DKH 6173 (HN, LE), HAL 8341 (HN); Ha Giang, Vi Xuyen, DKH
6451 (HN, LE); Ha Giang, Yen Minh, CBL 1940 (HN, LE), NTH 3494 (HN, LE); Tuyen Quang,
Na Hang, HAL 147 (HN), HLF 659 (HN, LE).
Notes. Very sensitive species, which vanish very fast under deforestation. It
is quite variable in coloration of leaves and flowers. Its populations occasionally include
albino forms and forms with more deep flower coloration.
that description of P. singchii reported from southern Yunnan is based on plants imported
from Vietnam.
6. P. emersonii Koop. et P.J. Cribb, 1986, Orchid Advocate, 12: 86; Aver. et
Averyanova, 2003, Updated Checklist Orch. Viet.: 48; Aver. et al. 2003, Slipper Orch.
Vietnam: 140. P. huonglanae N.T. Tich, 1998, Hoa Canh (1998), 3: 1011, nom.
inval. P. emersonii Koop. et P.J. Cribb var. huonglanae (N.T. Tich) N.T. Tich,
1998, Hoa Canh (1998), 4: 12, comb. inval.
Described from China (China). Type (cult. E.W. Charles) K.
Lithophytic herb with 47 coriaceous, ligulate, glossy green leaves, below with
sparse purple-violet marks near the base. Peduncle erect, 1-flowered, 1015 cm long,
yellowish-green, shortly white hairy. Pedicel and ovary about 3 cm long. Flower slightly
sweet fragrant, 810.5 cm wide, rather thick-textured. Median sepal ovate, 2.75.1
by 1.53.5 cm. Synsepal broadly elliptic, 24.5 by 1.84 cm. Petals broadly obovate,
3.55.4 by 24 cm, white villose and sometimes faintly flushed with pink at base. Lip
light yellow to dull orange-pink, obovoid, 35 by 23 cm. Staminode bright yellow with
reddish markings, 1.62 by 0.81.1 cm, deeply longitudinally sulcate. Fig. 17, g-i; 23, h.
Ecology. Shady broad-leaved evergreen dry forests on rocky solid slaty lime-
stone. 550750 m. Fl. May, June. Very rare (CR).
Distribution. Vietnam (Bac Kan, Thai Nguyen, Tuyen Quang). S China.
Studied specimens. Bac Kan, Na Ri, HAL 4963 (HN), DKH 7644 (HN, LE, MO);
Thai Nguyen, Dong Hy, HAL 031/1 (LE); Thai Nguyen, Vo Nhai, HAL 1581 (HN).
Notes. Variable species, particularly in size of flowers, petal shape and color
of the lip.
Distribution. Vietnam (Bac Kan, Bac Giang, Ha Nam, Ha Noi, Hai Phong,
Hoa Binh, Lang Son, Nam Dinh, Ninh Binh, Quang Binh, Quang Ninh, Quang Tri, Son
La, Thai Nguyen, Thanh Hoa, Vinh Phuc), SE Myanmar, S China, Thailand, Laos,
Cambodia?
Studied specimens. Bac Kan, Ba Be, HAL 118 (HN, LE), HLF 615 (HN,
LE); Bac Kan, Cho Don, HLF 841 (HN, LE); Hai Phong, Cat Ba, LX-VN 3393/7
(HN, LE); LX-VN 3747 (HN, LE); Hoa Binh, Mai Chau, DKH 8178 (HN, LE, MO);
Lang Son, Huu Lung, NTH 3259 (HN, K, LE); Ninh Binh, Cuc Phuong, NTH 5383
(HN), NMC 1600 (HN); Quang Binh, Minh Hoa, HAL 5840 (HN), HAL 6037 (HN);
Quang Ninh, Halong Bay, NTH 2631 (HN, LE); Quang Tri, Huong Hoa, HLF 5935
(HN); Son La, Moc Chau, NTH 2922 (HN, LE); Thanh Hoa, Quan Hoa, HAL 3705
(HN), HAL 3757 (HN).
Notes. Common lowland limestone species extinct in many places of its former
occurrence. Color of flowers varies from yellowish-white to bright yellow. Occasional
flowering sometimes observed in December.
Key to varieties
1. Flowers (10)1216 cm wide; petals (5)78 cm long, in basal half strongly undulate on
margins ............................................................................ 8a. P. hirsutissimum var. esquirolei
Flowers 68(10) cm wide; petals (2.6)34(5) cm long, straight or hardly undulate on
margins ........................................................................ 8b. P. hirsutissimum var. chiwuanum
shaped, 35 by 1.52.8 cm. Staminode pale purple with glossy olive-brown center,
sub-quadrate, 1 by 0.8 cm. Fig. 18, e-h; 24, a.
Ecology. Broad-leaved evergreen, mixed and coniferous forests on rocky
crystalline limestone. 3501250 m. Fl. March May. Not rare (VU).
Distribution. Vietnam (Bac Kan, Cao Bang, Ha Giang, Hoa Binh, Lao Cai,
Ninh Binh, Son La, Thai Nguyen, Thanh Hoa, Tuyen Quang). SW China, N Thailand,
N Laos.
Studied specimens. Bac Kan, Cho Don, HAL 4669 (HN), HLF 770 (HN, LE), HLF
835 (HN, LE); Bac Kan, Na Ri, HAL 4893 (HN), Cao Bang, Ha Lang, CBL 694 (HN, LE), CBL
1677 (HN, LE); Cao Bang, Nguyen Binh, CBL 1628a (LE photo); Cao Bang, Tra Linh,
Averyanov et al. s.n., 1996 (HN, LE), Averyanov et al. s.n., 1997 (HN, LE), CBL 982 (HN, LE), DH
4349 (HN, LE); Cao Bang, Trung Khanh, HAL 5545 (HN); Ha Giang, Cuan Ba, DKH 6213 (HN,
LE); Ha Giang, Dong Van, NTH 3532 (HN, LE); Hoa Binh, Da Bac, HAL 290 (HN, LE), HAL 318
(HN, LE); Hoa Binh, Mai Chau, DKH 7748 (HN, LE, MO), DKH 8187 (HN, LE, MO); Hoa Binh,
Tung Khe, Averyanov et al. s.n., 1995 (HN, LE); Lao Cai, Sapa, Tachtajan 56 (LE); Ninh Binh,
Cuc Phuong, HAL 1637 (HN), HAL 2854 (HN), DDS 11325 (HN), NMC 933 (HN), NMC 1593
(HN); Son La, Moc Chau, NTH 2971 (HN, LE) HAL 9362 (HN, LE), HAL 9506 (HN, LE), HAL
9410 (HN, LE); Son La, Yen Chau, HAL 9433 (HN, LE); Thai Nguyen, Dong Hy, HAL 28 (HN,
LE); Thanh Hoa, Ba Thuoc, HAL 927 (HN, LE), Thanh Hoa, Ba Thuoc, HAL 969 (HN, LE),
Thanh Hoa, Ba Thuoc, HAL 1077 (HN, LE), Thanh Hoa, Ba Thuoc, HAL 2986 (HN), Thanh
Hoa, Ba Thuoc, HAL 3288 (HN), Thanh Hoa, Ba Thuoc, HAL 4345 (HN); Thanh Hoa, Quan
Hoa, HAL 3716 (HN), HAL 3933 (HN); Tuyen Quang, Na Hang, HLF 663 (HN, LE).
Notes. In the past one of the most common lithophytic species on rocky
limestone formations all over northern Vietnam. Very rarely are observed albino forms
and forms with inflorescence bearing 2 flowers.
8b. P. hirsutissimum var. chiwuanum (T. Tang et F.T. Wang) P.J. Cribb,
1987, Gen. Paphiopedilum: 140; Aver. et al. 2003, Slipper Orch. Vietnam: 166. P. chi-
wuanum T. Tang et F.T. Wang, 1951, Acta Phytotax. Sin. 1: 56.
Described from S China (China, south-east Yunnan, Foo-ning Hsien, Ban-
loun, 700 m.). Type (15 April 1940, Wang 88252) PE.
Differs in small flowers, 68(10) cm wide. Dorsal sepal 1.82.6 by 1.82.5
cm. Synsepal 1.82.4 by 1.11.5 cm. Petals, straight or hardly undulate on margins,
2.64(5) by 11.3 cm. Lip 1.82.6 by 1.21.5 cm. Fig. 18, i-k; 24, b.
Ecology. Broad-leaved evergreen, mixed and coniferous forests on rocky
crystalline limestone. 7501450 m. Fl. March May. Not rare (VU).
Distribution. Vietnam (Cao Bang, Hoa Binh, Son La), S China.
Studied specimens. Cao Bang, Bao Lac, CBL 1370 (HN, LE), CBL 1588 (HN, LE);
Hoa Binh, Da Bac, HAL 319 (HN, LE); Hoa Binh, Mai Chau, HAL 624 (HN, LE); Hoa Binh, Tung
Khe, Averyanov et al. s.n., 1995 (LE); Son La, Yen Chau, DKH 7260 (HN, LE).
Notes. Montane race closely allied to previous variety.
9. P. barbigerum T. Tang et F.T. Wang, 1940, Bull. Fan. Mem. Inst. Bot. Ser.
10: 23; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 48; Aver. et al.
Turczaninowia 2008, 11(1) : 5168 71
Fig. 18. Paphiopedilum concolor: a flowering plant, b-d staminode; P. hirsutissimum var.
esquirolei: e flowering plant and flower, f-h staminode. P. hirsutissimum var. chiwuanum: i
flower, j, k staminode. P. barbigerum var. coccineum: l flowering plant, m staminode.
72 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
2003, Slipper Orch. Vietnam: 170. P. insigne (Wall. ex Lindl.) Pfitz. var. barbigerum
(T. Tang et F.T. Wang) Braem, 1988, Paphiopedilum: 113. P. areeanum Gruss,
2001, Die Orchidee, 52, 5: 117. P. rhizomatozum S.C. Chen et Z.J. Liu, 2002, Journ.
Wuhan Bot. Res. 20, 1: 12.
Described from S China (China, Kweichow (Guizhou)). Type (Cavalerie
& Fortunat 1794) PE (holotype), P (isotype).
Distribution. N Myanmar, N Thailand, S China, N Vietnam.
Key to varieties
1. Median sepal white with uniformly pale pink, pink, pink-brownish or carmine center ...
................................................................................................. 9a. P. barbigerum var. cocineum
Median sepal white or yellowish-pink with pink to pink-carmine center spotted with
numerous contrast dark brown to brown-purple blotches ... 9b. P. barbigerum var. aspersum
10. P. tranlienianum Gruss et Perner, 1998, Caesiana, 11: 66; Aver. et Averya-
nova, 2003, Updated Checklist Orch. Viet.: 49; Aver. et al. 2003, Slipper Orch. Vietnam:
175. P. caobangense Nguyen Thien Tich, 1999, Hoa Canh (1999), 1: 14, nom. illeg.
Described from N Vietnam (Bac Thai Province, October 1998 highly
doubtful). Type (fl. in cult. November 1998, leg. O. Gruss) RO.
Lithophytic herb with 36 oblong lanceolate, glossy green leaves, to 18 by
1.7 cm, with paler margin. Peduncle suberect, 1-flowered, 1018 cm tall, bright green,
yellow to purple-brown hairy. Flower 5.56 cm across. Pedicel and ovary 2.95 cm
long, green, brown to purple-brown hairy. Dorsal sepal white, with purple veins, sub-
orbiculate, 2.53.2 by 2.83.4 cm. Synsepal pale green, ovate, 2.42.6 by 1.11.8 cm.
Petals olive-green, veined with chocolate-brown, oblong, 33.4 by 0.70.9 cm, glossy,
white-ciliate, strongly undulate along margins. Lip olive-green, glossy, with a prominent
claw-like base, 3.73.9 by 1.6 cm. Staminode yellow, obcordate to obovate, 810 by
79 mm, with glossy umbo. Fig. 19, a-e; 24, e.
Ecology. Broad-leaved, evergreen forests on rocky crystalline and slaty
stratified. 400750 m. Fl. September November. Very rare (CR).
Distribution. N Vietnam (Thai Nguyen, Tuyen Quang). Endemic.
Studied samples. Thai Nguyen, Dong Hy, HAL 30 (HN); Tuyen Quang, Na Hang,
HAL 183 (HN).
Notes. Local endemic with very restricted distribution. The plants exhibit
essential variation in size of flower. Very rare they develop inflorescence with 2 flowers.
11. P. helenae Aver. 1996, Bot. Journ. (Leningrad) 81, 9: 108; Aver. et Averya-
nova, 2003, Updated Checklist Orch. Viet.: 49; Aver. et al. 2003, Slipper Orch. Vietnam:
182. P. helenae f. aureum Gruss et Rth, 1999, Die Orchidee, 50, 1: 3. P. delicatum
Z.J. Liu et J.Y. Zhang, 2001, Acta Phytotax. Sin. 39, 1: 78.
Described from N. Vietnam (Cao Bang Prov., Tra Linh Distr., Quoc Toan
Subdistr., 850900 m, near Thang Hen Lake, environs of Thang Hen and Lung Tao
74 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
villages). Type (21 October 1995, L. Averyanov, N.T. Hiep, D.D. Huyen CB 12)
LE (holotype), HN (isotype).
Dwarf lithophytic herb with 35 coriaceous to rigid, rather succulent, oblong
leaves 312 by 0.62 cm, green with yellowish margin. Peduncle erect to almost
horizontal, one-flowered, 47 cm long, shortly black-purple pubescent. Flower 2.56
cm across. Pedicel and ovary 23.5 cm long, shortly black-purple pubescent. Median
sepal dull to bright yellow with white margin, obovate to circular, 1.83.5 by 1.53 cm.
Synsepal white, ovate, 1.52.5 by 0.81.5 cm. Petals pale yellow to bright orange
with darker nerves, straight, ligulate, 1.53.5 by 0.40.8 cm. Lip pale pink to brownish-
orange, saccate, 23 by 1.52 cm. Staminode yellow, obovate to circular, 78 mm
across, with central bright yellow to greenish umbo. Fig. 19, f-h; 24, f.
Ecology. Broad-leaved evergreen, mixed and coniferous dry forests on rocky
crystalline limestone. 500900 m. Fl. September November. Rare (EN).
Distribution. Vietnam (Bac Kan, Cao Bang, Lang Son), S China?
Studied samples. Bac Kan, Na Ri, HAL 4981 (HN); NTH 3716 (HN, LE), DKH
7539 (HN, LE, MO); Cao Bang, Ha Lang, CBL 648 (HN, LE), CBL 744 (HN, LE), CBL 767/1
(LE), CBL 789 (HN, LE), CBL 1690b (LE); Cao Bang, Thach An, CBL 830 (HN, LE), CBL 911
(HN, LE); Cao Bang, Tra Linh, CB 12 (HN, LE), CBL 1140 (HN, LE), CBL 1222/1 (LE); Cao
Bang, Trung Khanh, HAL 5456 (HN), HAL 5628 (HN); Lang Son, Bac Son, HAL 6728 (HN).
Notes. Local endemic with very restricted distribution. Quite possible this species
occurred in the past in border territory of S China allied to Cao Bang and Lang Son
provinces of Vietnam. It is very variable in plant dimensions and flower coloration. Flowers
vary from very small, about 2.5 cm wide, to relatively large up to 67 cm across.
Fig. 19. Paphiopedilum tranlienianum: a flowering plant, b flower, c-e staminode; P. helenae:
f flowering plant, g flower, h staminode; P. henryanum: i flowering plant, j flower, k, l
staminode.
76 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Studied samples. Bac Kan, Cho Don, HAL 4767 (HN), HAL 4817 (HN), HLF 772
(HN, LE), HLF 805 (HN, LE); Ha Giang, Dong Van, NTH 3533 (HN, LE); Ha Giang, Meo Vac,
CBL 1875 (HN, LE), NTH 3323 (HN, LE), HAL 8497 (HN); Ha Giang, Quan Ba, NTH 3590 (HN,
LE), DKH 6159 (HN, LE); Ha Giang, Vi Xuyen, DKH 6460 (HN, LE).
Notes. Local endemic with very restricted distribution. Very variable species,
particularly in size of flowers, form and spotting of tepals. Rarely are observed albino
specimens.
13. P. gratrixianum Masters ex Rolfe, 1905, Orchid Rev. 13: 63; Seidenf.
1992, Orch. Indochina: 17; Aver. 1994, Identif. Guide Vietnam. Orch.: 20; Aver. et
Averyanova, 2003, Updated Checklist Orch. Viet.: 48; Aver. et al. 2003, Slipper Orch.
Vietnam: 199. Cypripedium gratrixianum Masters, 1905, Gard. Chron. Ser. 3, 37:
76, fig. 35, nom. inval., non B.C. Williams, 1897. P. villosum (Lindl.) Stein var.
gratrixianum (Masters ex Rolfe) Braem, 1988, Paphiopedilum: 119.
Described from Laos (Native of Annam). Type (Micholitz s.n., cult. San-
der) K.
Distribution. S Laos, Vietnam.
Key to varieties
1. Median sepal with green, olive-green to yellowish-green center, sometimes slightly
flushed with purple-violet, spotted with numerous small spots ...........................................
.................................................................................... 13a. P. gratrixianum var. gratrixianum
Median sepal pure white, occasionally with light green or purplw violet tint at the
base, spotted with few relatively large blotches .......... 13b. P. gratrixianum var. daoense
13b. P. gratrixianum var. daoense Aver. 2002, Orch. Rev. 110, 1247: 287.
Described from N Vietnam (Vinh Phuc Prov., Tam Dao Distr., vicinities of
Tam Dao town (2127N 10539E). Tam Dao ridge at an altitude of 9501000 m. ).
Type (10 November 1998. N.T. Hiep, Averyanov L. s.n.) HN (holotype.), LE
(isotype).
Differs from type variety in pure white median sepal, rarely light green or
purple violet tinted at the base, spotted with few relatively large purple-brown blotches.
Leaves and inflorescence shorter and all plant more compact and miniature. Fig. 20, a-
c; 24, i.
Ecology. Broad-leaved, evergreen humid forests on granite, rhyolite and similar
silicate rocks. 9001500 m. Fl. October December. Very rare (EN).
Distribution. Vietnam (Lai Chau, Lao Cai, Thai Nguyen, Vinh Phuc). Endemic.
Studied samples. Lao Cai, Sa Pa, Averyanov s.n. 2005 (LE photo); Vinh Phuc,
Tam Dao, 10 November 1998. N.T. Hiep, Averyanov L. s.n. (HN, LE), VH 5041 (HN, LE).
Notes. According to observations not confirmed by herbarium collections,
this variety may be rarely found also on rocky limestone.
14. P. villosum (Lindl.) Stein, 1892, Orchideenbuch: 490; Seidenf. 1992, Orch.
Indochina: 19; Aver. 1994, Identif. Guide Vietnam. Orch.: 19; Aver. et Averyanova,
2003, Updated Checklist Orch. Viet.: 49; Aver. et al. 2003, Slipper Orch. Vietnam:
208. Cypripedium villosum Lindl. 1854, Gard. Chron. (1854): 135.
Described from Myanmar (Burma). Type (Lobb s.n.) K.
Distribution. NE India, Myanmar, S China, Laos, Vietnam, Thailand.
Key to varieties
1. Median sepal broadly oblanceolate to narrowly obovate, boldly spotted with deep
brown fused blotches, with strongly reflexed lateral margins throughout its length;
petals often marked with purple-brown, narrowly spatulate, distinctly attenuate to
narrowing stipitate base ............................................................ 14e. P. villosum var. boxallii
Median sepal obovate to suborbicular, without distinct spots, with strongly reflexed
lateral margins at lower part; petals without purple-brown marks, with more or less
broad, not distinctly stipitate base ................................................................................... 2
2. Median sepal dark brown with more or less wide yellowish-green to whitish margin ....
.................................................................................................... 14a. P. villosum var. villosum
Median sepal light yellowish-green, dull pink to purple, or white with more or less
wide median longitudinal brown stripe ............................................................................ 3
3. Median sepal white, with more or less wide median longitudinal brown stripe,
sometimes with broad dark brown central area occasionally margining with green .......
................................................................................................ 14b. P. villosum var. annamense
Central area of median sepal dull pink to dark purple or dull yellowish-green tinted
with light brown along midvein ........................................................................................ 4
78 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
4. Central area of median sepal dull pink to dark purple, occasionally greenish at the
base; sepal margin commonly white ............................ 14d. P. villosum var. fusco-roseum
Median sepal dull yellowish-green, tinted with light brown along midvein, occasionally
with white or yellowish-white margin .......................... 14c. P. villosum var. fusco-viride
14b. P. villosum var. annamense Rolfe, 1907, Curtis Bot. Mag. 133: t. 8126;
Aver. et al. 2003, Slipper Orch. Vietnam: 215.
Described fromSVietnam(Annam, Micholitz s.n., cult. Sander). Lectotype
Ic. P. villosum var. annamense (1907, Curtiss Bot. Mag. 133, tab. 8126).
This variety differs from the type in pure white median sepal with narrow
chestnut-brown longitudinal stripe. Occasionally this central brown stripe on sepal
becomes wider forming broad chestnut to brown-purple central area margining with
green. Fig. 20, d-f; 25, b.
Ecology. Shady broad-leaved mixed and evergreen humid forests mainly on
silicate rocks. 13002000 m. Fl. November March. Occasional (VU).
Distribution. Vietnam (Dak Nong, Khanh Hoa, Lam Dong). Endemic.
Studied samples. Dak Nong, Dak Glong, HLF 5671 (HN); Gia Lai, Chu Pah, Phan
Ke Loc, P-9863 (HN); Khanh Hoa, Khanh Son, VH 4349 (HN, LE); Lam Dong, Dalat, Averyanov
s.n., 1995 (LE photo), Averyanov s.n., 2005 (LE photo), O. Zaitsev et al. 153 & 281 (LE); Lam
Dong, Lac Duong, VH 3120 (HN, LE), VH 3333 (HN, LE).
Notes. Eastern vicarious race of the species, which is rather common in
montane forests of southern Vietnam. Very variable. Grows commonly as humus
epiphyte.
Turczaninowia 2008, 11(1) : 5168 79
14e. P. villosum var. boxallii (Reichenb. f.) Pfitz. 1903, in Engler, Pflanzenr.
4, 50, 12, Orch. Pleon.: 73; Aver. et al. 2003, Slipper Orch. Vietnam: 217.
Cypripedium boxallii Reichenb. f. 1877, Gard. Chron. n. s. 7: 367.
Described from Myanmar (Burma, Moulmein). Type (Boxall s.n.) W.
This variety is distinguished by its deep colored flowers with a boldly spotted narrow
median sepal and narrow lateral petals heavily striped with purple-brown. Fig. 25, e.
Ecology. Shady broad-leaved evergreen humid mountain forests? Fl.? Probably
very rare (DD).
Distribution. Vietnam (Lao Cai, Lai Chau)?, Myanmar.
Studied samples. Lao Cai, Sa Pa, Averyanov s.n., 2004 (LE photo).
Notes. Many plants of this rare variety collected in nature are cultivated in Sa
Pa town. Meanwhile the occurrence of this plant in Vietnam needs confirmation by
herbarium collections.
80 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
4, 24: 551. P. amabile auct. non Hallier.: Seidenf. 1975, Contrib. Revis. Orch. Fl.
Cambod. Laos Vietnam: 88; Aver. 1988, Prelim. List Vietnam. Orch. 2: 31.
Described from Thailand (Siam). Type (Regnier s.n.) W.
Distribution. Thailand, Vietnam, Laos, Cambodia.
Key to varieties
1. Petals strongly sigmoid, distinctly down deflexed, usually more than 5 cm long,
commonly with ciliate warts along upper and lower margin; median sepal usually
morethan5cmwide................................................................. 16a.P. callosum var. callosum
Petals hardly sigmoid or straight, borne at an angle of 45 to the horizontal, usually
less than 5 cm long, with few ciliate warts along upper margin or without marginal
warts at all; median sepal usually less than 5 cm wide .................................................... 2
2. Petals bear few warts only on the upper margin ......... 16b. P. callosum var. warnerianum
Petals lack the marginal warts at all ........................ 16c. P. callosum var. potentianum
16b. P. callosum var. warnerianum (T. Moore) P.J. Cribb ex Aver. comb.
nov. Cypripedium barbatum var. warnerianum T. Moore, 1878, Warner Select.
Orchid. Ser. 3, Pl. 3, t. 11. C. callosum var. sublaeve Reichenb. f. 1888, Gard.
Chron. Ser. 3, 3: 331. P. callosum var. sublaeve (Reichenb. f.) P.J. Cribb, 1987,
Gen. Paphiopedilum: 188.
Origin of the type unknown. Lectotype Ic. Paphiopedilum callosum var.
Turczaninowia 2008, 11(1) : 5168 83
16c. P. callosum var. potentianum (O. Gruss et J. Roeth) P.J. Cribb, 1998,
Gen. Paphiopedilum, ed. 2: 337. P. potentianum O. Gruss et J. Roeth, 1995, Caesiana,
5: 39.
Described from Thailand (Thailand). Type (cult. E. et G. Potent, Roth
s.n.) HAL.
The variety differs from the type in unspotted petals lacking marginal warts
on the upper margin and in its narrower dorsal sepal, about 4 by 2.6 cm. Fig. 25, i.
Ecology. Shady evergreen broad-leaved lowland and submontane forests. Fl.
April June. Very rare (DD).
Distribution. Vietnam (Lam Dong), Thailand.
Studied samples. Lam Dong, Dalat, Averyanov, HAL s.n. 2004 (LE photo).
Notes. May be certainly found in lowlands of Laos and Cambodia. Rarely
forms sub-albino and albino forms.
2.53.5 by 2.24.3 cm. Synsepal green with darker veins, narrowly ovate, 23.5 by
1.21.6 cm. Petals glossy deep purple-maroon, heavily spotted with black-maroon
spots, spreading, narrowly elliptic, 3.54.6 by 0.91.3 cm. Lip purple, 3.34.4 by 2
2.7 cm, verrucose on incurved side-lobes. Staminode yellowish, purple tinted along
margin, with green veins, lunate, with narrow falcate acute side lobes, 8 by 811 mm.
Fig. 21, d-g; 26, a.
Ecology. Broad-leaved evergreen and semi-deciduous forests on highly eroded
crystalline limestone at elev. 6001400 m. Fl. September October. Very rare (EN).
Distribution. Vietnam (Cao Bang, Lao Cai, Tuyen Quang), SE China.
Studied samples. Cao Bang, Bao Lac, CBL 1458b (LE); Lao Cai, Sa Pa, P.K. Loc
s.n., 2000 (HN, LE photo); Tuyen Quang, Na Hang, HLF 662 (HN, LE).
Notes. Rare species, which populations commonly include very few scattered
samples. Leaf tessellation varies from whitish or pale green to deep green. Rarely
forms 2-flowered inflorescence.
18. P. dianthum T. Tang et F.T. Wang, 1940, Bull. Fan Mem. Inst. Biol. Ser.
10: 24; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 48; Aver. et al.
2003, Slipper Orch. Vietnam: 244. P. parishii (Reichenb. f.) Stein var. dianthum
(T. Tang et F.T. Wang) Karasawa et Saito, 1982, Bull. Hiroshima Bot. Gard. 5: 38.
Described from S China (China, Yunnan, Mengtze, ). Type (Wang et
Liu, 83446) PE.
Large lithophytic herb forming clumps of up to 15 growths, each with 46
coriaceous, ligulate, dark green leaves 2050 by 25 cm. Inflorescence arching or
suberect, with 17 flowers, green, sparsely papillose below. Pedicel and ovary glabrous,
47 cm long. Flowers 1215 cm across. Dorsal sepal white with green venation at the
base, obovate, 3.84.3 by 2.22.5 cm. Synsepal pale green to white, broadly ovate,
3.53.8 by 2.32.6 cm. Petals light greenish, olive-green striped, with few black hairy
marginal warts near base, linear-tapering, strongly twisted, pendent, 811 by 11.1 cm,
with roundish hairy apex. Lip olive-green, deeply saccate, 44.5 by 22.5 cm. Staminode
obcordate, with a blunt basal umbo, three-lobed at apex, 911 by 67.5 mm, white
with dark green venation in the center. Fig. 21, h-k; 26, b.
Ecology. Coniferous, mixed and broad-leaved, evergreen, humid forests on highly
eroded, crystalline limestone at elev. 6001450 m. Fl. September November. Rare (EN).
Distribution. Vietnam (Cao Bang, Ha Giang, Hoa Binh, Lai Chau, Lao Cai,
Son La), S China.
Studied samples. Cao Bang, Bao Lac, CBL 394 (HN, LE); Ha Giang, Dong Van,
NTH 3364 (HN, LE), NTH 3539 (HN, LE); Hoa Binh, Mai Chau, HAL 680 (HN, LE), DKH 7871
(HN, LE, MO); Lai Chau, Averyanov s.n. (LE photo); Lao Cai, Averyanov s.n. (LE photo);
Son La, Moc Chau, DKH 7396 (HN, LE), NTH 2970 (HN, LE), HAL 9328 (HN, LE), HAL 9409
(HN); Son La, Yen Chau, DKH 7279 (HN, LE), HAL 9500 (HN, LE).
Notes. Species is closely allied to P. parishii from Myanmar, Thailand and
SW Yunnan, but differs in its lithophytic habit, glabrous inflorescences, bracts and
ovary and in white dorsal sepal.
Turczaninowia 2008, 11(1) : 5168 85
Fig. 22. Paphiopedilum appletonianum P. villosum (P. cribbii): a flowering plant, b stami-
node; P. callosum P. villosum (P. dalatense): c flowering plant, d flower, e staminode;
P. helenae P. hirsutissimum (P. herrmannii): f flowering plant, g flower, h, i staminode.
88 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Fig. 23. Paphiopedilum delenatii: a (Averyanov s.n., 2005); P. vietnamense: b (HAL 31); P. ma-
lipoense var. malipoense: c (HAL 6207); P. malipoense var. jackii: d (HAL 169); P. malipoense
var. hiepii: e (Hiep s.n., 1995, type); P. micranthum: f (HAL 8341); P. hangianum: g (HAL
4789); P. emersonii: h (HAL 1581); P. concolor: i (HLF 5935).
Turczaninowia 2008, 11(1) : 5168 89
Fig. 24. Paphiopedilum hirsutissimum var. esquirolei: a (Averyanov et al. s.n., 1995); P. hirsu-
tissimum var. chiwuanum: b (CBL 1588); P. barbigerum var. coccineum: c (P-10546); P. barbi-
gerum var. aspersum: d (P-10547); P. tranlienianum: e (HAL 183); P. helenae: f (Averyanov et al.,
CB 12, 1995, type); P. henryanum: g (NTH 3590); P. gratrixianum var. gratrixianum: h (Ave-
ryanov s.n., 2005); P. gratrixianum var. daoense: i (Averyanov s.n., 2005).
90 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Fig. 25. Paphiopedilum villosum var. villosum: a (Averyanov s.n., 2005); P. villosum var. anna-
mense: b (Averyanov s.n., 1995); P. villosum var. fusco-viride: c (Averyanov s.n., 2005); P. villosum
var. fusco-roseum: d (Averyanov s.n., 2005); P. villosum var. boxallii: e (Averyanov s.n., 2004);
P. appletonianum: f (HLF 5800); P. callosum var. callosum: g (HLF 5575); P. callosum var. warne-
rianum: h (Averyanov, HAL s.n., 2004); P. callosum var. potentianum: i (Averyanov, HAL s.n., 2004).
Turczaninowia 2008, 11(1) : 5168 91
Fig. 26. Paphiopedilum purpuratum: a (CBL 1458b); P. dianthum: b (Averyanov s.n.); P. apple-
tonianum P. villosum: c (HAL 8626, type); P. barbigerum (var. aspersum) P. gratrixianum:
d (Averyanov s.n., 2005); P. callosum P. gratrixianum: e (Averyanov s.n., 2005); P. callosum
P. villosum: f (Averyanov s.n., 2005); P. gratrixianum P. villosum: g (Averyanov s.n., 2005);
P. helenae P. hirsutissimum: h (Averyanov s.n., 2002); P. malipoense P. micranthum: i
(Averyanov s.n., 2006).
92 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Corymborkis Thouars, 1809, Nouv. Bull. Sci. Soc. Philom. (Paris) 2 ser. 1:
318; Rasmussen, 1977, Bot. Tidskr. 71, 34: 161192.
Type: C. corymbis Thouars
Sympodial terrestrial herbs with short woody rhizome, fasciculate rigid roots
and erect semi-woody bamboo-like unbranched stems up to 4 m tall. Leaves sessile,
glabrous, plicate, sheathing at the base. Inflorescences lateral, panicle, with many
flowers. Flowers white, resupinate or not. Sepals and petals basally connate to form a
short tube, sub similar, long and narrow, widened toward the apex, spreading. Lip as
long as sepals, narrow and close to column, except apical ovate reflexed blade. Column
long, slender, straight, thickened to the apex. Anther erect, acute. Stigma broad, deeply
2-lobed. Pollinia 2, sectile, with linear caudicle and small peltate viscidium.
5(1) species. Tropical zone of the world.
Turczaninowia 2008, 11(1) : 5168 93
1. C. veratrifolia (Reinw.) Blume, 1859, Coll. Orch. Arch. Ind.: 125, pl. 43, 1,
pl. 42E; Seidenf. 1992, Orch. Indochina: 27; Aver. 1994, Identif. Guide. Vietnam.
Orch.: 29; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 24. Hysteria
veratrifolia Reinw. 1828, Syll. Pl. Nov. Ratisbon. 2: 5.
Described from Java? Neotype (Java: T. Lobb, 162 sec. Rasmussen,
1977, Bot. Tidsskr. 71, 34: 170) K.
Stem up to 5 m tall with spirally arranged leaves up to 45 cm long and 12 cm
wide, with long sheaths covering stem. Inflorescence up to 10 cm long. Sepals, petals
and lip 24 cm long. Reflexed apical blade of the lip ovate with finely crisped edges
and shortly pointed apex. Column to 23.5 cm long. Fruits cylindrical capsule 34 cm
long. Fig. 27, a,b; 31, a.
Ecology. All kinds of lowland and submontane forests on any soils. 501000 m.
Fl. May July. Not rare (LR).
Distribution. Vietnam (probably all provinces). Tropical and subtropical Asia,
Australia and Pacific islands to Samoa.
Studied specimens. Bac Kan, Cho Don, HAL 4773 (HN, LE); Bac Kan, Na Ri, HAL
4952 (HN, LE), DKH 7631 (HN, LE, MO); Ha Nam Ninh, Cuc Phuong, Kien 147 CP (HN, LE);
Ha Son Binh, Luong Son, Tiep et al., NOT-2765 (HN, LE); Hai Phong, Cat Ba, 1988, LX-VN
3340, LX-VN 3779 (HN, LE); Hoang Lien Son, Dan Khao, 1964, sine coll. 3178 (HN, LE); Kim
Son, Trung, 1702 (HN, LE); Ninh Binh, Cuc Phuong national park, N.T. Hiep, NTH 5364 (HN);
Quang Binh, Bo Trach, HAL 6101 (HN, LE); Quang Binh, Ke Bang, VH 4609; VH 4768 (HN,
LE); Quang Binh, Minh Hoa, HAL 5870 (HN, LE), Lai Chau, Muong Te, HAL 10087 (HN, LE),
Nghe An, Tuong Duong, HLF 6606 (HN, LE).
Note. This tallest, shrub-like orchid in Vietnam is sometimes truly named as
bamboo orchid.
Tropidia Lindl., 1833, Bot. Reg. 19, tab. 1618; Seidenf. 1978, Dansk Bot.
Ark. 32, 2: 108113. Schoenomorphus Thorel ex Gagnep. 1933, Bull. Soc. Bot. Fr.
80: 351.
Type: T. curculigoides Lindl.
Sympodial terrestrial herbs with short woody rhizome, fasciculate rigid roots
and erect semi-woody rod-like branched, or superposed leafy stems. Leaves sessile,
glabrous, plicate, sheathing at the base. Inflorescences lateral or terminal, a short,
dense raceme. Flowers white, not resupinate. Sepals and petals free, or lateral sepals
connate into synsepalum. Lip short, spurred or saccate at the base, not mobile, narrowed
to down turned, triangular, obtuse apex. Column short. Clinandrium thin-margined.
Anther erect, dorsal. Pollinia 2, sectile, with linear caudicle and viscidium.
20(2) species. Tropical zone of the World.
Key to species
1. Leaves 2, broadly elliptic-ovate, usually sub opposite at a top of stem; inflorescence
terminal, more than 2 cm long; lateral sepals connate into synsepalum ...... 1. T. angulosa
Numerous narrowly-ovate leaves distant throughout stem; inflorescence lateral, rare
terminal, less than 2 cm long; lateral sepals free or connate at the base only ... 2. T. curculigoides
94 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
1. T. angulosa (Lindl.) Blume, 1858, Coll. Orch. Arch. Ind. Jap.: 122; Seidenf.
1992, Orch. Indochina: 27; Aver. 1994, Identif. Guide Vietnam. Orch.: 28; Aver. et
Averyanova, 2003, Updated Checklist Orch. Viet.: 59. Cnemidia angulosa Lindl.
1833, Bot. Reg. 19, tab. 1618.
Described from NE India (India orientali; in montibus prov. Sylhet). Lecto-
type (Wallich, 7388) K.
Stem to 40 cm tall, simple or superposed, with 2 apical sub-opposite leaves.
Leaves ovate to broadly-ovate, or even circular, acute to shortly acuminate, 620 cm
long, 2.511 cm wide. Inflortescence dense terminal spike 14 cm long on erect stalk
28 cm tall. Floral bracts narrowly-cuneate to filiform, 514 mm long. Flowers white,
sweet fragrant. Sepals oblong, 67 mm long, 34 mm wide, shortly suddenly apiculate
at the apex; lateral sepals connate into bidentate synsepalum. Petals smaller, oblong,
obtuse. Lip ovate, obtuse, 3.54.5 mm long, concave at the base, with strongly turned
down brightly yellow apex. Anther ovate, 2 mm long. Fig. 27, c; 31, b.
Ecology. All kinds of lowland and submontane forests on any soils. 01600 m.
Fl. April July. Common (LR).
Distribution. Vietnam (probably all provinces). Tropical and subtropical main-
land Asia, S Japan, Taiwan, Hainan, Java, Sumatra.
Studied specimens. Bac Kan, Ba Be National Park, HLF 604 (HN, LE); Bac Kan,
Cho Don, HLF 879 (HN, LE); Cao Bang, Ba Be national park, Averyanov et al., CB 46 (HN, LE);
Cao Bang, Bao Lac, CBL 1547 (HN, LE); Cao Bang, Bao Lac, CBL 307 (HN, LE); Dak Lak, Chu
Yang Sin mt., HLF 5482 (HN, LE); Ha Tinh, Huong Son, HAL 1223, 5261 (HN, LE); Hai Phong,
Cat Ba, LX-VN 3266, 3335, 3396, 3447, 3660, 3679 (HN, LE, C); Hoa Binh, Lac Son, HAL
9033 (HN, LE); Hoa Binh, Pa Co, VH 2425 (HN, LE); Lai Chau, Sin Ho, HAL 9837 (HN, LE);
NgheAn, Tuong Duong, HLF 6625 (HN, LE), HLF 6708 (HN); Quang Binh, Ke Bang, VH
4606 (HN, LE); Quang Binh, Minh Hoa, HAL 5867 (HN, LE); Quang Ninh, Ha Long Bay, NTH
5621 (HN, LE); Son La, Mai Son, HAL 9586 (HN); Vungtau-Condao, Bay Canh, 1989, Averyanov
412 (HN, LE); Vungtau-Condao, Con Dao, 1987, Averyanov 881 (HN, LE); Vungtau-Condao,
Con Dao, 1989, Averyanov 512 (HN, LE).
2. T. curculigoides Lindl., 1840, Gen. Sp. Orch. Pl.: 497; Seidenf. 1992, Orch.
Indochina: 27; Aver. 1994, Identif. Guide Vietnam. Orch.: 27; Aver. et Averyanova,
2003, Updated Checklist Orch. Viet.: 60. T. pedunculata Blume, 1859, Coll. Orch.
Arch. Ind.: 122, pl. 40; Seidenf. 1992, l.c.: 27. T. graminea Blume, 1859, l.c.: 124, pl.
41, 3; Seidenf. 1992, l.c.: 27. Schoenomorphus capitatus Thorel ex Gagnep. 1933,
Bull. Soc. Bot. France, 80: 351.
Described from NE India (India orientali; montibus Sylhet, et secus ripas
fluminis Attran). Lectotype (Wallich, 7368) K.
Stem to 1.5 m tall, often branched, with numerous distant leaves. Leaves
narrowly-lanceolate to narrowly-ovate, acuminate, 525 cm long, 0.85 cm wide.
Inflorescence lateral axillary, rarely subapical, 11.8 cm long. Floral bracts triangular-
acuminate, acute, 510 mm long, densely imbricate, lowest sterile. Flowers white.
Sepals and petals free, narrowly-ovate, acute, recurved, 710 mm long, petals smaller.
Turczaninowia 2008, 11(1) : 5168 95
Lip ovate, obtuse, 57 mm long, concave at the base, with turned down apex. Anther
narrowly-ovate, acuminate, 56 mm long. Fig. 27, d; 31, c.
Ecology. All kinds of forests on any soils. 02000 m. Fl. April July. Very
common (LR).
Distribution. Vietnam (probably all provinces). Tropical and subtropical main-
land Asia, Taiwan, Hainan, Malaysia, Java, Kalimantan, Timor to New Caledonia.
Studied specimens. Bac Kan, Ba Be National Park, HLF 618 (HN, LE); Bac Kan,
Cho Don, Po Luong mt., HAL 4736 (HN, LE); Cao Bang, Ba Be national park, CB 43 (HN, LE);
Cao Bang, Tra Linh, CBL 1251 (HN, LE); Cao Bang, Yen Lac, CBL 307 (HN, LE); Ha Giang,
Quan Ba, NTH 3564 (HN, LE); Ha Son Binh, Luong Son, Tiep, NOT 2765 (HN, LE); Ha Tinh,
Huong Son, DKH 4381 (HN, LE, MO); Hai Phong, Cat Ba, LX-VN 3340a (HN, LE); Kon Tum,
Dak Gley, VH 1558 (HN, LE); Kon Tum, Dak Gley, VH 1383 (HN, LE); Lai Chau, Muong Te,
HAL 10094 (HN, LE); Lao Cai, Van Ban, DKH 6538, 6627 (HN, LE, MO); Nghe An, Tuong
Duong, HLF 6620 (HN, LE); Ninh Thuan, Ninh Hai, Nui Chua mt., HLF 4276 (HN, LE); Quang
Binh, Minh Hoa, HAL 5859 (HN, LE); Quang Ninh, Ba Mun, 1987, Averyanov 142 (HN, LE);
Quangnam-Danang, Phuoc Son, LX-VN 3088 (HN, LE); Thai Nguyen, Dong Hy, HAL 9093
(HN, LE); Thua Thien Hue, A Luoi, HAL 7888 (HN, LE); Vungtau-Condao, Con Dao, 1987,
Averyanov 770, 806, 880 (HN, LE); Vungtau-Condao, Con Dao, 1989, Averyanov 535 (HN, LE).
Anoectochilus Blume, 1825, Bijdr. 6, fig. 15; id. 1825, ibid. 8: 11 (sub Anoeco-
chilus sphalm.); id. 1828, Fl. Javae 6, emend. nom. cons.
Type: A. setaceus Blume
Terrestrial and lithophytic sympodial herbs with creeping succulent rhizome
rooting at nodes, ascending erect leafy floriferous stem and terminal lax, few flowered
spike. Leaves few, spirally arranged in rosette, petiolate, sheathing at the base, glabrous,
convolute, asymmetric, velvety dark green, brown-green to nearly black, with more or
less pronounced reticulation of white, yellow or pink nerves. Scape leafless, with 24
sterile yellowish bracts. Flowers, sessile, resupinate or not. Sepals free. Petals very
oblique, connivent to median sepal forming hood, lateral sepals spreading. Lip adnate
to column base, spurred. Spur conical to cylindric, with 2 short peltate glands on lateral
sides. Mesochile channeled, long to shortly fimbriate-toothed on flanges. Epichile with
96 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
2 divergent flabellate spreading lobes. Column short, fleshy, with 2 wings at front, 2
lateral stigmas and prominent rostellum. Anther triangular-ovate, acute. Pollinia 2,
sectile, with stipe and viscidium.
30(7) species. Tropical mainland Asia, Indonesia, Australia, New Guinea, Pacific
Islands to Polynesia and adjacent islands.
Key to species
1. Spur and lip lie in one line or placed at a broad angle; spur more or less parallel with
ovary ...................................................................................................................................... 2
Spur strongly bent down in right or acute angle with mesochile, placed in a wide
angle to ovary ..................................................................................................................... 3
2. Mesochile flanges long fimbriate; sepals flat without any callosities; epichile lobes
oblong-lanceolate, more or less flat ............................................................ 1. A. roxburghii
Mesochile flanges toothed; sepals outside with few fat papillae along midvein;
epichile lobes triangular-flabellate, truncate, plicate along margin .......... 2. A. papillosus
3. Leaves black or gray-black, rarely dark gray-brown with green tint, commonly dense
reticulated with net of silvery-white nerves; mesochile flanges pectinate with more
than 10 small dents or short fringes ................................................................ 3. A. calcareus
Leaves green, dark green to green-brown, rarely black-green, commonly reticulated
with net of white, yellowish or pink nerves; mesochile flanges with few blunt dents
on 26 long fringes ............................................................................................................. 4
4. Mesochile flanges with at least 12 long linear fringes 35 mm long ........................... 5
Mesochile flanges with at few short dents ....................................................................... 6
5. Fringes on mesochile flange 46, all more or less equal in length; sepals lanceolate or
broadly-lanceolate, slightly falcate; column wings broad, rectangular, undulate along
margin ............................................................................................................ 4. A. albolineatus
Mesochile with 13 pair of long fringes, other diminishing toward the base into small
narrow dents; sepals obovate, strongly oblique, with short suddenly attenuate apex;
column wings narrowly-triangular, straight, acute .................................. 5. A. annamensis
6. Spur narrowly-cylindrical, longer than 4 mm; epichile lobules lanceolate to broadly-
lanceolate, 45 times as long as broad; mesochile lobes insignificant, flanges narrow,
less than 1 mm wide, slightly erose or with few short irregular denticles near the base
of epichile ................................................................................................................. 6. A. lylei
Spur broadly-conical shorter than 4 mm; epichile lobules narrowly obovate, commonly
not more than twice as long as broad; mesochile lobes large, erect, triangular, flanges
broad, triangular more than 1 mm wide, with 34 broad blunt dents along margin .........
............................................................................................................................ 7. A. brevilabris
1. A. roxburghii (Wall.) Lindl. 1840, Gen. Sp. Orch. Pl.: 499; Seidenf. 1992,
Opera Bot. 114: 32; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 11.
Chrysobaphus roxburghii Wall. 1826, Tent. Fl. Napal.: 37. A. setaceus auct. non
Blume: Aver. 1994, Identif. Guide Vietnam. Orch.: 46.
Described from N India (India, Mt. Sylhet ...). Syntypes (coll. F. De Silva
Wall. Cat. 7387; E India Company, icon. Wallich 34) K.
Terrestrial herb with 34 petiolate leaves in basal rosette. Leaves ovate, acute,
Turczaninowia 2008, 11(1) : 5168 97
Fig. 27. Corymborkis veratrifolia: a flower, b sepal, petal, lip and column; Tropidia angulosa:
c flower, sideand frontal view; T. curculigoides: d flower; Anoectochilus roxburghii: e
flower and flattened column and tepals, f spur section with glands; A. papillosus: g
flowering plant and leaf, h, i flower, frontal and side view, j dorsal surface of median sepal
and petals, k flattened tepals without lip, l column, ventral and dorsal view, m pollinarium.
Turczaninowia 2008, 11(1) : 5168 99
3. A. calcareus Aver. 1996, Bot. Journ. (St. Petersburg) 81, 10: 75, fig. 2.
Described from northern Vietnam (Prov. Hoa Binh, near 1 km of Mai Chau
to the E). Type (20 May 1995, Averyanov et al. s.n.) LE.
Terrestrial and lithophytic herb with 34 rosulate petiolate leaves. Leaves ovate,
acute, to 7 cm long, dark velvety gray-brown to black, with dense network of silvery-
white to light pink nerves. Scape hairy, 812 cm long, with 37 lax flowers. Floral
bracts cuneate, tomentose, as long as ovary. Ovary tomentose, 810 mm long. Sepals
light green, sparsely hairy; dorsal sepal broadly-ovate, 45 mm long, acute; lateral
sepals ovate, oblique, 67 mm long. Petals white, greenish to the apex, strongly falcate,
attenuate to the base and apex, as long as median sepal. Lip white, 1315 mm long,
basally with small conical, flattened spur in a wide angle to a claw; spur 3 mm wide,
2.5 mm wide, with 2 capitate gland. Claw 45 mm long, with 1015 pair of dents or
short fringes on flanges. Epichile lobes obovate, 56 mm long 2.53 mm wide. Column
3 mm tall, with 2 small vertical wings broadening proximally, not entering into spur.
Anther ovate, acute. Fig. 28, a-e; 31, g.
100 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
4. A. albolineatus Par. et Reichenb. f. 1874, Trans. Linn. Soc. Bot. 30: 141.
A. siamensis Schleichter, 1906, Feddes Repert. 2: 83; Seidenf. 1992, Opera Bot. 114:
32; Aver. 1994, Identif. Guide Vietnam. Orch.: 46; Aver. et Averyanova, 2003, Updated
Checklist Orch. Viet.: 11.
Described from Myanmar (Burma Moulmein). Type (Parish 325) K
(holotype), W (isotype).
Terrestrial herb with 23 shortly petiolate leaves in basal rosette. Leaves
broadlyovate, acute, 1.54 cm long, velvety dark-green to brown-green or nearly
black, often with broad irregular median longitudinal white to yellowish stripe or
reticulation of pink to yellowish veins. Scape pubescent, with 23 acuminate yellowish
bracts, 615 cm tall, with 18 lax flowers. Floral bracts ovate, acuminate, pubescent.
Ovary pubescent, 510 mm long. Sepals sub-similar, ovate-oblong, 710 mm long,
acute, sparsely hairy. Petals narrowly-obovate, slightly oblique, as long as median
sepal, shortly acuminate. Lip 57 mm long, spurred. Spur directed parallel with ovary
and claw, 1.53 mm long, broadly-conical, notched at the apex, with 2 large peltate,
warty glands. Mesochile with 47 pairs of long fringes along flanges. Epichile lobes
narrowly-obovate to oblique-flabellate, more or less truncate, 57 mm long. Column at
front with 2 large narrowly-triangular, parallel wings not entering to the spur and 2 short
broad rostellar arms. Anther narrowly-ovate, acute, 34 mm long. Fig. 28, f-h; 31, h.
Ecology. Humid broad-leaved forests on silicate soils. 11001700 m. Fl. No-
vember January. Rare (VU).
Distribution. Vietnam (Kon Tum, Lam Dong, Lao Cai), Myanmar, Thailand.
Studied specimens. Kon Tum, Dak Gley, VH 1675 (HN, LE); Lam Dong, Lac
Duong, Hon Giao ridge, VH 4132 (HN, LE); Muang Xen, Hayata s.n. (TI).
Fig. 28. Anoectochilus calcareus: a flowering plant, b flower, c flattened tepals, d flat-
tened lip and section of spur, e column, dorsal and ventral view; A. albolineatus: f, g flat-
tened tepals and lip, h column ventral and side view; A. annamensis: i flowering plant, j flat-
tened tepals, k flattened lip and section of spur, l column and section of spur, side view;
A. lylei: m, n flattened tepals and lip, o column and spur section, frontal view, p column, side view.
102 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
6. A. lylei Rolfe ex Downie, 1925, Kew Bull. 1925: 411; Seidenf. 1992, Opera
Bot. 114: 33; Aver. 1994, Identif. Guide Vietnam. Orch.: 44; Aver. et Averyanova,
2003, Updated Checklist Orch. Viet.: 11.
Described from Thailand (Sim ... Me Awn, 750 m). Type (Lyle in herb.
Kerr, 297) K.
Terrestrial herb with 24 petiolate leaves in basal rosette. Leaves ovate to
almost circular, 26 cm long, shortly acute, dark green, green-brown to nearly black
with reticulated yellowish to pink nerves, rarely with only 1 or 3 longitudinal colored
veins. Scape densely pubescent, with 24 yellowish, broad, acuminate bracts, 615 cm
tall, with 18 lax flowers. Floral bracts ovate, acuminate, pubescent, 714 mm long.
Ovary 1016 mm long, densely pubescent. Sepals sub-similar, ovate-oblong, 711 mm
long, shortly acute, hairy. Petals obovate, as long as median sepal, strongly oblique,
Turczaninowia 2008, 11(1) : 5168 103
rising from narrow base to broad median part and oblique attenuate apex. Lip 15
18 mm long, spurred. Spur cylindric, directed parallel with ovary and claw, rare at
wide angle with claw, 45 mm long, with 2 large peltate, warty glands. Mesochile
boat-shaped, with short claw, along flanges slightly erose or with few short irregular
denticles. Epichile lobes lanceolate to broadly-lanceolate, 810 mm long, 1.21.8 mm
wide. Column 34 mm tall, at front with 2 narrowly-triangular, acute wings, not entering
to the spur and short bifid rostellum. Anther ovate, acute, 4 mm long. Fig. 28, m-p; 32, b.
Ecology. Closed humid broad-leaved forests on silicate soils. 6002000 m. Fl.
November January. Rare (VU).
Distribution. Vietnam (Dak Nong, Gia Lai, Khanh Hoa, Kon Tum, Lam Dong,
Quang Tri), Thailand.
Studied specimens. Da Tan La river, Col de Pren, Tixier drawing (P); Dak Nong,
Dak Glong, Ta Dung mt., HLF 5656 (HN, LE); Dalat, Evrard 1944, 2029 (P); Gialai-Kontum,
1985, Averyanov et al., LX-VN s.n. (HN, LE); Khanh Hoa, Khanh Son, Hon Giao ridge, VH 4276
(HN, LE); Kon Tum, Dak Gley, VH 2278a (HN, LE); Kontum, Kon Plong, Mang La, VH 5599a
(HN, LE); Kontum, Sa Tay, Chu Mom Ray, P-7906a (HN, LE); Langbian, Rolfe; Phu Sapoum,
Blao, Poilane 23804 (P); Quangtri, Poilane 11288 (P).
7. A. brevilabris Lindl. 1840, Gen. Sp. Orchid. Pl.: 499. A. griffithii Hook.
f. 1890, Fl. Brit. Ind. 6: 96. A. sikkimensis King et Pantl. 1896, Journ. As. Soc.
Bengal, 65: 124; id. 1898, Ann. Bot. Gard. (Clcutt) 8, tab. 391; Aver. 1994, Identif.
Guide Vietnam. Orch.: 46; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.:
11. A. chapaensis Gagnep. 1931, Bull. Mus. Hist. Nat. (Paris) 2 ser. 3, 7: 679;
Seidenf. 1992, Opera Bot. 114: 33; Aver. 1994, Identif. Guide Vietnam. Orch.: 44;
Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 10. A. tridentatus Seidenf.
1992, Opera Bot. 114: 33, fig. 11; Aver. 1994, Identif. Guide Vietnam. Orch.: 44; Aver.
et Averyanova, 2003, Updated Checklist Orch. Viet.: 11.
Described from NE India (India, Assam). Type (Mack s.n.) K.
Terrestrial herb with 34 petiolate leaves in basal rosette. Leaves dark green,
with yellow to pink nerve reticulation, ovate, to broadly-ovate, shortly acute, to 6 cm
long. Scape pubescent, to 15 cm long, with 16 lax flowers and 23 acuminate sterile
bracts. Floral bracts ovate, acuminate, 610 mm long. Sepals sub-similar, olive-green,
oblong, obtuse, 68 mm long. Petals oblique-falcate, as long as median sepal. Lip
white, spurred, 810 mm long; spur short, 34 mm long, broadly-conical, bifid at the
apex, with 2 rather small peltate glands. Mesochile with 34 pairs of short blunt teeth
on flanges. Epichile lobes divergent, flabellate-triangular, truncate, 78 mm long. Column
34 mm tall, with 2 narrowly-triangular, obtuse, parallel wings at front and broad bifid
acute rostellum. Fig. 29, a, b.
Ecology. Humid broad-leaved forests on silicate soils. 9001500 m. Fl.
December February. Rare (EN).
Distribution. Vietnam (Lao Cai, Vinh Phuc), Bhutan, NE India.
Studied specimens. Tamdao, Hayata s.n. (TI); Tonkin: Cha-p, Petelot 5147 (P);
Vinh Phu, Tam Dao, LX-VN 874 (HN, LE).
104 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Cheirostylis Blume, 1825, Bijdr. 6, fig. 16; id. 1825, ibid. 8: 413.
Type: C. montana Blume
Small terrestrial, lithophytic and epiphytic sympodial herbs. Rhizome creeping,
with very swollen, succulent internodes and roots modified into ridges or pillows densely
covered by root hairs. Stem ascending, leafy at the base, with erect scape and lax
terminal spike. Leaves few, spirally arranged, petiolate or sessile, sheathing at the
base, glabrous, convolute, usually dull green, acute. Scape leafless, with 15 sterile
hyaline bracts. Flowers small, sessile, resupinate, hardly opening. Sepals sub-similar,
connate for half of their length to form a swollen tube. Petals connivent or joined to
median sepal forming hood. Lip spurless. Hypochile concave, with various papillae on
lateral sides, or lacking any glands. Epichile reflexed, 2-lobed, rarely entire, connected
to hypochile by entire simple short claw. Column short, erect, with 2 thin prominent
stelidia and 2 rostellar arms. Anther triangular-ovate, acute. Pollinia 2, sectile, with
stipe and viscidium. Stigma with 2, small, convex, separated lobes.
50(13) species. Subtropical and tropical zone of the Eastern Hemisphere with
main center of diversity in mainland SE Asia.
Key to species
1. Leafy stem creeping; leaves distant, distichous, sessile, 47 mm long ....... 1. C. serpens
Leafy stem ascending to erect; leaves rosulate, spirally arranged, petiolate, usually
longer than 7 mm ................................................................................................................. 2
2. Inflorescence sub-sessile, scape less than 1 cm long, much shorter than apical leaves
................................................................................................................................... 2. C. inabai
Leafy stem ascending to erect; leaves rosulate, spirally arranged, petiolate, usually
longer than 7 mm; scape taller than 2 cm ........................................................................... 3
3. Stem erect; leaves distant in lower part of stem, erect to sub-erect, to 6 cm long,
cuneate at the base, sub-succulent, brightly uniform green, glossy; stigma lobes
cylindric-conical ..................................................................................................... 3. C. foliosa
Stem ascending; leaves usually rosulate at the base of stem, spreading, less than
4 cm long, broadly-triangular to cordate at the base, thin, velvety dull-green to dark
green, commonly with dark veins; stigma lobes hemispheric ........................................ 4
4. Epichile narrow, simple, ligulate-oblong, longer than broad; hypochile at the base
without glands or with hardly visible rudimentary warts ................................ 4. C. takeoi
Epichile distinctly broader than long, often 2-lobed, with toothed or fringed lobes;
hypochile at the base with slender finger-like glands placed along lateral veins ....... 5
5. Lobes of epichile deeply toothed of fringed along margin ............................................. 6
Lobes of epichile entire, finely irregular denticulate or shallowly crenulate ............... 10
6. Scape 69 cm long with 12 flowers .................................................................................. 7
Scape normally longer than 9 cm, with 110 flowers ....................................................... 8
7. Leaves glossy, more or less rigid, silvery-gray to light green, with network of dark
green nerves; petals narrowly-obovate to oblanceolate; epichile lobes with 58
Turczaninowia 2008, 11(1) : 5168 105
triangular dents; hypochile with numerous fat conic glands along lateral veins .................
....................................................................................................................... 5. C. marmorifolia
Leaves uniform dull gray-green, thin and soft; petals filiform-linear; epichile lobes
with 4 finger-like fringes; hypochile with 2 slender glands along lateral veins .....................
............................................................................................................................... 6. C. filipetala
8. Sepals white, 3.55 mm long; epichile 1.53.5 mm long and wide; epichile lobes usually
with 47 dents; hypochile on sides with 2(3) glands ..................................... 7. C. chinensis
Sepals light olive-green to greenish-pink, 4.5-6.5 mm long; epichile 47 mm long and
wide; epichile lobes usually with 712 dents or fringes; hypochile on sided with 35
or more glands ..................................................................................................................... 9
9. Epichile lobes deep lacinate, with long finger-like fringes; leaves usually dark green
with light green median stripe .............................................................. 8. C. cochinchinensis
Epichile lobes deeply dentate, with narrowly-triangular teeth; leaves usually light
dull gray-green with dark veins .................................................................. 9. C. yunnanensis
10. Scape less than 5 cm tall; lip 55.5 mm long; epichile entire, transverse-rhomboid ........
..................................................................................................................... 10. C. thanmoiensis
Scape normally taller than 5 cm; lip longer than 6 mm; epichile distinctly emarginate or
2-lobed ................................................................................................................................... 11
11. Epichile 23 mm wide, emarginate to 2-lobed, with entire semi-circular lobes .....................
.......................................................................................................................... 11. C. bipunctata
Epichile more than 5 mm wide, distinctly 2-lobed, with flabellate to oblique-rectangular
lobes, crenulate or denticulate along margin ................................................................. 12
11. Epichile lobes spreading, to 4 mm long, shallowly crenulate or broadly dentate, with
24(5) blunt-roundish indistinct dents; petals obovate, much broadening to the apex
............................................................................................................................ 12. C. latipetala
Epichile lobes convergent, 1014 mm long, irregular denticulate, with low irregular rounded
teeth; petals elliptic, narrowing to the base and apex .............................. 13. C. spathulata
2. C. inabai Hayata, 1914, Ic. Pl. Formos. 4: 108, fig. 56. C. pingbianensis
K.L. Lang, 1996, Acta Phytotax. Sin. 34, 6: 635. C. phamhoangii N.T. Tich, 2001,
Orch. Vietnam, 1: 197, nom. inval.
Described from Taiwan (Rinkoho, ad 5000 ped. alt.). Type (H. Inaba,
Aug., 1911) TI?
Terrestrial creeping herb with ascending fleshy pink stem 1015 cm tall. Leaves
distant and small at the base, clustered at the apex with 45 narrowly-ovate, attenuate,
petiolate, green leaves, 34 cm long, 1.52 cm wide. Petiole 11.5 cm long. Inflo-
rescence very short, 12 cm long, secund, with 37 flowers, clustered in dense head-
like spike. Ovary narrowly-obovoid, to 1 cm long, glabrous. Flowers pure white. Sepals
sub-similar, forming cylindric tube 1 cm long, with 3 teeth at apex. Petals broadly-
lanceolate, 1 cm long, 3 mm wide. Lip 1.3 cm long. Hypochile saccate, boat-shaped,
at the base with few small close glands along each lateral vein. Epichile conduplicate,
broad, 2-lobed, 68 mm wide; lobes spreading; each lobe dissected into 5 close finger-
like segments 34 mm long, 0.50.6 mm wide. Column short. Stelidia narrow-parallel,
4 mm long, sub-equal in length with rostellum arms. Anther broafly-ovate, much shorter
than rostellum. Fig. 29, f-i.
Ecology. Ericaceous thickets on silicate soils along tops of ridges. 20002150 m.
Fl. September. Very rare (EN).
Distribution. Vietnam (Lam Dong), S China, Taiwan.
Studied specimens. Lam Dong, Lac Duong, Langbian mt., T.07.09.93a (icon
Cheirostylis phamhoangii, N.T. Tich, 2001, l.c.: 199).
Note. Very rare species known in Vietnam by the only collection.
and wide. Hypochile concave, cup-like, 1.5 mm across, with 1 large longitudinal cristate
straight ridge along lateral veins. Epichile papillose, 1.2 mm wide, 2-lobed, lobes square,
entire or with 2 indistinct rounded dents. Column 2 mm long, 1.4 mm wide; anther cup
lanceolate; rostellum arms narrowly spathulate, slightly shorter than anther cup; stelidia
shorter, thick, cylindrical, obtuse, with papillose surface. Fig. 29, j-o; 32, d.
Ecology. Broad-leaved forests on rocky limestone, commonly on mountain
tops. 12001300 m. Fl. September November. Very rare (EN).
Distribution. Vietnam (Son La).
Note. Very rare species known by the only type collection.
Fig. 29. Anoectochilus brevilabris: a flattened tepals and lip, b column, frontal and side view;
Cheirostylis serpens: c flowering plant, d flattened tepals, e column, side view; C. inabai:
f flowering plant, g flowers, h flattened sepals, petal and lip, i column and lip, side view;
C. foliosa: j flowering plants, k flower, l, m flattened tepals and lip, n column, dorsal,
frontal and side views, o floral bract; C. takeoi: p flowering plant and leaf, q flower, r
flattened tepals, s lip, side view.
Turczaninowia 2008, 11(1) : 5168 109
Dwarf lithophytic herb with creeping chain rhizome and ascending short stem
to 6 mm tall with 24 rosulate leaves. Leaves shortly petiolate, ovate, rigid, to 2 cm
long, light green to silvery white with contrasting dark green veins. Scape erect,
pubescent, 68 cm tall, with 12 flowers. Floral bracts pubescent, 34 mm long, shorter
than ovary. Ovary densely pubescent, 4 mm long. Sepals olive-green, pubescent, ovate,
5 mm long, joined on 2/3 of their length forming tube. Petals white, oblanceolate, falcate,
5.5 mm long. Lip 8 mm long, white, with 2 green spots at the base of epichile. Hypochile
concave, with fleshy central keel and numerous fleshy glands on each side. Epichile
2-lobed, lobes deeply fringed, with 67 cuneate teeth. Column 2.5 mm tall. Fruit ovoid
capsule 4 mm long. Fig. 30, a-e; 32, f.
Ecology. Wet broad-leaved and coniferous mossy forests on rocky limestone.
4001500 m. Fl. December March. Rare (EN).
Distribution. Vietnam (Bac Kan, Cao Bang, Son La).
Studied specimens. Bac Kan, Ba Be national park, HLF 508 (HN, LE), HLF 650
(HN, LE); Cao Bang, Bao Lac, CBL 1485 (HN, LE); Cao Bang, Tra Linh, 27 May 1997,
L. Averyanov, N.T. Hiep, VH s.n. (LE); Son La, Yen Chau, DKH 7203 (HN, LE).
Terrestrial and lithophytic herb. Stem 630 cm tall, with 15 leaves at the
base. Leaves dull gray-green, narrowly-ovate to ovate-cordate, acute, 13 cm long.
Scape tomentose, to 22 cm long. Rachis to 1.5 cm long, tomentose, with 110 flowers.
Floral bracts triangular-ovate, acuminate, 510 mm long, sparsely hairy to sub-glabrous,
longer than ovary. Ovary narrowly-obovoid, sparsely pubescent, 47.5 mm long. Sepals
white to light greenish, sub-similar, narrowly-ovate, obtuse, 45 mm long, joined on 1/2
2
/3 of their length, sparsely hairy to sub-glabrous. Petals white, narrowly-obovate,
oblique-falcate, obtuse, narrowing to the base, same length as sepals. Lip 67 mm
long, white, with 2 greenish spots at the base of epichile. Hypochile concave, boat
shaped, at the base with 2(3) slender finger-like recurved glands along lateral veins.
Epichile 2-lobed; lobes spreading, flabellate, 1.53.5 mm long and wide, with 47
triangular to narrowly-triangular teeth. Column 22.5 mm tall, with sub-equal narrowly-
triangular, acute stelidia and subulate rostellar arms. Fruit obovoid capsule to 9 mm
long. Fig. 30, j, k; 32, g.
Ecology. Wet broad-leaved and coniferous mossy forests on rocky limestone.
3001550 m. Fl. December April. Not rare (LR).
Distribution. Vietnam (Bac Kan, Ha Giang, Lang Son, Quang Binh, Son
La), Myanmar, Hong Kong, Taiwan.
Studied specimens. Bac Kan, Cho Don, HLF 744 (HN, LE); Ha Giang, Dong Van,
HAL 8598 (HN, LE); Ha Giang, Quang Ba, DKH 4894 (HN, LE, MO), DKH 5098 (HN, LE, MO);
Ha Giang, Vi Xuyen, DKH 6309 (HN, LE, MO), DKH 6511 (HN, LE, MO); Lang Son, Bac Son,
HAL 6708 (HN, LE); Quang Binh, Bo Trach, HAL 6056 (HN, LE), HAL 6114a (HN, LE), HAL
6196 (HN, LE), HAL 6352 (HN, LE); Quang Binh, Minh Hoa, HAL 5869 (HN, LE); Son La, Moc
Chau, DKH 7509 (HN, LE, MO); Son La, Yen Chau, DKH 7111 (HN, LE, MO).
Note. Typical element of herbaceous lithophytic vegetation on mossy rocky
solid limestone.
8. C. cochinchinensis Blume, 1859, Coll. Orch. Arch. Ind. Jap.: 55; Seidenf.
1992, Opera Bot. 114: 35; Aver. 1994, Identif. Guide Vietnam. Orch.: 41; Aver. et
Averyanova, 2003, Updated Checklist Orch. Viet.: 20. C. flabellata auct. non Wight:
Gagnep. 1933, Fl. Gen. Indochine 6, 4: 547.
Described from southern Vietnam (Tourane, Vietnam). Type (Gaudichaud
62) P.
Terrestrial and lithophytic herb. Stem 825 cm tall, with 14 leaves at the
base. Leaves dark green with light green median stripe, triangular-ovate to cordate,
acute, 13.5 cm long. Scape sparsely hairy, to 22 cm long. Rachis to 1.5 cm long,
glabrous or sparsely hairy, with 110 flowers. Floral bracts triangular-ovate, acuminate,
510 mm long, glabrous, shorter than ovary. Ovary narrowly-obovoid, glabrous, 5
8 mm long. Sepals dull greenish to pink, sub-similar, narrowly-ovate to ovate, obtuse,
4.56 mm long, joined on 1/22/3 of their length, glabrous. Petals white, narrowly-
obovate, oblique-falcate, obtuse, narrowing to the base, same length as sepals. Lip 8
12 mm long, white, with triangular green papillose area at the base of epichile. Hypochile
concave, boat shaped, at the base with 35 slender finger-like recurved glands along
Turczaninowia 2008, 11(1) : 5168 111
lateral veins. Epichile 2-lobed; lobes spreading, flabellate, 46 mm long and wide,
deeply laciniate into 610 narrowly-triangular finger-like segments. Column 2.53 mm
tall, with sub-equal narrowly-triangular, acute stelidia and subulate rostellar arms. Fig.
30, l; 32, h.
Ecology. Wet broad-leaved and coniferous mossy forests on rocky limestone.
700900 m. Fl. January February. Occasional (VU).
Distribution. Vietnam (Quang Binh, Quang Nam Da Nang), Taiwan.
Studied specimens. Quang Binh, Bo Trach, HAL 6197 (HN); d-EXSICCATES OF
VIETNAMESE FLORA 0006/HAL6197.
Note. Typical element of herbaceous lithophytic limestone vegetation in central
Vietnam.
9. C. yunnanensis Rolfe, 1896, Kew Bull. 1896: 201; Seidenf. 1992, Opera
Bot. 114: 35; Aver. 1994, Identif. Guide Vietnam. Orch.: 41; Aver. et Averyanova,
2003, Updated Checklist Orch. Viet.: 20.
Described from S China (Yunnan: ... at Mengtse). Type (Hancock, 25) K.
Terrestrial and lithophytic herb. Stem 830 cm tall, with 15 leaves at the
base. Leaves triangular-ovate to cordate, acute, 14 cm long. Scape tomentose, to
25 cm long. Rachis to 1.5 cm long, tomentose, with 18 flowers. Floral bracts triangular-
ovate, acuminate, 511 mm long, tomentose, longer than ovary. Ovary narrowly-
obovoid, pubescent, 48 mm long. Sepals dull greenish to pink, sub-similar, narrowly-
ovate to ovate, obtuse, 56.5 mm long, joined on 1/22/3 of their length, pubescent.
Petals white, narrowly-obovate, oblique-falcate, obtuse, narrowing to the base, same
length as sepals. Lip 812 mm long, white, with 2 green spots at the base of epichile.
Hypochile concave, boat shaped, at the base with 35 slender finger-like recurved
glands along lateral veins. Epichile 2-lobed; lobes spreading, flabellate, 47 mm long
and wide, with 610 triangular to narrowly-triangular teeth. Column 2.53 mm tall,
with sub-equal narrowly-triangular, acute stelidia and subulate rostellar arms. Fruit
obovoid capsule to 12 mm long. Fig. 30, m, n; 32, i.
Ecology. Wet broad-leaved and coniferous mossy forests on rocky limestone.
501200 m. Fl. February April. Not rare (LR).
Distribution. Vietnam (Bac Kan, Cao Bang, Ha Giang, Hanoi, Lang Son,
Quang Binh, Tuyen Quang), NE India, S China, Thailand.
Studied specimens. Bac Kan, Ba Be national park, HLF 510 (HN, LE); Cao Bang,
Bao Lac, CBL 1371 (HN, LE); Ha Giang, Dong Van, NTH 3526 (HN, LE); Ha Giang, Quang Ba,
DKH 4893 (HN, LE, MO), DKH 5097 (HN, LE, MO); Hai Phong, Cat Ba, LX-VN 3600 (HN, LE),
LX-VN 3426/16 (HN, LE); Lang Son, Bac Son, HAL 6709 (HN, LE); Quang Binh, Quang Ninh,
HAL 6307 (HN, LE); Quang Ninh, Ha Long Bay, NTH 5468 (HN, LE); Tonkin, Notre Dame, Tu-
vu, Balansa 2003 (P); Tuyen Quang, Na Hang, NTH 2060 (HN, LE).
Note. Vicarious species replacing C. cochinchinensis in limestone areas of
northern Vietnam where it is typical element of herbaceous lithophytic vegetation on
rocky solid limestone.
112 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
11. C. bipunctata Aver. 1996, Bot. Journ. (St. Petersburg), 81, 10: 78, fig. 4;
Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 20.
Described from northern Vietnam (N Vietnam, Prov. Ha Giang, Distr. Quan
Ba, around 4 km to SW from Can Ty town.). Type (Averyanov et al. HG 72, 27
October 1995) LE.
Lithophytic herb 1025 cm tall with 24 basal, shortly petiolate leaves. Leaf
blade dull silvery-green with unclear green vein network, narrowly-ovate to ovate, to
4 cm long. Scape and rachis pubescent, rachis 23 cm long, normally with 38 lax,
hardly opening flowers. Floral bracts triangular-ovate, acuminate, 48 mm long, not
longer than ovary, dense tomentose. Ovary 58 mm long, densely pubescent. Sepals
outside sparsely pubescent, light green, narrowly-ovate, 57 mm long, joined on 2/3 of
their length into a tube. Petals white or light greenish, free, lanceolate, 56.5 mm long,
1.5 mm wide. Lip white or light greenish, 67.5 mm long. Hypochile concave, basally
with numerous small glands on each side along lateral veins. Claw with 2 brightly
green spot on bended part. Epichile entire, emarginate, obcordate, or divided into 2
semi-circular lobes with entire margins. Column 1 mm tall. Anther red. Fruit obovoid,
810 mm long, 56 mm wide. Fig. 33, a-c.
Ecology. Wet broad-leaved and coniferous mossy forests on rocky limestone.
5001000 m. Fl. March April. Rare (VU).
Distribution. Vietnam (Ha Giang).
Studied specimens. Ha Giang, Quang Ba, DKH 4895/1 (HN, LE, MO).
Fig. 30. Cheirostylis marmorifolia: a flowering plant and leaf, b flower, c sepals and
petals, d flattened lip, e column, frontal and dorsal view; C. filipetala: f flowering plant
and leaf, g flower, h flattened tepals and lip, i fruit and floral bract; C. chinensis: j
flattened tepals and lip, k column, ovary and floral bract; C. cochinchinensis: l flattened lip;
C. yunnanensis: m flattened tepals and lip, n column, frontal view; C. thanmoiensis: o
flowering plant, p flower, q column and lip, side view.
114 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
village). Type (25 November 2002, D.K. Harder, N.T. Hiep, L. Averyanov, DKH
7659c) HN (holotype), LE (isotype).
Terrestrial and lithophytic herb. Stem 520 cm tall, with 15 leaves at the
base. Leaves velvety dark green to nearly black, petiolate, triangular-ovate to cordate,
acute, to 2 cm long. Scape tomentose, to 22 cm long, with 14 flowers. Floral bracts
triangular-ovate, acuminate, 58 mm long, much shorter than ovary. Ovary narrowly-
ellipsoid, sparsely pubescent, 56 mm long. Sepals white, glabrous, concave, obtuse, 4
mm long, dorsal sepal broadly ovate, lateral sepals narrowly-ovate, all joined on 2/3 of
their length. Petals white, much broadening in apical part, broadly obtuse to slightly
emarginate, 4 mm long, 1.5 mm wide. Lip 67 mm long, white, with 2 greenish spots
at the base of epichile. Hypochile concave, at the base with 23 slender finger-like
recurved glands along lateral veins. Epichile 2-lobed, 3 mm long, 67 mm wide; lobes
spreading, each with 34(5) roundish to broadly triangular blunt teeth. Column 2.5 mm
tall, with sub-equal narrow, acute stelidia and rostellar arms. Anther narrowly-ovate,
subacute, 1.5 mm long. Fig. 33, d-f; 36, a.
Ecology. Wet broad-leaved and coniferous mossy forests on rocky limestone.
4001300 m. Fl. November December, March. Occasional (LR).
Distribution. Vietnam (Bac Kan, Ha Giang, Son La).
Studied specimens. Ha Giang, Meo Vac, HAL 8478 (HN, LE), HAL 8526 (HN, LE);
Ha Giang, Quan Ba, HAL 8381 (HN, LE); Son La, Muong Lum, DKH 7111 (HN, LE, MO).
13. C. spathulata J.J. Smith, 1927, Bull. Jard. Bot. Buitenzorg, 3 ser. 9: 32;
Seidenf. 1992, Opera Bot. 114: 35; Aver. 1994, Identif. Guide Vietnam. Orch.: 40;
Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 20. C. grandiflora
auct. non Blume: Gagnep. 1933, Fl. Gen. Indochine, 6, 4: 548.
Described from Java (Java, Soerabaj: Goenoeng Kembangan bei Gresik).
Type (Dorgelo 2072) BO, L?
Stem 1520 cm tall, with 46 leaves at the base. Leaves pale gray-green, with
dark gray nerves, petiolate, narrowly-ovate to ovate, acute, 1.52.5 cm long. Scape
1216 cm long, tomentose, with 25 flowers. Sepals sub-similar, narrowly-ovate, 7
10 mm long, joined on 1/22/3 of their length, pubescent. Petals oblique-elliptic, narrowing
to the apex an base, the same length as sepals. Lip 1.51.7 cm long. Hypochile concave,
boat-shaped, at the base with 46 small terete glands along lateral veins. Epichile 2-
lobed, lobes divergent, triangular-flabellate, truncate, apex wavy, with low irregular
rounded teeth. Fig. 33, g; 36, b.
Ecology. Limestone at low elevations. Fl.? Occurrence in Vietnam needs
confirmation (EX?).
Distribution. Vietnam (probably extinct). Thailand, Java (extinct), Kalimantan.
Studied specimens. Cochinchina, Mt. Chao-lo, Pierre 145 (P).
Note. Insufficiently known species, occurrence of which in Vietnam needs
confirmation.
Turczaninowia 2008, 11(1) : 5168 115
Fig. 31. Corymborkis veratrifolia: a (LX-VN 3779); Tropidia angulosa: b (Averyanov 512
1988 and HAL 5261); T. curculigoides: c (DKH s.n., 1999 Ha Tinh); Anoectochilus roxburghii:
d (HAL 9735), e (HAL 3428, HAL 2303, HAL 2002); A. papillosus: f (HAL 8978, type);
Anoectochilus calcareus: g (DKH s.n., 2002, HAL 5866, HAL 1696, HAL 2994); A. alboli-
neatus: h (VH 4132); A. annamensis: i (HAL 8177, type).
116 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Fig. 32. Anoectochilus annamensis: a (HAL 6897); A. lylei: b (P 7906a); Cheirostylis serpens:
c (HAL 6092, type); C. foliosa: d (HAL 9415, type); C. takeoi: e (HAL 6708); C. marmorifolia:
f (Averyanov et al., CBL s.n., 1998, type and HLF 508); C. chinensis: g (HAL 6196 and HAL
8598); C. cochinchinensis: h (HAL 6197); C. yunnanensis: i (HAL 6389).
Turczaninowia 2008, 11(1) : 5168 117
Erythrodes Blume, 1825, Bijdr.: 410. Physurus L.C. Rich ex Lindl. 1840,
Gen. Sp. Orch. Pl.: 501.
Type: E. latifolia Blume
Terrestrial sympodial herbs with creeping succulent rhizome rooting at nodes,
ascending erect leafy floriferous stem and terminal many flowered raceme. Leaves
few, spirally arranged at the base of stem, petiolate, sheathing at the base, glabrous,
convolute, asymmetric, uniformly green, acute. Scape hairy, leafless, with 24 sterile
bracts. Flowers small, resupinate. Sepals free, pubescent, lateral spreading; petals
free or connivent to median sepal forming hood. Lip lying alongside the column,
concave, saccate or spurred, not hidden by the sepals; spur or sac with 1 small flat
protuberance or flat papillose wart on each side (usually breaking off in dry herbarium
material). Epichile in form of small reflexed lamina. Column narrow, erect, apically
setaceous, acuminate, with 1 hollow stigma and 2 sectile pollinia having caudicle and
viscidium.
20(2) species. Tropical and subtropical Southeast Asia to islands of SW Pacific.
Key to species
1. Lip narrowly-oblong, boat-shaped, longer than 5 mm, about 2.5 mm wide, glabrous, at
the base with bifid spur longer than 2.5 mm; epichile sub circular, as wide as lip;
sepals 5.57 mm long ............................................................................................. 1. E. hirsuta
Lip ovate, concave, 45 mm long, 4 mm wide, with dark papillose areas inside, at the
base saccate, sac less than 2.5 mm long; epichile broadly-obovate to triangular, as
broad as 1/2 of the lip; sepals 45.5 mm long ................................................... 2. E. blumei
at the apex, inside with 1 low insignificant callosity on each side. Column 45 mm
long. Fruit elliptic beaked capsule, 1215 mm long. Fig. 33, h; 36, c.
Ecology. Wet closed forests, often with bamboo, on any soils. 300500 m. Fl.
March May. Not rare (VU).
Distribution: Vietnam (Ha Tinh, Ninh Binh, Quang Tri). Bhutan, NE India,
Myanmar, Thailand, S China.
Studied specimens. Ha Tinh, Huong Son Distr., Son Hong Municipality, HAL
5215 (HN, LE); Ninh Binh, Cuc Phuong national park, M.V. Xinh et al. MVX 173, CP 1076 (HN,
LE); LX-VN 1766, 1767 (HN, LE); Quang Tri, Da Krong, HLF 5776 (HN, LE).
flowers in more or less dense, often secund spike. Sepals all parallel to the floral axis,
or 2 lateral sepals spreading. Petals oblique, usually connivent and forming hood with
median sepal, sometimes joined near their apex. Lip divided into hypochile and epichile.
Hypochile saccate, with bristly hairs or lamellar protuberance inside. Epichile simple,
entire, oblong-ligulate, usually reflexed. Column short, without appendages at base.
Rostellum long, deeply cleft into long arms. Pollinia 2, sectile, with caudicle and elongate
viscidium. Stigma entire, hemispheric, on the front of the column.
80100(9) species. Europe, Mainland Asia, Indonesia, N Australia, islands of
W Pacific, N Australia, North and Central America.
Key to species
1. Rhizome very short, floriferous stem erect; leaves more or less erect, broadly-lanceolate
to narrowly ovate; scape, rachis and flowers glabrous; inflorescence dense many
flowered spike; sepals 23 mm long ................................................................... 1. G. procera
Rhizome more or less long, creeping, floriferous stem ascending; leaves spreading,
narrowly-ovate to broadly-ovate; scape, rachis and flowers mre or less hairy;
inflorescence lax or sub-dense, few- to many-flowered spike; sepals longer than 3 mm ...... 2
2. Large plants, with inflorescence 0.51 m tall; spike with more than 30 flowers; leaves
1020 cm long, creeping stem 510 mm thick .................................................... 2. G. fumata
Plants smaller, with inflorescence less than 0.5 m tall; spike with less than 30 flowers;
leaves less than 10 cm long; creeping stem less than 5 mm thick .................................. 3
3. Leaves distinctly reticulated with white, yellow or pink veins ...................................... 4
Leaves uniform green, sometime with 1 median white stripe or mottled with white
irregular spots ...................................................................................................................... 5
4. Inflorescence many-flowered spike 37 cm long on scape longer than 5 cm; sepals 3
3.5 mm long, densely tomentose ......................................................................... 3. G. hispida
Inflorescence subsessile with 13 flowers on scape less than 1 cm long; sepals 18
22 mm long, glabrous or sparsely hairy ............................................................... 4. G. biflora
5. Inflorescence with 13 flowers 22.5 cm across, sepals glabrous, spreading, 12
14 mm long ........................................................................................................ 5. G. viridiflora
Inflorescence normally with 425 flowers, flowers less than 1 cm across, sepals
pubescent or sparsely hairy, not spreading, less than 13 mm long ............................... 6
6. Floral bracts exceeding half of flowers, longer than 1.5 cm; sepals usually longer than
10 mm; leaves 2.510 cm long ............................................................................... 6. G. foliosa
Floral bracts not exceeding half of flowers, shorter than 1.5 cm; sepals usually less
than 10 mm long; leaves 1.54 cm long ............................................................................. 7
7. Sepals 89 mm long, pubescent; leaves mottled with white irregular spots, rare pure
green ................................................................................................... 7. G. schlechtendaliana
Sepals less than 8 mm long, glabrous or sparsely hairy; leaves uniform dark green to
dark olive-green, sometimes with white median stripe .................................................... 8
8. Leaves dark uniform green; inflorescence dense secund spike with 1025 flowers;
hypochile glabrous, with 2 lamellate parallel protuberance near the base .... 8. G. arisanensis
Leaves dark green to dark olive-green, with white median stripe; inflorescence lax
spike, with few spirally arranged flowers; hypochile pilose with numerous bristly
hairs ............................................................................................................ 9. G. rhombodoides
120 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
1. G. procera (Ker-Gawl.) Hook. 1823, Exot. Fl. 1, 3, tab. 39; Seidenf. 1992,
Opera Bot. 114: 38; Aver. 1994, Identif. Guide Vietnam. Orch.: 36; Aver. et Averyanova,
2003, Updated Checklist Orch. Viet.: 37. Neottia procera Ker-Gawl. 1822, Bot.
Reg. 8, tab. 639.
Described from Nepal (Nepal). Type icon (Wallich 619) K.
Erect terrestrial and lithophytic herb up to 60 cm tall, with 48 petiolate erect
leaves in basal third. Leaves broadly-lanceolate to narrowly-ovate, 515 cm long, 1
4 cm wide, pure light green. Scape glabrous, with 24 sterile narrow, acuminate bracts
and dense, many-flowered spike 520 cm long; scape finely papillose. Floral bracts
glabrous, ovate-lanceolate, acuminate, 58 mm long, ciliate along margin. Ovary
glabrous, 35 mm long. Flowers hardly opening, sub-globose, 34 mm across. Sepals
similar, concave, ovate, 23 mm long, glabrous, light greenish, brownish or olive-green.
Petals white, as long as sepals, oblique-spathulate. Lip white, sub-globose, saccate,
1.52 mm long, with few slender glands within; apex triangular, blunt recurved, with 2
fleshy tubercles at the base. Column 11.5 mm high, with acute rostellar arms, and
ovate, acute anther. Fruit ovoid capsule, 45 mm long. Fig. 33, j, k; 36, e.
Ecology. Wet open mossy places among secondary grasslands and open shrubs,
commonly along rocky stream valleys and on spring seepages, on any kinds of soils.
1001600 m. Fl. February May. Common (LR).
Distribution. Vietnam (probably all provinces), Sri Lanka, Nepal, Bhutan,
India, Japan, China, Taiwan, Hainan, Myanmar, Thailand, Laos, Cambodia, Malaysia,
Java, Sumatra, Philippines.
Studied specimens. Bac Kan, Ngan Son, CBL 1259 (HN, LE); Binh Tri Thien, Bo-
giang, Dak Lac, Krong Bong, VH 6461 (HN, LE); Gia Lai Kontum, Dakgley, Bien 565 (HN,
LE), Phoong 426 (HN, LE); Gia Lai Kontum, Kbang, LX-VN 2085 (HN, LE); Gia Lai Kontum,
Kon Ha Nung, LX-VN 1991 (HN, LE); Ha Giang, Bac Me, HAL 6608 (HN, LE); Ha Giang,
Quang Ba, DKH 4834 (HN, LE), Bien 14248 (LE); Hoang Lien Son, Sapa, 19.5.61, sine coll.,
2312, 2318 (LE); Kon Tum, Chu Mom Ray, DKH 4553 (HN, LE); Kon Tum, Ngoc Linh mt, VH
297 (HN, LE), VH 1190 (HN, LE); Lai Chau, Muong Te, HAL 10120 (HN, LE); Lai Chau, Phong
Tho, HAL 10713 (HN, LE); Ninh Thuan, Nui Chua mt., HLF 4179 (HN, LE); Quang Tri, Da
Krong, HLF 5756 (HN, LE), HLF 6163 (HN, LE); Quang Tri, Huong Hoa, HLF 5901 (HN, LE),
d-EXSICCATES OF VIETNAMESE FLORA 0080/HLF5901; Thua Thien Hue, A Luoi, HAL
7266 (HN, LE), HAL 7502 (HN, LE); Thua Thien Hue, Bach Ma national park, HLF 1732 (HN,
LE); Thua Thien Hue, Huong Thuy, HAL 8221 (HN, LE); Thua Thien Hue, Nam Dong, HAL
6924 (HN, LE), HAL 10867 (HN, LE); Thuan Hai, Ka-rom.
2. G. fumata Thwaites, 1861, Enum. Pl. Zeylan.: 314; Seidenf. 1992, Opera
Bot. 114: 28; Aver. 1994, Identif. Guide Vietnam. Orch.: 36; Aver. et Averyanova,
2003, Updated Checklist Orch. Viet.: 37. Allochilus eberhardtii Gagnep. 1932,
Bull. Mus. Hist. Nat. 2 ser. 4: 591.
Described from Sri Lanka (Ceylon... Central Province). Type (Thwaites
CP 3668) K (holotype), PDA (isotype).
Terrestrial and lithophytic herb with ascending fleshy leafy stem 610 mm
Turczaninowia 2008, 11(1) : 5168 121
thick and terminal inflorescence 50100 cm tall. Leaves 37; petiole 26 cm long;
leaf blade oblique-elliptic, acute, 1020 cm long, 48 cm wide, pure green. Scape
leafless, pubescent, with few narrow, acuminate sterile bracts, bearing dense, many-
flowered spike 1545 cm long. Floral bracts cuneate, 0.51 cm long. Flowers widely
opening, light brownish to olive-greenish, lip white. Ovary densely glandular pubescent,
11.2 cm long. Sepals sub-similar, narrowly ovate, obtuse, pubescent, 68 mm long;
lateral sepals reflexed. Petals oblique lanceolate-spathulate, dilated to the apex, as
long as sepals. Lip concave, cup-shaped, 56 mm long, 45 mm wide, hairy inside,
with attenuate narrowly-oblong, obtuse, much reflexed apex. Column stalked dilated
above with emarginate rostellum, 45 mm long. Fruits obovoid capsule, 11.2 cm
long, 45 mm wide. Fig. 33, l; 36, f.
Ecology. Wet closed forests on any kind of soils, commonly along humid
rocky stream canyons. 4002000 m. Fl. JanuaryApril. Occasional (LR).
Distribution. Vietnam (Bac Kan, Dak Nong, Kon Tum, Lam Dong, Ninh
Binh), Sri Lanka, India, Myanmar, China, Ryukyu Islands, Taiwan, Thailand, Malaysia,
Philippines, Java.
Studied specimens. Bac Kan, Ba Be, HLF 555 (HN, LE); Bac Kan, Cho Don, HAL
4696 (HN, LE); Bac Thai, Chu-bo; Siam-Khong, Dak Nong, Dak Glong, HLF 5610 (HN, LE); Ha
Nam Ninh, Cuc Phuong, 1989, Bao 631 (HN, LE), 20.03.1971, Trong 155 (HN, LE), N.M. Cuong,
H.L. Quyen, NMC 289 (HN), D.D. Soejarto et al. DDS 11264 (HN); sine date, MAJ 514 (HN);
Ha Nam Ninh, Cuc Phuong, May bac, LX-VN 1811 (HN, LE); Kon Tum, Dak Gley, VH 2344
(HN, LE); Kontum, Ngoc Linh mt., VH 833 (HN, LE); Lam Dong, Lac Duong, VH 3329 (HN, LE).
Note. Widespread and common species growing in shady wet woods at dif-
ferent elevations, commonly along stream and wet valleys. Rarely observed in flowers.
Typical herbaceous element of humid forest understory.
3. G. hispida Lindl. 1857, Journ. Linn. Soc. London (Bot.), 1: 183; Seidenf.
1992, Opera Bot. 114: 29; Aver. 1994, Identif. Guide Vietnam. Orch.: 36; Aver. et
Averyanova, 2003, Updated Checklist Orch. Viet.: 37.
Described from NE India (... Khasia Hills). Type (Hooker & Thomson
2110) K.
Terrestrial and lithophytic herb 620 cm tall. Leaves 25, petiolate; petiole
0.63 cm long; leaf blade ovate to lanceolate, acuminate, 414 cm long, 1.22.7 cm
wide, velvety dark brown-green, reticulated with silvery, yellow or pinkish veins, often
with light broad median stripe. Scape pubescent, with 24 sterile narrow, acuminate
bracts and dense, many-flowered spike. Rachis hairy, 37 cm long. Floral bracts
glabrous to sparsely hairy, lanceolate, acuminate, 711 mm long. Ovary densely hairy,
46 mm long. Flowers hardly opening. Sepals ovate, concave, 34 mm long, 11.5 mm
wide, hairy, olive-green, with white tips. Petals white, of the same length, oblique-
oblong, to lanceolate, with wavy obtuse apex. Lip white, concave, cup-shaped, 2
3 mm long, 2 mm wide, coarsely pubescent within, with attenuate recurved apex.
Column 1 mm high, with linear rostellar arms. Fruit broadly elliptic to almost round
capsule, 58 mm long. Fig. 33, m-o; 36, g.
122 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Ecology. Closed forests on any kind of soils. 1501500 m. Fl. June August.
Not rare (VU).
Distribution. Vietnam (probably all provinces), India, Thailand, Peninsular
Malaysia.
Studied specimens. Bac Kan, Cho Don, HAL 4700 (HN, LE), HLF 760 (HN, LE);
Bac Kan, Na Ri, HAL 4934 (HN, LE); Cao Bang, Bao Lac, CBL 310 (HN, LE); Cao Bang,
Nguyen Binh, CBL 1304b (HN, LE); Dak Lac, Chu Yang Sinh mt., VH 6102 (HN, LE); Ha Giang,
Bac Me, HAL 6574 (HN, LE), HAL 6421 (HN); Ha Giang, Quang Ba, DKH 4788 (HN, LE), DKH
4823 (HN, LE); Ha Giang, Vi Xuyen, DKH 6346 (HN, LE, MO); Ha Nam Ninh, Cuc Phuong,
Luan Van Bao 707 (HN, LE), LX-VN 1808 (HN, LE), 23 July 1965, Bich et al. 948, CPNP 5149
(HN), 25 March 1971, Hien B-50, CPNP 5123 (HN), 17 July 1971, Hop, Khoi 565, CPNP 5122
(HN), 12 July 2002, NMC 1680 (HN), sine date, Aung 784 (HN), sine date, Bich, Nhu, Liem 948
(HN), sine date, Dung 784, CPNP 5124 (HN), sine date, Hien B-51(HN), sine date, N.T. Hiep,
NTH 3892 (HN), sine date, Hop, Khoi 565 (HN); Hai Phong, Cat Ba, LX-VN 3446 (HN, LE);
Khanh Hoa, Hon Giao, 1997, VH s.n. (LE photo); Ninh Binh, Cuc Phuong, HAL 1659 (LE
photo); Ninh Thuan, Nui Chua national park, HLF 4286 (HN, LE); Quang Binh, Phong Nha-Ke
Bang national park, HAL 6189 (HN, LE photo); Son La, Moc Chau, HAL 9385 (HN); Son La,
Yen Chau, HAL 9463 (HN); Thanh Hoa, Ba Thuoc, HAL 337 (HN, LE), HAL 977 (HN), HAL
3010 (HN, LE), HAL 3182 (HN), HAL 3312 (HN, LE), HAL 3371 (HN), Thanh Hoa, Co Lung,
2003, HAL 3312 (LE photo); Thua Thien Hue, Bach Ma national park, HLF 1624 (HN, LE),
Lai Chau, Sin Ho, HAL 9881 (HN, LE).
Note. Typical shade-loving element of forest understory. It is easily distinguished
by its deep velvety brown-green leaves reticulated with silvery, yellow or pink veins.
The species is widely collected for traditional oriental medicine and as ornamental plant
known under common name jewel orchids. Needs special attention for protection.
4. G. biflora (Lindl.) Hook. f. 1890, Fl. Brit. India 6: 114. Georchis biflora
Lindl. 1840, Gen. Sp. Orch. Pl.: 496.
Described from Nepal? Type (Wall. Cat. 7379) K.
Terrestrial and lithophytic herb 58 cm tall. Leaves 46, shortly petiolate; leaf
blade ovate to broadly-ovate, acute, 3.55 cm long, velvety dark green, reticulated
with silvery veins. Inflorescence sub-sessile with hairy scape less than 1 cm long and
13 terminal secund flowers. Floral bracts glabrous to sparsely hairy, ovate to obovate,
shortly acute, 0.91.4 cm long. Ovary sessile, hairy, 812 mm long. Flowers odorless,
not widely opening. Sepals oblanceolate, concave, 1.82.2 cm long, sparsely hairy,
white to light pink. Petals white, of the same length, oblique-rhomboid, with finely
plicate margin. Lip white, oblong-ligulate, 1.82 cm long, with concave, coarsely hairy
base and recurved acute apex plicate along margin. Column erect 2.53.5 mm tall;
linear rostellar arms and narrowly cuneate anther close each other, to 1 cm long. Fig.
33, p-r; 36, h.
Ecology. Primary evergreen broad-leaved wet forests on silicate rocks. 1500
1800 m. Fl. April May. Very rare (VU).
Distribution. Vietnam (Lao Cai), Nepal, NE India, Japan, Korea, China,
Taiwan.
Turczaninowia 2008, 11(1) : 5168 123
Fig. 33. Cheirostylis bipunctata: a flowering plant and leaf, b flower, c flattened tepals
and lip; C. latipetala: d flowering plant, e flattened tepals, flattened lip and column, side
view, f ovary; C. spathulata: g flowers and base of spur; Erythrodes hirsuta: h lip, frontal
view; E. blumei: i flattened tepals, column and lip; Goodyera procera: j flower and lip, k
column, side view; G. fumata: l flattened tepals, flattened lip and column, half-side view;
G. hispida: m flowering plant, n flower without lip and petals, o lip; G. biflora: p
flattened tepals and lip, q column, side view, r floral bract; G. viridiflora: s flattened flower
without petals and lip, flattened lip and flattened petals.
124 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
6. G. foliosa (Lindl.) Benth. ex Clarke, 1889, Journ. Linn. Soc. London (Bot.)
25: 73; Seidenf. 1992, Opera Bot. 114: 29; Aver. 1994, Identif. Guide Vietnam. Orch.:
35; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 37. Georchis foliosa
Lindl. 1840, Gen. Sp. Orch. Pl.: 406.
Described from Myanmar (in regno Burmano). Type (Griffith s.n.) K.
Creeping terrestrial herb 1535 cm tall. Leaves 46, petiolate, oblique-elliptic,
acute, 410 cm long, 1.54 cm wide, pure green, often finely undulate along margin.
Petiole 14 cm long. Scape leafless, with dense, many-flowered slightly secund spike.
Rachis sparsely hairy, 1.510 cm long. Floral bracts ovate to lanceolate, acuminate,
glabrous, 1.52 cm long, as long as half of flowers or longer. Ovary densely hairy, 5
12 mm long. Flowers odorless, not widely opening. Sepals sub-similar, olive-brown to
pinkish, ovate, concave, obtuse, 913 mm long, pubescent. Petals white, oblique
rhomboid-spathulate, 812 mm long, 23 mm wide, forming hood with dorsal sepal.
Lip white to light greenish or pinkish, ovate, 710 mm long, 45 mm wide, with concave,
cup-shaped, long-papillose base and attenuate, obtuse white recurved obtuse apex.
Column 33.5 mm tall, with straight, finger-like rostellum arms and narrowly ovate
attenuate anther 44.5 mm long. Fruit ellipsoid capsule, 1014 mm long, 45 mm
Turczaninowia 2008, 11(1) : 5168 125
3495 (HN, LE); Hoa Binh, Mai Chau, DKH 7923 (HN, LE, MO); Kontum, Ngoc Linh mt., VH
581 (HN, LE); Lai Chau, Tam Duong, HAL 10186 (HN), HAL 10341 (HN, LE); Lam Dong, 14 km
to NNW from Dalat, VH 2576 (HN, LE); Lam Dong, Bi Doup Nui Ba national park, HLF 5257
(HN, LE); Lam Dong, Lac Duong, Bi Doup mt., HLF 5257 (HN, LE), d-EXSICCATES OF
VIETNAMESE FLORA 0038/HLF5257; Lam Dong, Lac Duong, Bi Dup mt., VH 3304 (HN,
LE), VH 4454 (HN, LE); Lam Dong, Lac Duong, VH 2868 (HN, LE); Lam Dong, peak of Bi Dup
mt., VH 2999 (HN, LE); Lao Cai, Fan Si Pang mt., NTH 2317a (HN, LE).
concave, obtuse to blunt. Column 2.53 mm tall, with 2 short rostellar wings. Anther
narrowly-ovate, acuminate, 1.61.8 mm long. Fig, 34, eh.
Ecology. Primary broad-leaved evergreen forests on granite and shale. 1900
2400 m. Fl. September November. Very rare (DD).
Distribution. Vietnam (Lai Chau).
Note. Very rare species known by the only type collection. On floral structure
it resembles Goodyera foliosa, but distinctly differs in twice smaller flowers, small
size of plant and in small, dark velvety green leaves with distinct white longitudinal
median stripe.
1. H. longicaulis Lindl. 1833, Bot. Reg. 19, tab. 1618; Seidenf. 1992, Orch.
Indochina: 28; Aver. 1994, Identif. Guide Vietnam. Orch.: 33; Aver. et Averyanova,
2003, Updated Checklist Orch. Viet.: 40.
Described from Nepal (Napalia). Type (Wallich 7389) K, C.
Ecology. Broad-leaved forests on silicate soils. 15002000 m. Fl. September
November. Very rare (DD).
Distribution. Vietnam (Lam Dong), Nepal, Bhutan, NE India, Myanmar, S
China, Thailand, Sumatra.
Studied specimens. Annam, sine loc, Vernet, 12 (P); Dankia, Langbian, Poilane,
18666 (P).
Note. Very rare species not observed in Vietnam during last decades.
Hetaeria Blume, 1828, Fl. Jav. Orch. 1 Pref.: 7, nom. emend. id. 1858, Coll.
128 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Orch. Arch. Ind. Jap.: 102, tab. 32; Seidenf. 1978, Dansk Bot. Ark. 32, 2 : 94104.
Etaeria Blume, 1825, Bijdr. 6, fig. 14; id. 1825, ibid. 8: 409, orth. var. rej.
Lectotype: H. oblongifolia (Blume) Blume (Etaeria oblongifolia Blume)
Terrestrial and lithophytic sympodial herbs with creeping succulent rhizome
rooting at nodes, ascending erect leafy floriferous stem and terminal many-flowered
raceme. Leaves few, spirally arranged, petiolate, sheathing at the base, glabrous,
convolute, asymmetric, uniformly green, acute. Scape leafless, with 24 sterile bracts.
Flowers small, not or hardily resupinate. Sepals free; petals free or connivent to median
sepal forming hood. Lip spurless. Hypochile concave with various papillae on lateral
sides. Epichile straight, acute or reflexed in form of small circular plate. Column short,
erect, with prominent rostellar arms and two wing-like appendages at front. Anther
narrowly triangular-ovate, acute. Pollinia 2, sectile, with stipe and viscidium. Stigma
with 2 convex, separated lobes.
30(5) species. Tropical Africa and Asia, Australia, Pacific islands to Tahiti.
Key to species
1. Epichile of the lip minute, less than 5 mm wide, heart-shaped, triangular acute or
obtuse .................................................................................................................................... 2
Epichile of the lip large, 68 mm wide, with large orbicular upwards-backwards turned
white lobules ....................................................................................................... 1. H. anomala
2. Wings of column incurved or folded, not straight, not reaching apex of rostellar arms ....... 3
Wings of column narrow, straight, widening into apical spathulate prolongation as
long as rostellar arms, or longer ........................................................................... 2. H. alta
3. Hypochile with 56 slender papillae placed on each side of the sac .... 3. H. oblongifolia
Hypochile with 13 fat irregularly branching or flattened irregularly dented glands on
each side of the sac ............................................................................................................. 4
4. Sepals 45 mm long; petals narrowly-obovate, hammer-shaped, very oblique, widening
from narrow base into semicircular widening on upper edge faced to dorsal sepal ........
.................................................................................................................................... 4. H. affinis
Sepals 56 mm long; petals ovate to broadly-ovate, without semicircular widening,
not oblique or slightly oblique .............................................................................. 5. H. nitida
1. H. anomala (Lindl.) Reichenb. f. 1874, Trans. Linn. Soc. Bot. 30: 142;
Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 40. Aetheria anomala
Lindl. 1857, Journ. Linn. Soc. 1: 394. H. grandiflora Ridl. 1923, Journ. Mal. Br.
Roy. As. Soc. 1, 1: 98. H. rotundiloba J.J. Smith, 1926, Svensk Bot. Tidsskr. 20:
470. H. hainanensis T. Tang et F.T. Wang, 1974, Act. Phytotax. Sin. 1, 1: 34.
H. biloba (Ridl.) Seidenf. et Wood, 1992, Orch. Pen. Mal. Sing.: 95, fig. 38; Aver. et
Averyanova, 2003, l.c.: 40. Zeuxine biloba Ridl. 1909, Journ. Fed. Mal. St. Mus. 4: 73.
Described from NE India (Assam: Tangree). Type (Griffith s.n.) K (holotype).
Terrestrial or lithophytic herb up to 35 cm tall, with 35 narrowly-ovate leaves
58 cm long. Scape to 20 cm long, shortly hairy. Rachis white hairy, to 7 cm long, with
420 flowers. Bracts ovate, 57 mm long, shorter than ovary, acuminate. Sepals oblong,
56 mm long, olive-brown with white and red apex, lateral sepals spreading. Petals
Turczaninowia 2008, 11(1) : 5168 129
lanceolate, 5 mm long, white with reddish-brown base, connivent to median sepal. Lip
white with saccate greenish-pink base containing 2 short gland-like keels on each
side, mesochile short, channeled with inrolled edges, epichile with 2 recurved broadly
obovate lobes, each lobes 34.5 mm long, 2.5 mm wide. Column with short narrow
wings and prominent rostellum arms. Fig. 34, m, n; 37, d.
Ecology. Wet primary and secondary closed forests, often with bamboo, on
any soils. 8001300 m. Fl. January April, August. Not rare (VU).
Distribution: Vietnam (Dac Lac, Hoa Binh, Lao Cai, Quang Binh, So La),
NE India, Burma, Laos, S China, Malaya, Sumatra, Borneo, Sulawesi.
Studied specimens. Dac Lac, Chu Yang Sinh mt., 1996, Vu Ngoc Long s.n. (LE);
Hoa Binh, Da Bac, Phu Canh mt., HAL 396 (HN, LE); Lao Cai, Van Ban, HAL 2287, DKH 6987
(HN, LE); Quang Binh, Bo Trach, HAL 6060 (HN, LE); Quang Binh, Phong Nha Ke Bang
national park, HAL 6241 (HN, LE); So La, Yen Chau, DKH 7225 (HN, LE, MO).
3. H. oblongifolia (Blume) Blume, 1858, Coll. Orch. Arch. Ind. Jap.: 102,
tab. 32; Seidenf. 1992, Orch. Indochina: 39; Aver. 1994, Identif. Guide Vietnam. Orch.:
32; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 40. Etaeria
oblongifolia Blume, 1825, Bijdr. 6, fig. 14; id. 1825, ibid. 8: 410.
Described from Java (Solassie Provinciae Tjanjor). Type L (BO, P)?
Terrestrial herb up to 40 cm tall, at the base with 47 shining narrowly-ovate
leaves 46 cm long. Scape shortly hairy, to 16 cm long with many hardly opening
flowers. Floral bracts narrowly-cuneate, 35 mm long, shorter than ovary. Sepals light
yellowish, ovate, 33.5 mm long, hairy. Petals white, oblique narrowly-obovate, 3
3.5 mm long, connivent to median sepal. Lip 33.5 mm long, yellowish, with about 5
6 slender papillae on each side, epichile brightly yellow, triangular, fat, finely papillose,
apiculate. Column at front with strongly sigmoid, triangular-acute wings, shorter than
rostellar arms. Fig. 35, a-c; 37, f.
130 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Fig. 34. Goodyera foliosa: a flower, lip section and petals; G. schlechtendaliana: b flower,
lip section and petals; G. arisanensis: c flowering plant, d flattened tepals and lip section,
side view; G. rhombodoides: e flowering plant and leaf, f flattened tepals, g flattened lip
and lip section, side view, h column, dorsal and frontal views; Herpysma longicaulis: i
flowering plant, j flattened sepals and flattened petals, k lip, side view, l column, frontal
and half-side views; Hetaeria anomala: m flower, n flattened tepals, column and lip;
H. alta: o flattened tepals and column, p lip, frontal view, q column, side view.
Turczaninowia 2008, 11(1) : 5168 131
5. H. nitida Ridl. 1896, Journ. Linn. Soc. Bot. 32: 404; Seidenf. Wood, 1992,
Orch. Pen. Mal. Sing.: 95, fig. 37 g-i, pl. 2, d.
Described from Malacca Peninsular (Pulau Butong, Penang). Type (Curtis
s.n.) SING (holotype).
Terrestrial herb up to up to 45 cm tall, with 57 narrowly-ovate to ovate leaves
710 cm long. Scape hairy, to 25 cm long, with many hardly opening flowers. Floral
bracts cuneate, hairy, 58 mm long, shorter than ovary. Pedicel and ovary straight,
densely adpressed to the rachis. Sepals olive to pink, ovate, 56 mm long, hairy. Petals
132 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Ludisia A. Rich. 1825, in Bory de St.-Vincent, Dict. Class. Hist. Nat. 7: 437.
Type: L. discolor (Ker-Gawl.) A. Rich. (Goodyera discolor Ker-Gawl.)
Lithophytic and occasionally epiphytic sympodial herb with fleshy succulent
creeping rhizome and ascending short leafy stem with 26 petiolate leaves and terminal
erect leafless inflorescence. Petiole and sheath 13 cm long; leaf blade ovate to broad-
elliptic, 410 cm long, 1.54 cm wide, dark velvety green to almost black, with 17
yellow or pink longitudinal veins, occasionally reticulated with veins network. Scape
1018 cm tall, with 23 pale pink, hyaline bracts. Rachis 214 cm long, hairy, with 1
12 laxy flowers. Floral bracts cuneate, 11.5 cm long, a little shorter than ovary.
Flowers resupinate, widely open, 1.21.7 cm across, white, with yellow anther. Sepals
ovate, 10 mm long. Petals narrowly-ovate of same length, connivent to dorsal sepal
forming hood. Lip twisted, with short 2-lobed spur; spur inside with 2-lobed sessile
gland on each side; lip blade 79 mm long, clawed, claw channeled, epichile 2-lobed,
with spreading oblong lobes. Column twisted, stigma single on front of column. Fruit
ellipsoid capsule, 1.5 cm long, 45 mm wide. Pollinia 2, sectile, with caudicle and
viscidium. Fig. 35, h-k; 43, a-c.
Monotypic genus.
Described from S Vietnam (Dalat, cult. Hort. Bot. Parisiensis, 1959). Type
(Tixier M-918) P.
Rachis pubescent, laxy 45 flowered. Floral bracts ovate-elliptic, acute,
pubescent, 9.5 9 mm long, 4 mm wide. Pedicel and ovary pubescent, to 12 mm long.
Flowers glabrous, not resupinate. Sepals narrowly-ovate, acute, 8 mm long, 4 mm
wide; lateral sepals recurved, oblique. Petals oblique narrowly-ovate, 8 mm long, 3
mm wide. Lip 3-lobed, 8.5 long, 5 mm wide. Hypochile subquadrate, saccate, sidelobes
broadly rounded, with the apical quarter membranaceous; inside along median vein
hypochile with a low fleshy keel terminating apically in an arrow shape about 4 mm
long, 5 mm wide; hypochile glands terete with an enlarged apex. Claw linear, 4 mm
long. Epichile cordate, acute, 1.5 mm long, 2 mm wide. Column erect, 6.5 mm tall.
Anther basally ovoid, long rostrate, 5 mm long. Rostellar arms linear, 4 mm long.
Stigma ventral, at the base of rostellum. Column wings subquadrate, ventral, 1 mm
across. Fig. 35, l-n.
Distribution. Vietnam (Lam Dong?).
Note. Insufficiently known species described on the base of fragment of
inflorescence collected from alone cultivated specimen, origin of which needs
verification. It may be related to widely cultivated M. petola, from which differs in
broader narrowly-ovate petals and twice longer anther and rostellum.
Fig. 35. Hetaeria oblongifolia: a flattened tepals and lip, b column and lip, side view, c
column, frontal view; H. affinis: d flattened tepals and lip, e column, frontal view; H. nitida:
f flattened tepals and lip, g column, frontal view and section of spur base. Ludisia discolor:
h flowering plant, i flattened tepals and lip, j flower, k column, side view and lip;
Macodes cupida: l flattened sepals and petal, m flattened lip, n column, side view;
Myrmechis pumila: o flowering plant, p flattened sepals, petal and lip, q column and lip,
side view, r floral bract.
136 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
narrowed near the apex, pyriform, 5.8 mm long, 2.2 mm wide. Lip 5.6 mm long.
Hypochile saccate, with 1 prominent midvein and 1 bilobed gland on each lateral vein.
Epichile 2-lobed, with spreading oblique obovate lobes. Column short, with small horn-
like frontal wings. Fig. 35, o-r.
Ecology. Wet broad-leaved closed mossy forests on silicate soils. 21002150 m.
Fl. July. Very rare (EN).
Distribution. Vietnam (Kon Tum, Lam Dong), Bhutan, Nepal, NE India,
Myanmar, S China, Thailand.
Studied specimens. Kon Tum, Ngoc Linh mt., VH 627a (specimen lost); Lam
Dong, Lac Duong, Langbian mt., T.20.07.94b (icon Zeuxine langbianensis N.T. Tich, 2001,
l.c.: 215).
Odontochilus Blume, 1859, Coll. Orchid.: 79. Evrardi Ggnep. 1932, Bull.
Mus. Hist. Nat. (Paris) 2 ser. 4, 5: 596. Pristiglottis Cretz. et J.J. Smith, 1934, Acta
Fauna Fl. Universali, ser. 2, Bot. 1, 14: 4. Evrardianthe Ruschert, 1983, Feddes
Repert. 94: 433. Evrardian Aver. 1988, Bot. Journ. (Leningrd) 73, 3: 432.
Lectotype: O. flavescens (Blume) Blume (Anoectochilus flavescens Blume)
Terrestrial and lithophytic sympodial herbs with creeping succulent rhizome
rooting at nodes, ascending erect leafy floriferous stem and terminal lax, few- to many
flowered spike. Leaves few, spirally arranged, petiolate, sheathing at the base, glabrous,
convolute, asymmetric, uniformly green. Scape leafless, with 24 sterile bracts.
Flowers, sessile, resupinate or not. Sepals free. Petals very oblique, connivent to median
sepal forming hood, lateral sepals spreading, concave, enclosing saccate base of lip.
Lip adnate to column base. Hypochile concave to saccate to globose, with 2 fleshy
branched papillae on lateral sides, rare without glands. Mesochile channeled, long to
shortly fimbriate-toothed. Epichile with 2 divergent flabellate spreading lobes. Column
short, fleshy, winged or keeled, with 2 rostellar arms. Anther triangular-ovate, acute.
Pollinia 2, sectile, with stipe and viscidium. Stigma ventral, 1 or 2.
40(9) species. Tropical mainland Asia, Indonesia, New Guinea, Pacific Islands
to Polynesia and adjacent islands.
Key to species
Fig. 36. Cheirostylis latipetala: a (HAL 8526 and DKH 7659c, type); C. spathulata: b (Thai-
land, s. coll.); Erythrodes hirsuta: c (HAL 2266 and HLF 5755); E. blumei: d (Vietnam, s. loc.);
Goodyera procera: e (HAL 6924 and HLF 5756); G. fumata: f (LX-VN 1811); G. hispida: g (HAL
9385); G. biflora: h (HAL 11135); G. viridiflora: i (DKH 7400 and HAL 6057).
138 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Fig. 37. Goodyera foliosa: a (HAL 3967 and HAL 10027); G. schlechtendaliana: b (HLF 5257),
c (HAL 8404); Hetaeria anomala: d (HAL 6241); H. alta: e (HAL 7156); Hetaeria oblongifolia:
f (Averyanov 817, 1987); H. affinis: g (HLF 6591); H. nitida: h (HLF 6255 and NTH 5420G), i
(HLF 6255).
Turczaninowia 2008, 11(1) : 5168 139
4. Lip at the base straight, not concave or hardly concave, without any warts or glands
inside .............................................................................................................. 4. O. acalcaratus
Lip at the base distinctly concave to saccate, with distinct lateral warts or glands
inside ..................................................................................................................................... 5
5. Ovary glabrous; column with 2 stigmas ........................................................................... 6
Ovary tomentose or hairy; column with 1 stigma ............................................................ 7
6. Lip white, with convergent semi-circular to broadly-obovate lobes, slightly erose
along margin, 89 mm long; spike few-flowered, 24 cm long; hypochile glands more
or less straight, not uncinnate ........................................................................ 5. O. brevistylus
Lip yellow, with rather triangular-flabellate lobes, divided by broad sinus, entire along
margin, 912 mm long; spike many-flowered, 412 cm long; hypochile glands uncinnate
.......................................................................................................................... 6. O. lanceolatus
7. Ovary and sepals densely hairy; lip 1012 mm long; flanges of mesochile entire or
with few low insignificant teeth; lateral sepals not spreading, about 8 mm long; epichile
lobes 6 mm long .................................................................................................. 7. O. umbrosus
Ovary and sepals sparsely short hairy; lip 1620 mm long; flanges of mesochile with
35 long fringes; lateral sepals, 810 mm long; epichile lobes 812 mm long ............. 8
8. Sepals and petals uniform greenish, with 13 longitudinal veins, without distinct
pinnate venation from midvein ............................................................................. 8. O. tortus
Sepals and petals brownish to brownish-pink, with unclear whitish pinnate tessellation
and distinct pinnate venation from mid-vein ..................................................... 9. O. inabai
3. O. elwesii C.B. Clarke ex Hook. f. 1890, Fl. Brit. Ind. 6: 100. Anoec-
tochilus elwesii (C.B. Clarke ex Hook. f.) King et Pantl. 1898, Ann. Roy. Bot. Gard.
(Calcutta), 8: 297; Seidenf. 1992, Opera Bot. 114: 31; Aver. 1994, Identif. Guide Vietnam.
Orch.: 44; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 10. A. daoensis
Gagnep. 1951, Bull. Mus. Natl. Hist. Nat. 2 ser. 22, 5: 632; Seidenf. 1992, l.c.: 30;
Aver. 1994, Identif. Guide Vietnam. Orch.: 44.
Described from NE India (Darjeeling, Rishap; Khasia Hills, Shillong; Munipore,
Kohima, Naga Hills). Syntypes (Clarke 12251; Clarke 44609; Prain 17) K.
Terrestrial and lithophytic herb to 20 cm tall with 25 petiolate leaves at the
base. Leaves ovate, acute, purple-brown to black-violet, 1.54.5 cm long. Scape to 15
cm long, hairy, with 13 acute, hyaline bracts and 12(3) lax not resupinate flowers.
Floral bracts lanceolate, acuminate, hairy, to 12 mm long. Ovary 1015 mm long,
densely pubescent. Sepals olive to brown-green, long hairy, with acute, strongly recurved
apex; median sepal ovate, 911 mm long, with light pinkish apex; lateral sepals oblique-
ovate, 1012 mm long, with, concave, hemi-spherical base, embracing lip base. Petals
Turczaninowia 2008, 11(1) : 5168 141
white, hemi-circular, strongly oblique, broad at the middle, attenuate toward the apex,
as long as median sepal. Lip white, 1.62.5 cm long, basally globular, bifid, with 2
prominent conical glands. Mesochile pectinate with 46 purple-brown fringes. Epichile
lobes triangular-flabellate, irregular-undulate along margin, 11.5 cm long, 68 mm
wide, divided by a broad sinus. Column twisted, 23 mm tall, with 2 stigmas and 2
appendages at the base. Anther beaked. Fig. 38, i-o; 43, e.
Ecology. Primary and secondary wet broad-leaved forests, often with bamboo
on any kind of soils. 3001200 m. Fl. May July. Not common (VU).
Distribution. Vietnam (Bac Kan, Lai Chau, Lao Cai, Quang Binh, Thanh
Hoa, Tuyen Quang, Vinh Phuc), Bhutan, Nepal, NE India, Myanmar, Thailand, S China,
Taiwan.
Studied specimens. Bac Kan, Ba Be, HLF 612 (HN, LE); Bac Kan, Cho Don, HAL
4791 (HN, LE); Lai Chau, Sin Ho, HAL 8835 (HN, LE); Lao Cai, Van Ban, HAL 2003 (HN, LE);
Quang Binh, Minh Hoa, HAL 5957 (HN, LE); Thanh Hoa, Ba Thuoc, HAL 3372 (HN, LE);
Tuyen Quang, Na Hang, HAL 1720 (HN, LE); Vinh Phu, Tam Dao, 23 Jan. 1988, Averyanov et
al. LX-VN s.n. (LE), P-7452 (HN, LE).
Fig. 38. Odontochilus poilanei: a flowering plant, b flattened tepals and lip, c column,
side view; O. saprophyticus: d flowering plant, e flattened tepals and lip, f fragment of
mesochile, g column, side view, h pollinarium; O. elwesii: i inflorescence, j flower and
flower without lateral sepals and epichile, k section of spur and hypochile, l flattened lip,
m section of spur base, n column, frontal view, o pollinarium; O. acalcaratus: p flowering
plant, q flower and fragment of inflorescence, r flower without lateral sepal and column with
lip, side view, s flattened tepals and lip, t column, ventral and dorsal views.
Turczaninowia 2008, 11(1) : 5168 143
Herb to 30 cm tall with 56 distant pure green petiolate leaves. Leaves nar-
rowly-ovate to ovate, acute, 4.57 cm long. Scape short, 24 cm long, with lax spike
24 cm long of 610 flowers. Floral bracts narrowly-ovate, acuminate, to 10 mm long,
a little shorter than ovary. Ovary glabrous. Sepals sub-similar, ovate, 78 mm long,
green, sparsely hairy; lateral sepals swollen at the base, enclosing base of lip. Petals,
strongly oblique, falcate with attenuate apex, as long as median sepal. Lip white, 10
14 mm long, basally shortly globular, bifid, with prominent longitudinal ridge and flat
quadrangular gland on each side. Mesochile 78 mm long, laterally with 34 long
fringes. Epichile lobes 89 mm long, 56 mm wide, semi-circular to broadly-obovate,
slightly irregularly toothed, divided by broad sinus. Column short, 2.53 mm tall, with 2
stigmas, short rostellar arms and 2 small insignificant wings at the base. Fig. 39, a, b.
Ecology. Humid broad-leaved forests. 14001600 m. Fl.? Very rare (DD).
Distribution. Vietnam (Lao Cai), Thailand, S China, Taiwan, Malacca Peninsula.
Studied specimens. Chapa, Tonkin, Petelot s.n. (P).
Described from southern Vietnam (Kontum Prov., Dak Gley Distr., ...).
Type (VH 1932) LE (holotype), HN, MO (isotypes).
Terrestrial and lithophytic herb to 25 cm tall with 57 more or less distant
petiolate leaves. Leaves ovate, dark green, sometimes finely undulate along margin, to
8 cm long, 3.5 cm wide. Scape 57 cm long, usually with 1 rudimentary leaf. Rachis
pubescent, 38 cm long, with 814 lax flowers. Floral bracts pubescent, 810 mm
long, not longer than ovary. Ovary 1012 mm long, densely pubescent. Sepals brown
to brown-green, sparsely pubescent; median sepal ovate, acute, 67 mm long; lateral
sepals ovate, oblique, 8 mm long. Petals white, strongly oblique, with broad base and
attenuate apex, 6 mm long, 3 mm wide at the base. Lip white, 1012 mm long, basally
shortly globular, bilobulate, with 23 close glands on each side. Mesochile 6 mm long,
laterally with 45 short fringes. Epichile lobes spreading, ovate, 6 mm long, 4 mm
wide. Column twisted, about 1 mm long, with 1 stigma and 2 small vertical wings at
the base. Anther narrowly-ovate, brightly red, 4 mm long. Fig. 39, f-k; 43, f.
Ecology. Primary humid broad-leaved forests, usually along shady stream
canyons on silicate soils. 10001500 m. Fl. October January. Rare (EN).
Distribution. Vietnam (Dak Nong, Kon Tum).
Studied specimens. Dak Nong, Dak Glong, HLF 5611a (HN, LE).
8. O. tortus King et Pantl., 1896, Journ. As. Soc. Bengal, 2, Nat. Hist. 65:
125. Anoectochilus tortus (King et Pantl.) King et Pantl. 1898, Ann. Roy. Bot.
Gard. (Calcutta) 8: 298, tab. 396. Pristiglottis torta (King et Pantl.) Aver. 1996,
Bot. Journ. (St. Petersburg) 81, 10: 78. A. repens auct. non (Downie) Seidenf. et
Smitinand: Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 11.
Described from northern India (Darjeeling (Duars) Kumai). Type (Pantling
354) CAL (holotype), BM, K (isotypes).
Terrestrial and lithophytic herb to 30 cm tall with 47 distant petiolate leaves.
Leaves ovate, uniform green, 47 cm long. Scape short, 45 cm long, pubescent with
lax many flowered spike 410 cm long. Floral bracts narrowly-ovate, acuminate, finely
pubescent, 812 mm long. Ovary hairy, 812 mm long. Sepals and petals greenish.
Sepals sparsely long hairy; median sepal ovate, acute, 58 mm long; lateral sepals
ovate-oblong, oblique, spreading, 710 mm long. Petals strongly oblique, falcate with
attenuate apex, as long as median sepal. Lip pure white, 1216 mm long, basally
shortly globular, with 2 thick close massive straight conical glands on each side.
Mesochile 57 mm long, laterally with 57 pair long fringes. Epichile lobes spreading,
hemi-circular to broadly-obovate, 47 mm long and wide. Column stout, 2.53.5 mm
tall, with 1 semi-circular stigma, conical rostellar arms and 2 small vertical wings at
the base. Anther narrowly-ovate, beaked, 2.53 mm long. Fig. 39, l-n.
Ecology. Closed humid broad-leaved forests on rocky limestone in shady, wet
places. 400950 m. Fl. June August. Very rare (EN).
Distribution. Vietnam (Ninh Binh). Bhutan, N India, S China, Taiwan.
Studied specimens. Ninh Binh, Cuc Phuong national park, HAL 1670 (HN, LE).
Turczaninowia 2008, 11(1) : 5168 145
9. O. inabai (Hayata) T.P. Lin, 1975, Nat. Orch. Taiwan: 216. Anoectochilus
inabai Hayata, 1914, Icon. Pl. Formos. 4: 102, pl. 14.
Described from Taiwan (Rinkiho, ad 5000 ped. alt.). Type (H. Inaba, Aug.
1911) TI?
On its vegetative features this species resembles O. tortus, but differs in
slightly larger flowers, as well as brownish to brownish-pink sepals and petals, with
irregular pinnate whitish tessellation and pinnate venation. Fig, 39, o; 43, g.
Ecology. Closed humid broad-leaved forests on rocky limestone. 9001000 m.
Fl. June August. Very rare (EN).
Distribution. Vietnam (Hoa Binh), Taiwan.
Studied specimens. Hoa Binh, Mai Chau, HAL 9002 (HN).
Orchipedum Breda, Kuhl et Hasselt, 1827, Gen. Sp. Orch. Asclep.: tab. 10.
Type: O. plantaginifolium Breda
Terrestrial sympodial herbs with creeping succulent rhizome rooting at nodes,
ascending erect leafy floriferous stem and terminal many flowered spike. Leaves
few, spirally arranged, petiolate, sheathing at the base, glabrous, convolute, asymmetric,
uniformly green. Scape leafless, with 24 large sterile bracts. Flowers, sessile,
resupinate. Sepals free. Petals oblique-obovate, joined in apical part and connivent to
median sepal forming hood; lateral sepals spreading, with concave base, enclosing
saccate base of lip. Lip adnate to column base, spurred, 3-lobed. Spur globose, slightly
bifid, curved forwards under the blade, containing dense group of fleshy hairs on each
side. Base of lip with erect rounded side lobes embracing column base, inside with 2
erect semi-circular parallel keels. Median lobe with narrow clawed base suddenly
broadening into spreading lobules. Column narrow, bended, with 1 stigma and narrow
winged rostellar arms; anther narrowly ovate acute. Pollinium sectile, very long, to 5
mm long, with stipe and viscidium.
3(1) species. Vietnam, Malacca Peninsular, Java, Sumatra, Philippines.
irregularly grooved on sides, narrowing toward epichile. Epichile lobes spreading, flat,
rectangular to obovate, 1.53 mm long, 1.52 mm wide, irregularly serrate along margin.
Column simple, narrow, 910 mm long, bent forward at the middle, with narrowly-
ovate stigma at front. Anther and pollinarium 6 mm long. Fig. 40, a-g; 43, h.
Ecology. Coniferous, mixed and broad-leaved evergreen forests on shale and
sandstone, commonly along shady streams. 10001100 m. Fl. April May. Very rare (EN).
Distribution. Vietnam (Thua Thien Hue).
Note. Species is known up to now only by alone type collection.
Key to species
1. Flowers more than 5 mm across; lip 912 mm long; sepals dark purple-brown .....................
................................................................................................................................ 1. R. petelotii
Flowers less than 5 mm across; lip less than 5 mm long; sepals pale brown or greenish
.................................................................................................................................................. 2
2. Epichile entire, broadly-ovate to circular; ovaries erect, densely adpressed to the
rachis ......................................................................................................................... 2. R. tokioi
Epichile 2-lobed, with spreading acute lobes; ovaries spreading, not adpressed to the
rachis ................................................................................................................ 3. R. lanceolata
1. R. petelotii (Gagnep.) Ormerod, 1998, Austral. Orchid Rev. 63: 11; Aver.
et Averyanova, 2003, Updated Checklist Orch. Viet.: 11. Haemaria petelotii Gagnep.
1933, Bull. Soc. Bot. France, 80: 349. Anoectochilus petelotii (Gagnep.) Seidenf.
1975, Contr. Revis. Orch. Fl. Cambodia. Laos Vietnam: 7. Ludisia discolor auct.
non (Ker Gawl.) A. Rich.: Seidenf. 1992, Orch. Indochina: 29, p. p.
Described from northern Vietnam (Tonkin: Cha-pa, 1.5001.600 m.). Type
(Petelot 5166) P (holotype), C (isotype).
148 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Terrestrial and lithophytic herb 2035 cm tall with 35 petiolate leaves in basal
rosette. Leaves ovate, 47 cm long, acute velvety dark green, purple-brown to nearly
black, with 1 (rarely 3) white to light pink longitudinal stripes. Scape 712 cm tall,
shortly pubescent, with 23 cuneate, acuminate, yellowish bracts. Rachis 615 cm
long, with many not resupinate flowers. Ovary glabrous, 1015 mm long. Floral bracts
sparsely pubescent at the base, ciliate along margin, narrowly-ovate, acuminate, as
long as, or slightly shorter than ovary. Sepals purple-brown, free, sparsely pubescent
at the base; median sepal narrowly ovate, acute, 79 mm long, with recurved apex;
lateral sepals oblique-ovate, narrowing to the apex, 810 mm long. Petals brown,
narrowly-ovate, oblique, falcate, acuminate, as long as median sepal. Lip 910 mm
long. Hypochile purple to purple-brown, saccate, 56 mm long, 45 mm wide, with 2
roundish, erect side lobes, 2 large stalked hemi-circular gland at the base, 1 short tall
triangular lamellate keel along midrib and lamellate, finely serrulate lateral longitudinal
flanges outside. Epichile joined to hypochile with a short claw, sometimes twisted on
180. Epichile lobes circular, spreading, irregular wavy, 56 mm across. Column conical,
45 mm tall, with 2 large broad, triangular wavy wings at front, acute bifid rostellum
and 2 lateral stigmas touching at front. Anther acuminate at the apex. Fruits ellipsoid
capsule, 1215 mm long, 35.5 mm wide. Fig. 40, h-l; 43, i.
Ecology. Humid forests on rich soils, mainly on rocky limestone. 7002000 m.
Fl. July August (VU).
Distribution. Vietnam (Bac Kan, Cao Bang, Ha Giang, Hoa Binh, Kon Tum,
Lai Chau, Lam Dong, Lao Cai, Quang Binh, Son La, Thanh Hoa, Thua Thien Hue).
Studied specimens. Bac Kan, Ba Be national park, HLF 514 (HN, LE), HLF 552
(HN, LE); Bac Kan, Cho Don, HLF 761 (HN, LE), HLF 804 (HN), HAL 4790 (HN, LE); Bac Kan,
Na Ri, HAL 4924 (HN); Cao Bang, Bao Lac, CBL 312 (HN, LE); Cao Bang, Tra Linh, CBL 983
(HN, LE); Cao Bang, Trung Khanh, HAL 5561 (HN, LE); Ha Giang, Bac Me, HAL 6577 (HN,
LE); Ha Giang, Meo Vac, NTH 3442 (HN, LE); Ha Giang, Quan Ba, NTH 3603 (HN, LE); Ha
Giang, Yen Minh, NTH 3459 (HN, LE), NTH 3459 (HN, LE); Hoa Binh, Da Bac, HAL 467/a (HN,
LE); Hoa Binh, Mai Chau, HAL 641 (HN, LE), HAL 690 (HN, LE), DKH 7716 (H, LE, MO), DKH
7921 (HN, LE, MO); Kon Tum, Dak Gley, VH 1850 (HN, LE); Kon Tum, Kon Plong, VH 5439
(HN); Lai Chau, Sin Ho, HAL 9962 (HN), HAL 10050 (HN); Lai Chau, Tam Duong, HAL 10254
(HN); Lam Dong, Lac Duong, Hon Giao ridge, VH 4132 (HN, LE); Lao Cai, Van Ban, HAL 2013
(HN, LE), HAL 2316 (HN), HAL 2656 (HN, LE); Quang Binh, Bo Trach, HAL 6059 (HN, LE),
HAL 6141 (HN, LE photo); Son La, Moc Chau, DKH 7322 (HN, LE, MO), DKH 7357 (HN, LE,
MO), DKH 7452a (HN, LE, MO), DKH 7453 (HN, LE, MO); Son La, Moc Chau, HAL 9304
(HN); Son La, Thuan Chau, HAL 9632 (HN); Son La, Yen Chau, DKH 7204 (HN, LE, MO), HAL
9435 (HN); Thanh Hoa, Ba Thuoc, HAL 3495 (HN, LE); Thua Thien Hue, A Luoi, HAL 7682
(HN), HAL 7683 (HN, LE); Thua Thien Hue, Bach Ma national park, HLF 943 (HN, LE), HLF
1002 (HN, LE).
Note. Taxonomically very isolated species, which differs in touching stigma
lobes, serrulate lamellate flanges of mesochile, large alone median keel on the lip disc
and usually twisted claw. The plant is not rare in limestone areas of northern Vietnam
and may be certainly found in allied limestone regions of Laos and China.
Turczaninowia 2008, 11(1) : 5168 149
Fig. 40. Orchipedum echinatum: a flowering plant, b flower, c flattened tepals, d lip, side
view and lip longitudinal section, e spur cross section and spur glands, f column, side and
frontal views, g pollinarium; Rhomboda petelotii: h fruiting plant, i flattened tepals and lip,
j lip, without side lobe, k column, side view and longitudinal section of lip base, l column,
frontal view; R. tokioi: m flattened flower, without column, n column, side and frontal views.
Turczaninowia 2008, 11(1) : 5168 151
Vrydagzynea Blume, 1858, Coll. Orch. Arch. Ind. Jap.: 71. Vridagzenia
Benth. 1881, Journ. Linn. Soc. London (Bot.) 18: 344, orth. var. rej.
Lectotype: V. albida (Blume) Blume (Etaeria albida Blume)
Terrestrial and lithophytic sympodial herbs with creeping succulent rhizome
rooting at nodes, ascending erect leafy floriferous stem and terminal few flowered,
rather dense spike. Leaves few, spirally arranged, petiolate, sheathing at the base,
more or less distant or crowded at the stem apex, glabrous, convolute, asymmetric,
green to purple, sometimes with median whitish stripe, acute. Scape short, leafless,
with 12 sterile bracts. Flowers small, sessile, resupinate, not widely opening. Sepals
free, very fleshy, succulent. Petals connivent to median sepal forming hood. Lip simple,
ovate, fleshy, conical, with spur. Spur saccate to cylindric, with 2 glands at the base.
Column short, erect, with short bifid rostellum, wingless or with 2 insignificant wings
at front. Anther ovate, obtuse. Pollinia 2, sectile, with caudicle and viscidium. Stigma
with 2 separated lobes.
35(1) species. Tropical and subtropical Asia, northeastern Australia to islands
of W Pacific.
V. formosana Hayata, 1916, Icon. Pl. Formos. 6: 88. V. albida auct. non
(Blume) Blume, 1858: Seidenf. 1992, Opera Bot. 114: 30; Aver. 1994, Ident. Guide
Vietnam. Orch.: 34; Aver. et Averyanova, 2003, Updated Checkl. Orch. Vietnam: 61.
Described from Taiwan (Inter Urai et Agioku ). Type (leg. D. Hayata,
Mai. 1916) TI?
Herb 420 cm tall. Leaves 46, petiolate, ovate, 24 cm long, uniform green,
acute. Scape sparsely pubescent, ebracteate or with 13 acuminate bracts, 25 mm
long. Rachis densely tomentose, 13 cm long, elongate in fruits up to 12 cm, with 320
close flowers. Floral bracts narrowly-ovate, acuminate, 610 mm long. Ovary 58
mm long. Flowers hardly opening, 34 mm long, white, later yellowish, base of sepals
and spur green to olive-green, later dull orange to red. Sepals sub-similar, narrowly-
ovate, very fleshy, 3.55 mm long, 1.52 mm wide, blunt. Petals lanceolate, of the
same length, connivent to dorsal sepal. Lip 33.5 mm long, spurred, fleshy and finely
papillose along midvein, attenuate to short fleshy blunt tip, lateral sides thin incurved;
lip blade being flattened broadly ovate to almost orbicular. Spur broadly-conic, 3.54
mm long, with 2 stalked glands. Fruit erect elliptic capsule, 810 mm long, 3 mm wide.
Fig. 41, b; 44, b.
Ecology. Humid forests on any kind of soils, commonly along wet, shady
stream and river valleys. 3001100 m. Fl. January April. Not rare (LR).
152 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Distribution. Vietnam (Ha Tinh, Hoa Binh, Lao Cai, Nghe An, Ninh Binh,
Quang Binh, Quang Tri, Thua Thien Hue, Tuyen Quang), SE China, Taiwan, Laos.
Studied specimens. Ha Nam Ninh, Cuc Phuong, sine date, Bong 146 (HN), LX-VN
1767 (LE); Ha Tinh, Huong Son, HAL 5215a (HN, LE); Hoa Binh, Da Bac, HAL 440 (HN, LE);
Lao Cai, Van Ban, DKH 6678a (HN, LE, MO), HAL 2748 (HN, LE); Nghe An, Tuong Duong,
HLF 6738 (HN), HLF 6930 (HN); Quang Binh, Bo Trach, HAL 6058 (HN, LE), HAL 6148 (HN,
LE), HAL 6229 (HN, LE); Quang Binh, Quang Ninh, HAL 6316 (HN, LE) d-EXSICCATES OF
VIETNAMESE FLORA 0005/HAL6316; Quang Tri, Da Krong, HLF 6192 (HN); Thua Thien
Hue, Nam Dong, HAL 11129 (HN), HAL 6896a (HN, LE); Tuyen Quang, Na Hang, HAL 221
(HN, LE).
Note. Species closely related to V. albida (Blume) Blume, from which it
differs in hairy rachis, floral bracts and ovary, as well as almost rounded to emarginate
lip (when flattened) with very thin incurved lateral margins. Fairly common species in
northern Vietnam, particularly in wet forests on rocky limestone. Easily overlooked in
botanical studies and collecting due to dwarf habit and unattractive appearance of
flowers. All seen collections from Vietnam belong to this species, which replaces
V. albida (distributed in Myanmar, Thailand, Peninsular Malaysia, Indonesia, Philippines
and New Guinea) in Eastern Indochina, mainland SE China and in Taiwan.
Zeuxine Lindl. 1826, Collect. Bot. App. N 18 (sub Zeuxina orth. var.
rej.); id. 1840, Gen. Sp. Orch. Pl.: 485, nom. cons.
Type: Z. strateumatica (L.) Schleichter (Pterygodium sulcatum Roxb.;
Z. sulcata (Roxb.) Lindl.).
Terrestrial and lithophytic sympodial herbs with creeping succulent rhizome
rooting at nodes, ascending erect leafy floriferous stem and terminal few- to many
flowered spike. Leaves few, spirally arranged, petiolate, sheathing at the base, glabrous,
convolute, asymmetric, uniformly green or with median whitish stripe, acute, rarely
conduplicate, grass-like. Scape leafless, with 24 sterile bracts. Flowers small, sessile,
resupinate. Sepals free. Petals connivent to median sepal forming hood, lateral sepals
spreading. Lip spurless. Hypochile concave to saccate, with various papillae on lateral
sides. Epichile reflexed, circular-obovate or 2-lobed, connected to hypochile by a short
claw. Column short, erect, with 2 prominent rostellar arms and 2 wings at front. Anther
narrowly triangular-ovate, acute. Pollinia 2, sectile, with caudicle and viscidium. Stigma
with 2, small, convex, separated lobes.
70(7) species. Africa, tropical and subtropical Asia, Indonesia, Philippines,
Australia, New Guinea, Pacific Islands to Samoa.
Key to species
1. Leaves linear, conduplicate, grass-like ................................................... 1. Z. strateumatica
Leaves lanceolate to almost circular, plicate .................................................................... 2
2. Epichile entire, circular or obcordate ....................................................... 2. Z. goodyeroides
Epichile distinctly 2-lobed ................................................................................................ 3
3. Epichile with divergent lobes 68 mm long; sepals 78 mm long ................. 3. Z. grandis
Epichile with spreading lobes less than 3 mm long; sepals less than 6 mm long ........ 4
Turczaninowia 2008, 11(1) : 5168 153
4. Column with large, protruding quadrate or triangular flat vertical wings on front;
petals very oblique, suddenly narrowing toward the obtuse apex ................ 4. Z. nervosa
Column with insignificant, hardly visible wings on front; petals not much oblique
narrowing to the apex ......................................................................................................... 5
5. Lip white, 34 mm long; epichile lobes rectangular, about twice as long as wide ...........
.............................................................................................................................. 5. Z. parvifolia
Lip yellow, 23 mm long; epichile lobes squarish to obovate, less than twice as long
as wide ........................................................................................................................ 6. Z. flava
1. Z. strateumatica (L.) Schleichter, 1911, Bot. Jahrb. 45: 394; Seidenf. 1992,
Opera Bot. 114: 37; Aver. 1994, Identif. Guide Vietnam. Orch.: 38; Aver. et Averyanova,
2003, Updated Checklist Orch. Viet.: 61. Orchis strateumtica L. 1753, Sp. Pl.:
943. Z. bonii Gagnep. 1931, Bull. Mus. Natl. Hist. Nat. 2. ser. 3: 326. Z. evrardii
Gagnep. 1931, l.c.: 326.
Described from Sri Lanka (in Zeylon). Type (Hermann s.n.) BM.
Stem erect, slender, to 20 cm tall, leafy. Leaves erect, conduplicate, linear,
grass-like, acute, to 6 cm long. Inflorescence many-flowered cylindric dense spike 1
4 cm long. Ovary glabrous, 57 mm long. Floral bracts triangular-ovate, acuminate,
reddish-yellow, 410 mm long. Sepals white, sub similar, ovate, obtuse, 34 mm long,
glabrous. Petals of the same length, white, oblique narrowly-obovate, obtuse. Lip shortly
saccate, with 2 glands, 33.5 mm long; claw short, fleshy, papillose and grooved.
Epichile small, transversely oblong-reniform, papillose, blunt, yellow-green. Column
short, 1.52 mm tall. Fig. 41, c-f.
Ecology. Wet secondary grasslands, open wet places along stream and river
valleys. 501500 m. Fl. January February. Very rare (DD).
Distribution. Vietnam (Cao Bang, Ha Noi, Ha Tinh, Lam Dong, Nam Dinh,
Nghe An, Ninh Binh, Thanh Hoa). Tropical and subtropical Asia, Japan, Taiwan,
Philippines, Indonesia to New Guinea.
Studied specimens. Cao Bang; Ha Noi, Kien-Khe, mont Den; Ha Son Binh, Rocher
de Notre Dame, riviere Noire; Lam Dong, Dalat, Camly; Lang-bian; Nghe Tinh; Thanh Hoa;
Tonkin, delta du Fleuve Rouge; Tonkin, Mong-ho, Fleuve Rouge.
Note. Very variable and widespread species escaped and naturalized in Florida
and Cuba. Recently it, probably, become much rarer as did not been collected in
Vietnam during last 50 years.
3. Z. grandis Seidenf. 1978, Dansk Bot. Ark. 32: 90. Z. affinis auct. non
(Lindl.) Benth. ex Hook. f.: Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 61.
Described from Thailand (Omkoi Road at km 13). Type (GT 7730) C.
Terrestrial and lithophytic herb. Stem 825 cm tall with 26 leaves, usually
withered at the time of flowering. Leaves petiolate; narrowly-ovate, 1.54 cm long,
uniform dull green, acute. Scape and rachis densely pubescent, rachis to 6 cm long,
with 110 lax flowers. Floral bracts triangular-ovate, acuminate, pubescent, 612 mm
long. Ovary pubescent, 59 mm long. Sepals olive-green to light pinkish, pubescent
outside, sub-similar, ovate, 78 mm long, concave, obtuse. Petals of the same length
or longer, white, oblique narrowly-obovate to oblong-spathulate, roundish at the apex.
Lip white, with light yellow-brownish central spot on epichile, 1013 mm long. Hypochile
small, saccate, with 1 small gland on each lateral side. Epichile 2lobed, lobes flat, at
an acute angle to the axis, convergent, oblique-obovate, each 68 mm long, 3.5
4.5 mm wide. Column short, without frontal wings. Fruit erect elliptic capsule, 8
12 mm long, 23 mm wide. Fig. 41, l, m; 44, d.
Ecology. Wet primary and secondary closed forests, often with bamboo, on shale,
usually along rocky stream slopes. 10001200 m. Fl. February March. Rare (EN).
Distribution. Vietnam (Quang Tri), Thailand, Hainan.
Studied specimens. Quang Tri, Huong Hoa, Sa Mui pass, HLF 5755a (HN, LE),
HLF 6278 (HN, LE); d-EXSICCATES OF VIETNAMESE FLORA 0069/HLF6278.
Terrestrial herb. Stem 1025 cm tall with 35 leaves. Leaves petiolate; petiole
and sheath 0.51.5 cm long; leaf blade ovate to narrowly-ovate, 26 cm long, pure dull
green, often with wide median whitish stripe, acute. Scape and rachis sparsely pubescent,
rachis 16 cm long, with 314 laxy flowers. Floral bracts triangular-ovate, acuminate,
510 mm long. Ovary sparsely hairy, 510 mm long. Sepals dull greenish to greenish-
brown, sub-similar, ovate, 3.55.5 mm long, concave, obtuse. Petals of the same length,
white, oblique, broadly lunate, attenuate at the apex. Lip white, with 2 dull green to
brown spots on epichile, 4.56 mm long. Hypochile saccate, with (1)23 close parallel
fat calluses on each lateral side. Epichile 2-lobed, lobes spreading, rectangular to obovate.
Column short, with 2 large rectangular to triangular acute wings at front. Fruit erect
elliptic capsule, 912 mm long, 3.54.5 mm wide. Fig. 41, n, o; 44, e.
Ecology. Wet primary and secondary closed forests, often with bamboo, on
any soils. 2001500 m. Fl. January April. Not rare (LR).
Distribution. Vietnam (probably all provinces), Bhutan, Nepal, India, S China,
Thailand, Ryukyu Islands, Taiwan, Philippines.
Studied specimens. Cuc Phuong national park, NTH 3890 (HN, LE); Ha Giang,
Hoang Su Phi, HAL 6619 (HN, LE); Ha Giang, Quan Ba, DKH 5008 (HN, LE, MO); Ha Nam
Ninh, Cuc Phuong national park, LX-VN 1810 (HN, LE); Ha Tinh, Huong Son, HAL 5215a
(HN, LE); Hoa Binh, Da Bac, HAL 393 (HN, LE); Lao Cai, Van Ban, HAL 2001 (HN, LE); Nghe
An, Quy Chau, HLF 3095 (HN, LE); Nghe An, Tuong Duong, HLF 6602 (HN, LE); Quang
Binh, Minh Hoa, HAL 5868 (HN, LE); Son La, Moc Chau, DKH 7357 (HN, LE, MO); Son La,
Yen Chau, DKH 7225 (HN, LE, MO); Tuyen Quang, Na Hang, HAL 181 (HN, LE).
Note. Widespread and common species in Vietnam, which easily survives in
wet secondary forests, particularly with bamboo.
5. Z. parvifolia (Ridl.) Seidenf. 1978, Dansk Bot. Ark. 32, 2: 82; Seidenf.
1992, Opera Bot. 114: 37; Aver. 1994, Identif. Guide Vietnam. Orch.: 37; Aver. et
Averyanova, 2003, Updated Checklist Orch. Viet.: 61. Hetaeria parvifolia Ridl.
1903, Journ. Straits Branch Roy. As. Soc. 39: 87. Z. tonkinensis Gagnep. 1931,
Bull. Mus. Natl. Hist. Nat. 2. s. 3, 7: 328.
Described from Malacca Peninsula (Penins. Mal. sec. Ind. Kew.). Type
(Ridley s.n.) K.
Terrestrial herb. Stem 1030 cm tall with 35 leaves. Leaves shortly petiolate,
ovate to narrowly-ovate, 1.54.5 cm long, uniform dull green, or sometimes with median
whitish stripe, acute. Scape and rachis sparsely pubescent, rachis 18 cm long, few-
to many flowered. Floral bracts triangular-ovate, acuminate, sparsely pubescent to
glabrous, 510 mm long. Ovary sparsely pubescent or glabrous, 48 mm long. Sepals
light greenish-brown, sub-similar, ovate to narrowly-ovate, 34.5 mm long, concave,
obtuse. Petals of the same length, white, oblique-oblong, roundish at the apex. Lip 3
4 mm long, white, with yellowish to yellowish-brown spot on epichile. Hypochile saccate,
with 1 small gland on each lateral side. Epichile 2-lobed, lobes spreading, rectangular,
about twice as long as wide. Column short, with 2 narrow insignificant wings at front.
Fig. 41, p; 44, f.
156 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Fig. 41. Rhomboda lanceolata: a flower and floral bract; Vrydagzynea formosana: b flattened
flower and longitudinal section of spur; Zeuxine strateumatica: c flowering plant, d flower, e
flattened sepals and petals, f column and lip, side view; Z. goodyeroides: g flowering plant,
h flattened tepals and lip, i column, frontal fiew, j anther, ventral view, k ovary and floral bract;
Z. grandis: l flattened tepals and lip, m column, frontal view; Z. nervosa: n flattened tepals
and lip, o column, side and frontal views; Z. parvifolia: p column, flattened tepals and lip.
Turczaninowia 2008, 11(1) : 5168 157
Ecology. Wet primary and secondary closed forests, often with bamboo, on
any soils. 3001800 m. Fl. December April. Not rare (LR).
Distribution. Vietnam (probably all provinces), Myanmar, Thailand, Laos,
Cambodia, Malacca Peninsula, Ryukyu Islands, Taiwan, Philippines.
Studied specimens. Bavi mt., Tonkin, Balansa 2002 (P); Dak Nong, Dak Glong,
HLF 5503a (HN, LE); Gia Lai Kontum, Chu Pa, LX-VN 2531 (HN, LE); Ha Giang, Hoang Su
Phi, HAL 6620 (HN, LE); Hoa Binh prov., Da Bac, HAL 556 (HN, LE); Kon Tum, Dak Gley, VH
2281 (HN, LE); Lam Dong, Dalat, VH 2523 (HN, LE); Lam Dong, Lac Duong, Bi Dup mt, VH
3236 (HN, LE); Lang Son, Bac Son, HAL 6790 (HN, LE); Lao Cai, Van Ban, HAL 2265 (HN, LE),
DKH 6740 (HN, LE, MO); Thanh Hoa, Ba Thuoc, HAL 3310a (HN, LE); Thua Thien Hue,
Nam Dong, HAL 6896b (HN, LE).
Note. Widespread and common species in Vietnam, which easily survives in
wet secondary forests, particularly with bamboo.
6. Z. flava (Wall. ex Lindl.) Trimen, 1885, Syst. Cat. Fl. Pl. Ceylon: 90.
Monochilus flavus Wall. ex Lindl. 1840, Gen. Sp. Orch. Pl.: 487.
Described from Nepal. Type (Wallich, Wall. Cat. 7380A) K.
Terrestrial and lithophytic herb. Stem 1545 cm tall with 58 leaves. Leaves
petiolate; ovate to narrowly-ovate, 48 cm long, uniform light green, acute. Scape and
rachis pubescent, rachis to 22 cm long, many flowered. Floral bracts triangular-ovate,
acuminate, sparsely pubescent, 511 mm long. Ovary sparsely pubescent to sub-
glabrous, 610 mm long. Flowers odorless. Sepals dull greenish-brown, sub-similar,
ovate to narrowly-ovate, 3.54 mm long, concave, obtuse, sparsely pubescent to glabrous,
lateral sepals shorter. Petals of the same length, light greenish, oblique narrowly-obovate
to oblong-lanceolate, obtuse. Lip yellow, 23 mm long. Hypochile saccate, with 1
small gland on each lateral side. Epichile 2-lobed, lobes spreading, oblong-obovate,
0.81.2 mm long. Column short, with 2 narrow, fat, insignificant wings at front. Fruit
erect elliptic capsule, 69 mm long, 23 mm wide. Fig. 42, a, b; 44, g.
Ecology. Wet primary and secondary closed forests, often with bamboo, on
any soils. 5001500 m. Fl. February April. Rare (VU).
Distribution. Vietnam (Ninh Thuan, Quang Tri, Son La), Bhutan, Nepal, N
India, Myanmar, Thailand.
Studied specimens. Ninh Thuan, Ninh Hai, Nui Chua mt, HLF 4445 (HN, LE),
HLF 4538 (HN, LE); Quang Tri, Huong Hoa, Sa Mui pass, HLF 5939 (HN, LE); Son La, Moc
Chau, DKH 7515 (HN, LE, MO).
Zeuxinella Aver. in Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 96.
Type: Z. vietnamica (Aver.) Aver. (Zeuxine vietnamica Aver.)
Terrestrial creeping sympodial herb with succulent rhizome rooting at nodes,
short ascending leafy stem and erect thin terminal leafless inflorescence 1018 cm
tall. Leaves 35, petiolate. Petiole and sheath 0.51 cm long. Leaf blade velvety
carbon-black above, pale dull purple below, broadly ovate to orbicular, 1.53 cm long,
shortly apiculate. Scape hairy, with 34 acuminate bracts; rachis pubescent, 14 cm
158 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
long, with 38 laxy sessile flowers. Floral bracts pubescent, cuneate, acuminate, 46
by 11.5 mm. Ovary densely glandular hairy, 57 mm long. Flowers hardly opening,
odorless, resupinate, 35.5 mm long. Sepals dull greenish to greenish-brown, sub-
similar, narrowly-ovate to ovate, 35 mm long, concave, obtuse. Petals white, narrowly
ovate, incurved, 35 mm long, connivent to dorsal sepal forming hood. Lip yellow,
spurred, fleshy, 3-lobed, 3 mm long. Hypochile boat-shaped; sidelobes narrowly ovate,
rising and incurved. Epichile reflexed, 3-lobulate, with rounded-triangular subequal
lobules, with 4 cushions on the disk. Spur straight, sac-like, 2.53 by 2 mm, with 1
cylindric gland on each lateral side. Column small, with 2 insignificant frontal wings, 2
narrow rostellar arms and 2 stigmas. Fig. 42, c-h; 44, h.
Monotypic genus endemic to northern Vietnam.
Spiranthes L.C. Rich. 1817, Orch. Europ. Annot.: 20, nom. cons.
Type: S. spiralis (L.) K. Koch (Ophrys spiralis L.), type cons.
Terrestrial herbs with few fleshy fasciculate root tuberoids, rosulate sessile
conduplicate leaves and erect inflorescence bearing terminal many-flowered, secund,
spirally twisted, dense spike. Flowers sessile, resupinate, not widely opening. Sepals
free, sub-similar; median sepal erect, lateral sepals spreading, oblique at the base.
Petals connivent to sepal forming hood. Lip sessile or shortly claved, entire or lobed,
undulate along margin, enveloping column at the base. Column short, erect, with bifid
rostellum and 1 convex, entire stigma at front; clinandrium with prominent margins.
Anther erect, dorsal. Pollinia 2, mealy, with caudicle and viscidium.
Turczaninowia 2008, 11(1) : 5168 159
Fig. 42. Zeuxine flava: a flattened tepals and lip, b column, frontal view; Zeuxinella
vietnamica: c flowering plant, d ovary, floral bract and flower, e flattened tepals and lip,
f lip, side view, g gland inside spur, h column, view from above (with removed operculum)
and frontal view; Spiranthes sinensis: i flowering plants, j flower, k flattened tepals and
lip, l flattened median sepal and petals, m flattened lip, n ovary, column and lip, side view,
o column, side view, p column, view from above and frontal view, q fruit, r floral bract.
160 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
40(1) species. Tropical, subtropical and temperate areas of the world except
tropical and southern Africa.
1. S. sinensis (Pers.) Ames, 1908, Orch. 2: 53; Seidenf. 1992, Opera Bot.
114: 25; Aver. 1994, Identif. Guide Vietnam. Orch.: 48; Aver. et Averyanova, 2003,
Updated Checklist Orch. Viet.: 56. Neottia sinensis Pers. 1807, Syn. Pl. 2: 511.
Aristotelea spiralis Lour. 1790, Fl. Cochinchin.: 522. N. australis R. Br. 1810,
Prodr.: 319. N. amoena M. Bieb. 1819, Fl. Taur.-Caucas. 3: 606. S. australis
(R. Br.) Lindl. 1824, Bot. Reg. 10: tab. 823. S. amoena (M. Bieb.) Spreng. 1826, Syst.
Veg. 3: 708. S. spiralis (Lour.) Makino, 1926, Journ. Jap. Bot. 3: 25, nom. illeg.
Described from southern China (prope Gantonem Sinarum). Type (Loureiro
s.n.) P.
Herb 840 cm tall with 310 linear-lanceolate to broadly-lanceolate, glossy
green, acute leaves, 212 cm long, 38 mm wide. Scape glandular hairy, with 24
bracts. Rachis 414 cm. Floral bracts lanceolate, acuminate, glandular hairy, 47 mm
long. Ovary 24 mm long. Flowers pink-purple to white. Sepals and petals sub-similar,
narrowly ovate to lanceolate, 34 mm long, 11.5 mm wide, obtuse, petals slightly
smaller. Lip simple, oblong, longitudinally concave, 3.54.5 mm long, 23 mm wide,
plicate along margin, with 2 large spherical glands at the base. Column 23 mm tall,
with 2 erect, acute rostellar arms. Fruit ovate, glandular pubescent capsule, 67 mm
long, 3 mm broad. Fig. 42, i-r; 44, i.
Ecology. Secondary short-tall grasslands, pastures, road-cuts, old rice field in
agricultural areas, open grassy swamps along stream and river valleys, usually on wet
alluvial soils. 52000 m. Fl. February April. Very common (LR).
Distribution. Vietnam (probably all provinces). Widespread in temperate,
subtropical and tropical Asia to Australia, New Zeeland, Tasmania and south-west
Pacific Islands.
Studied specimens. Annam, sine loc., Eberhardt s.n. (P); Bac Kan, Ngan Son,
CBL 1258 (HN, LE); Cao Bang, Bao Lac, CBL 1373 (HN, LE); Cao Bang, Nguyen Binh, CBL
131 (HN, LE); Cao Bang, Tien Ha, sine coll. (P); Cochimchina s.n., Loureiro (P); Dalat, Boden,
Kloss s.n. (BM); Ha Giang, Bac Me, HAL 6607 (HN, LE); Ha Giang, Dong Van, CBL 1772 (HN,
LE); Ha Giang, Quang Ba, DKH 4832 (HN, LE); Ha Giang, Yen Minh, CBL 1979 (HN); Hanoi,
Bat Bac, Balansa; Hanoi, Kien Khe, Bon; Hoa Binh, Da Bac, HAL 285 (HN); Hoa Binh, Da Bac,
HAL 554 (HN); Ja Lao Mahay, Pierre s.n. (P); Kon Tum, Kon Plong, DKH 4750 (HN, LE);
Kontum, Kon Plong, Mang La, VH 5456 (HN); Kontum, Ngoc Linh mt., VH 10 (HN, LE); Lac
Duong, Langbian, sine coll.; Lang Son, Bac Son, HAL 6791 (HN, LE); Langbian, Andre; Lao
Cai, Van Ban, HAL 2448 (HN); Lao Cai, Van Ban, HAL 2845 (HN, LE); Ninh Binh, Cuc Phuong
national park, 20 Feb. 1971, Quynh 41, CPNP 5120 (HN); Pagode de Bamly, Tonkin, Petelot;
Quang Yen, dAlleizette; Santay, Balansa; Son La, Moc Chau, DKH 7449 (HN); Thai Nguyen,
Eberhardt; Tonkin delta, Eberhard drawing, 218; Tonkin, sine loc., Bon; Tu Phap, Balansa.
Note. Very common, variable and widespread weedy species, often spreading
into secondary habitats on agricultural lands. Used in traditional oriental medicine.
Turczaninowia 2008, 11(1) : 5168 161
Fig. 43. Ludisia discolor: a (HLF 6570), b (HAL 6350), c (HAL 1340, HAL 1655 and HAL 5865);
Odontochilus saprophyticus: d (HAL 8194); O. elwesii: e (HAL 1720); O. umbrosus: f (HLF
5611a); O. inabai: g (HAL 9002); Orchipedum echinatum: h (HLF1296, type); Rhomboda
petelotii: i (NTH 3459).
162 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
Fig. 44. Rhomboda tokioi: a (LX-VN s.n., 1985); Vrydagzynea formosana: b (HAL 6148); Zeu-
xine goodyeroides: c (HAL 10672); Z. grandis: d (HLF 6278); Z. nervosa: e (Averyanov, HAL
s.n., 2004); Z. parvifolia: f (HAL 6620). Z. flava: g (HLF 5939); Zeuxinella vietnamica: h (HLF
638); Spiranthes sinensis: i (HAL 11131a).
Turczaninowia 2008, 11(1) : 5168 163
LITERATURE
Averyanov L.V. Introduction and Fam. Orchidaceae Juss. Vascular plants synopsis of
Vietnamese flora, 1. Leningrad. Nauka, 1990. 167 p.
Averyanov L.V. Identification guide to Vietnamese orchids (Orchidaceae Juss.). S.-Pe-
tersburg: World and Family, 1994. 432 p.
Averyanov L.V., A.L. Averyanova. Updated checklist of the orchids of Vietnam. Hanoi.
Vietnam National University Publishing House, 2003. 102 p.
Dressler Robert L. Phylogeny and classification of the Orchid Family. Portland, Oregon.
Dioscorides Press, 1993. 314 p.
Gagnepain F., A. Guillaumin. Orchidacees, Apostasiacees. In Lecomte, H. (ed). Flore
Generale de lIndo-chine 6. Paris, 1934. P. 142654.
Nguyen Thien Tich. The orchids of Viet Nam. Part 1. Ho Chi Minh. Nha Xuat Ban Nong
Nghiep, 2001. 424 p.
Nguyen Tien Ban, L.V. Averyanov, Duong Duc Huen. 248. Orchidaceae Juss. 1789. The
orchids. P. 512666. Conspectus of vascular plants of Viet Nam. Vol. 3. Ha Noi. Agriculture
Publishing House, 2005. 1248 p.
Pham-hoang Ho. Cay Co Mien Nam. Saigon, 1960. 630 p.
Pham-hoang Ho. Cay Co Mien Nam. 2 ed. Vol. 2. Saigon. Nam Quoc-Te Phat-Trien Sach,
1972. 1139 p.
Pham-hoang Ho. An Illustrated Flora of Vietnam. Vol. 3. Part 2. - Santa Ana. Mekong Printing,
1993. 1176 p.
Pham-hoang Ho. An Illustrated Flora of Vietnam. Vol. 3. Part 2. Ho Chi Minh. Nha Xuat
Ban Tre, 2000. 1020 p.
Seidenfaden G. The Orchids of Indochina // Opera Botanica, 1992. Vol. 114. 502 p.
Tran Hop. The orchids of Vietnam. Ho Chi Minh. Nha Xuat Ban Nong Nghiep, 1998. 703 p.
(Orchidaceae) . -
, ,
,
Apostasioideae, Cypripe-
dioideae Spiranthoideae, 21 85 .
, ,
, , ,
. -
. 4 -
. Neuwiedia zollingeri var. annamensis
(Gagnep.) Aver., N. zollingeri var. balansae (Gagnep.) Aver., Paphiopedilum barbigerum var.
aspersum (Aver.) Aver. P. callosum var. warnerianum (T. Moore) P.J. Cribb ex Aver.
164 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.
CONTENTS
PREFACE ........................................................................................................ 5
ACKNOWLEDGEMENTS ............................................................................... 35