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582.594 (597)

L. Averyanov ..

THE ORCHIDS OF VIETNAM

ILLUSTRATED SURVEY

Part 1

SUBFAMILIES APOSTASIOIDEAE, CYPRIPEDIOIDEAE

AND SPIRANTHOIDEAE

The article opens serial publication of illustrated critical taxonomical survey of

orchids in the flora of Vietnam. The first part of this monograph includes introduction,

illustrated glossary of terminology used in identification keys and in description of

taxa, key for identification of genera, as well as taxonomical treatment of three

subfamilies Apostasioideae, Cypripedioideae and Spiranthoideae with 21 genera

and 85 species. Valid name, necessary synonyms, type material citation, short

description, data on ecology and distribution, as well as list of studied voucher specimens

for each species are provided. Mentioned species and varieties are illustrated withline drawings and color photographs.

PREFACE

Basic primary knowledge on orchid taxonomy and diversity in Eastern Indochina

and in Vietnam was obtained from landmark surveys (Gagnepain, Guillaumin, 1943;

Pham-hoang, 1960, 1972, 1993, 2000; Averyanov, 1990, 1994; Seidenfaden, 1992) based

mainly on collections of French and post-war periods housed now in largest part in

Herbaria of Paris (P) and Saint Petersburg (LE). Mentioned publications were used

in some compilative reviews few later (Tran Hop, 1998; Nguyen Thien Tich, 2001;

Nguyen Tien Ban, Averyanov, Duong Duc Huen, 2005). According to most recentassessment (Averyanov, Averyanova, 2003) flora of Vietnam includes about 158 genera

and 900 discovered orchid species, which may be estimated as about 80% of all orchids

occurring on the territory of the country. Meanwhile, recent exploration of numerous

earlier inaccessible territories of the country brings last time many more new

discoveries, which essentially expand our knowledge about regional plant diversity.

Illustrated taxonomical survey of all species hitherto reported for Vietnam including

most recent discoveries based on collected voucher herbarium specimens is the main

goal of presented monograph.

Monograph treatment will successively follow family system of R. Dressler

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Subtrib. 5.2.2. Vanillinae Lindl.

Vanilla Sw.

Subtrib. 5.2.3. Lecanorchidinae Dressler

Lecanorchis Blume

Trib. 5.3. Gastrodieae Lindl.

Subtrib. 5.3.1. Gastrodiinae Lindl.

Didymoplexiella Garay, Didymoplexiopsis Seidenf., Didymoplexis Griff.,

Gastrodia R. Br., Yoania Maxim.

Subtrib. 5.3.2. Epipogiinae Schlecht.

Epipogium R. Br., Stereosandra Blume

Trib. 5.4. Nervilieae Dressler

Nervilia Gaudich.

Epidendroid group

Trib. 5.5. Arethuseae Lindl.

Subtrib. 5.5.1. Arundinae DresslerArundina BlumeSubtrib. 5.5.2. Bletiinae Benth.

Acanthephippium Blume,Anthogonium Lindl.,Bletia Ruiz et Pavon, CalantheR. Br., Cephalantheropsis Guillaum., Eriodes Seidenf., Hancockia Rolfe,Mischobulbon Schlecht.,Nephelaphyllum Blume,Pachystoma Blume,Phaius Lour.,

Plocoglottis Blume, Spathoglottis Blume, Tainia BlumeTrib. 5.6. Malaxideae Lindl.

Subtrib. 5.6.1. Malaxidinae Benth. et Hook. f.

Liparis L.C. Rich., Malaxis Sw.

Subtrib. 5.6.2. Oberoniinae Aver.

Hippeophyllum Schlecht., Oberonia Lindl.

Trib. 5.7. Coelogyneae Pfitz.

Subtrib. 5.7.1. Thuniinae Schlecht.Thunia Reichenb.Subtrib. 5.7.2. Coelogyninae Bentham

Coelogyne Lindl., Neogyna Reichenb. f., Otochilus Lindl., Panisea Steud.,Pholidota Hook., Pleione D. Don

Trib. 5.8. Epidendreae Humb., Bonpl. et Kunth

Subtrib. 5.8.1. Glomerinae Schlecht.

Agrostophyllum BlumeSubtrib. 5.8.2. Polystachyinae Pfitz.

Polystachya Hook.

Dendrobioid group

Trib. 5.9. Podochileae Pfitz.

Subtrib. 5.9.1. Eriinae Benth.

Callostylis Blume, Ceratostylis Blume, Cryptochilus Wall., Eria Lindl.,Porpax Lindl., Trichotosia Blume

Subtrib. 5.9.2. Podochilinae Benth. et Hook.

Appendicula Blume, Podochilus Blume

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8 Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Subtrib. 5.9.3. Thelasiinae Schlecht.

Phreatia Lindl., Thelasis Blume

Subtrib. 5.9.4. Dendrobiinae Lindl.

Dendrobium Sw., Epigeneium Gagnep., Flickingeria Hawkes

Subtrib. 5.9.5. Bulbophyllinae Schlecht.

Bulbophyllum Thouars,Hamularia Aver., Monomeria Lindl., Sunipia Smith,Trias Lindl.

Cymbidioid group

Trib. 5.10. Cymbidieae Pfitz.

Subtrib. 5.10.1. Bromheadiinae Dressler

Bromheadia Lindl.

Subtrib. 5.10.2. Eulophiinae Benth.Eulophia R. Br., Dipodium R. Br., Geodorum Jackson

Subtrib. 5.10.3. Collabiinae Schlecht.

Chrysoglossum Blume, Collabium Blume, Diglyphosa Blume

Subtrib. 5.10.4. Thecostelinae Schlecht.

Thecopus Seidenf., Thecostele Reichenb.

Subtrib. 5.10.5. Cymbidiinae Benth.

Cremastra Lindl., Cymbidium Sw.

Subtrib. 5.10.6. Acriopsidinae Dressler

Acriopsis Blume

Subfam. 6. VandoideaeTrib. 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.,HygrochilusPfitz.,Kingidium P.F. Hunt,Luisia Gaudich., Malleola J.J. Smith, Micropera Lindl.,Microsaccus Blume, Ornithochilus Lindl., Papilionanthe Schlecht., ParapterocerasAver.,Pelatantheria Ridl.,Pennilabium J.J. Smith,Phalaenopsis Blume,PomatocalpaBreda, 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, TuberolabiumYamamoto, Uncifera Lindl., Vanda Jones, Vandopsis Pfitz.

Illustrated survey of three subfamilies Apostasioideae, Cypripedioideae

and Spiranthoideae is presented here in form of standard taxonomic treatment, which

includes identification keys for all mentioned taxa and their short characterization.

Correct name (with standard taxonomic reference), type, data about volume and

distribution is reported for each taxonomic group. Data for each genus also include

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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. NationalGeographic 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 accordingto 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,

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VA - Vietnam-American Series collections of international group with Dr.

N.T. Hiep as principal investigator.

VH Vietnamese Highlands collections on program of U.S.A. National

Geographic Society Flora of Highlands of South Vietnam (years 1993-2001, # 5094-

93, 5803-96, 6383-98) with principal investigator Prof. Leonid V. Averyanov.

WP Vietnam-Chinese expedition of 2003 year with Prof. Phan Ke Loc as

co-leader from Vietnamese Side.

Modern administrative divisions of Vietnam into provinces and city territories,

as well as list of their names, which are used in indication of species distribution within

Vietnam, are presented on map 1.

Illustrated glossary of orchid terminology used in identification and plant

descriptions are presented below in the next chapter.

ILLUSTRATED GLOSSARY OF THE ORCHID

IDENTIFICATION TERMINOLOGY

Additional explanations of terms are available on cited figures (Fig. 112).

Abaxial the side away from the axis, normally the lower surface.

Acaulescent becoming stemless.

1 Dien Bien

2 Lai Chau

3 Lao Cai

4 Yen Bai

5 Son La

6 Ha Giang

7 Cao Bang

8 Tuyen Quang

9 Bac Can

10 Phu Tho

11 Thai Nguyen

12 Lang Son13 Bac Giang

14 Quang Ninh

15 Vinh Phuc

16 Ha Noi city

17 Bac Ninh

18 Ha Tay

19 Hung Yen

20 Hai Duong

21 Hai Phong

22 Hoa Binh

Names of provinces, names of city areas and figures designating these units on the map:

23 Ha Nam

24 Thai Binh

25 Ninh Binh

26 Nam Dinh

27 Thanh Hoa

28 Nghe An

29 Ha Tinh

30 Quang Binh

31 Quang Tri

32 Thua Thien-Hue

33 Da Nang city

34 Quang Nam35 Kon Tum

36 Quang Ngai

37 Gia Lai

38 Binh Dinh

39 Dak Lak

40 Phu Yen

41 Dak Nong

42 Khanh Hoa

43 Binh Phuoc

44 Lam Dong

45 Ninh Thuan

46 Tay Ninh

47 Binh Duong

48 Dong Nai

49 Binh Thuan

50 Long An

51 Ho Chi Min city

52 Ba Ria-Vung Tau

53 An Giang

54 Dong Thap

55 Tien Giang

56 Kien Giang57 Can Tho

58 Vinh Long

59 Ben Tre

60 Hau Giang

61 Tra Vinh

62 Ca Mau

63 Bac Lieu

64 Soc Trang

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Map 1. Modern administrative division of Vietnam territory (for 2007 year) (except Hoang Saand Truong Sa Archipelagos).

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Acaulous stemless or nearly so.

Achlorophyllous without chlorophyll, as in plants or plant structures lacking

green color.

Actinomorphic radially symmetrical.

Aculeate prickly; covered with prickles.

Acuminate having a gradually tapering point. Fig. 12.

Acute distinctly and sharply pointed, but not drawn out. Fig. 12.

Adaxial the side toward the axis, normally the upper surface.

Adnate united or joined together.

Adpressed (appressed) lying flat for the whole length of the organ.

Amplexicaul clasping the stem.

Androclinium see clinandrium.

Androecium all of the stamens in a flower.

Anther the expanded, apical, pollen bearing portion of the stamen. Fig. 47, 10.

Anther cup (operculum) highly modified, easily removable lid-like anther

wall, which covers pollinia placed on the top of column in many representatives of

Dendrobioideae and Vandoideae subfamilies. Fig. 810.

Antrorse directed forward or upward.

Aphyllous without leaves.

Apiculate with a short, sharp, but not stiff, point. Fig. 12.

Appressed see adpressed.

Approximate drawn close together, but not united.

Aristate awned, sharply terminated by a hard, straight point. Fig. 12.Articulate jointed. Fig. 3.

Articulation point of attachment. Fig. 3.

Attenuate tapering gradually to a narrow base. Fig. 12.

Auricle 1 a small lateral outgrowth on the top of column in representatives

of subfamily Orchidoideae. Fig. 7.

Auricle 2 a small, ear-shaped appendage, commonly at the base of lip or

leaf. Fig. 12.

Auriculate with a small ear-like lobe. Fig. 12.

Axial placentation placentation when ovules are attached to the central

axis of an ovary with two or more locules. Fig. 2, a.

Axil the point at the angle between a leaf and a stem.Axillary borne in the axil.

Barkepiphytes (trunk epiphytes) epiphytes growing on branches of middle

size in medium zone of tree canopies. Fig. 1, f.

Berry a fleshy, many-seeded fruit composed of more than one carpel.

Bicuspidate having 2 sharp points.

Bidentate with two teeth. Fig. 12.

Bifid divided into 2 shallow segments, usually at the apex. Fig. 12.

Bifurcate forked.

Bipartite divided nearly to the base into two portions.

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Fig. 1. Ecological groups of orchids and their habitats.Orchids in the flora of Vietnam may be terrestrial plants (a, about 20%), achlorophyllous

obligate mycotrophic (saprophytic) herbs (b, about 2%), lithophytes (c, about 8%), lianas orvines (d, about 2%), humus epiphytes (e, about 15%), bark epiphytes (f, about 48%) and branch epiphytes (g, about 5%).

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Fig. 2. Basic orchid morphology: a, b transversal section of ovary with axial placentation in

Apostasia odorata (a) and parietal placentation inDendrobium 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 inPholidota rubra (h) and hysteranthous (i) inflorescence in

C. fimbriata; j, k types of leaf development (transversal section of developing leaves) with

convolute leaves inLiparis 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).

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Bract more or less reduced leaf-like organ bearing a flower, inflorescence

or partial inflorescence in its axil. Fig. 3.

Branch epiphytes (canopy epiphytes) epiphytes growing on small branchlets

and perennial leaves in peripheral zone of tree canopies. Fig. 1,g.

Canopy epiphytes branch epiphytes.

Broadly elliptic spindle-shaped, two-dimensional; with an index from 1 to

about 1.4. Fig. 11.

Broadly lanceolate lance-shaped, narrow, tapering toward the apex more

than toward the base; with an index from 3 to around 4. Fig. 11.

Broadly oblanceolate inversely broadly lanceolate. Fig. 11.

Broadly obovate inversely broadly ovate. Fig. 11.

Broadly ovate egg-shaped in outline, broader at the base, two-dimensional;

with an index less than 1.4. Fig. 11.

Broadly triangular three-angled, with an index less than 0.85. Fig. 11.

Bursicle a cup-like covering over the viscidium in some representatives of

subtribe Orchidinae. Fig. 10.

Caducous falling off early, like floral bracts in some species ofCalanthe or

Coelogyne.

Caespitose tufted.

Callus (pl. calli) a thickening or protuberance, commonly on the lip. Fig. 6.

Calyculus a small cup or circle of bract-like structures outside of the sepals

in species ofLecanorchis. Fig. 8.

Campanulate bell-shaped.Canaliculate channeled, with a longitudinal groove.

Canopy epiphytes see branch epiphytes. Fig. 1, g.

Capilletium the sterile kapok-like hairs within the fruit mellowing seed

masses, particularly in epiphytes. Fig. 9.

Capitate head-like.

Capsule a dry, many-seeded, dehiscent fruit composed of more than one

carpel. Fig. 9.

Carinate keeled.

Carpel highly modified megasporophyll, formed from one modified leaf

bearing ovules (megasporangia) and seeds; in the orchids, the three carpels are so

united and modified as to be nearly unrecognizable. Fig. 2, a, b.Caudate tailed; with a tail-like appendage. Fig. 12.

Caudicle a slender elastic extension of the pollinium, or a mealy portion at one

end of the pollinium; the structure is a part of the pollen, which is formed within the

anther. Fig. 6, 7, 10.

Central relating to the central part of a body.

Ciliate having fine hairs at the margin, refers usually to the margin of leaf or

tepal. Fig. 12.

Clavate club-shaped, pointed at one end and rounded at the other.

Claw the conspicuously narrowed and attenuate base of an organ.

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Fig. 3. Structure of orchid shoot.

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Cleistogamous refers to flowers which regularly self-pollinate without

opening.

Clinandrium (androclinium) the anther bed, that part of the column under,

or surrounding, the anther. Fig. 9.

Column more or less modified structure formed by union of filaments and

style. Fig. 46, 810.

Column wing a wing- or arm-like appendage of the column, usually lateral.

Columnfoot a ventral extension of the base of the column which has the lip

attached at its tip. Fig. 8.

Complanate flattened or compressed.

Conduplicate leaf or leaf-like organs, with a single median fold, with each

half being flat. Fig. 2, m.

Congeneric belong to the same genus.

Connate united.

Connective the sterile portion of the filament connecting the two pollen

sacs of an anther. Fig. 7, 10.

Connivent coming into close contact or converging, but not united.

Conspecific belonging to the same species.

Convolute rolled up longitudinally with parts in an overlapping arrangement.

Fig. 2,j.

Cordate heart-shaped with the base at the broad, notched end. Fig. 11, 12.

Coriaceous leathery.

Corolla the inner whorl of the perianth, petals.Corymb flat-topped inflorescence.

Crenate scalloped, toothed with crenations. Fig. 11.

Crenulate crenate, but the teeth small. Fig. 11.

Cristate crested.

Cucullate hooded, hood-shaped.

Cuneate wedge-shaped. Fig. 12.

Cuspidate tipped with a sharp, rigid point. Fig. 12.

Decurved curved or bent down.

Deflexed bent outwards.

Dentate toothed. Fig. 11.

Denticulate minutely toothed. Fig. 11.Determinate a habit of growth in which each unit has a limited growth,

usually in sympodial shoot.

Disc the face of any flat organ with special reference to the lip.

Distal away from the base, toward the apex.

Distichous having leaves or other organs in two opposite rows.

Dorsal refers to the back or outward surface of an organ in relation to the

axis; in orchids refers to the upper side of the flower; see abaxial.

Dorsal sepal see median sepal.

Dorsiventral an organ which has more or less distinct dorsal and ventral

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Fig. 5. Orchid morphology (subfamily Cypripedioideae).

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Fig. 6. Orchid morphology (subfamily Spiranthoideae).

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surfaces, as a non-terete leaf.

Duplicate refers to the folding of leaves during development, folded once

with each half flat. Fig. 2, k.Echinate with numerous stiff hairs or spines.

Elastoviscin a very elastic substance found in pollinia and especially in

caudicles.

Ellipsoid spindle-shaped; narrow and tapering at the ends, three-dimensional.

Elliptic spindle-shaped, two-dimensional; with an index around 1.43. Fig. 11.

Emarginate notched, usually at the apex. Fig. 12.

Endemic confined to a given region.

Ensiform sword-shaped. Fig. 11.

Entire simple and with a smooth edge, as contrasted to toothed or lobed.

Fig. 11.

Ephemer living a very short time (usually annual plants of arid areas).

Ephemeral of flowers, referring to those which last only a few hours.

Ephemeroid perennial plants of arid areas with very short vegetative and

reproductive periods.

Epichile the terminal part of a complex lip when it is distant from the basal

portion. Fig. 6.

Epigeous growing upon the ground.

Epiphyte a plant growing on another plant but not parasitic. Fig. 1, e-g.

Equitant said of conduplicate or laterally flattened leaves or bracts which

overlap each other in two ranks.Erose bitten or gnawed. Fig. 11.

Extrafloral occurring outside of a flower; refers usually to glands that are

found outside of flower buds.

Falcate sickle-shaped. Fig. 11.

Farinose mealy.

Fasciculate clustered or bundled.

Filament the stalk of the stamen which bears the anther. Fig. 4, 5.

Filiform thread-like.

Fimbriate fringed, refers usually to the margin of leaf or tepal. Fig. 11.

Flabellate fan-shaped. Fig. 11.

Flexuose bent alternately in opposite directions.Flora the array of all plant species occurring in a region.

Floral bract more or less reduced leaf-like organ bearing a flower (or

reduced floral bud) in its axil. Fig. 3, 5, 6, 8.

Fovea a small depression or pit, like in lip ofMalaxis species.

Fusiform spindle-shaped.

Geniculate abruptly bent like a knee-joint.

Geophyte plants, in which new annual shoot develops from underground

bud, usually tuberiferous or bulb ephemeroids.

Gibbous swollen or enlarged on one side; ventricose.

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Fig. 7. Orchid morphology (subfamily Orchidoideae).

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Fig. 8. Orchid morphology (subfamilyEpidendroideae).

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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.

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Fig. 9. Orchid morphology (subfamily Vandoideae).

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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 ofDoritis pulcherrima, thatillustrate stipe (tegula) formation; e, f sagittal section of column (e) and pollinarium (f) of

flower ofDactylorhiza incarnata, that illustrate caudicle formation.

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Node the point on a stem at which a leaf or bract is attached. Fig. 3.

Non-resupinate flower the flower with not-resupinate ovary having the lip

turned upwards.

Obcordate inversely cordate. Fig. 11.

Oblanceolate inversely lanceolate. Fig. 11.

Obligate restricted to particular conditions or circumstances, like

achlorophyllous mycotrophic plants. Fig. 1, b.

Oblique with unequal sides; slanting. Fig. 12.

Oblique bifid divided into 2 unequal shallow segments, usually at the apex.

Fig. 12.

Oblong much longer than broad, with nearly parallel sides, rounded at each

end. Fig. 11.

Obovate inversely ovate. Fig. 11.

Obsolete wanting or rudimentary.

Obtuse blunt or rounded at the apex. Fig. 12.

Ombrophilous adapted to shady habitats of humid rainy forest.

Operculum see anther cup. Fig. 8.

Orbicular of a flat body with a circular outline. Fig. 11.

Oval oval-shaped, not much longer than broad, with nearly parallel sides,

rounded at each end. Fig. 11.

Ovary the part of the flower that contains ovules and develops into the fruit.

Fig. 35, 8.

Ovate egg-shaped in outline, broader at the base, two-dimensional; with anindex around 1.41.6. Fig. 11.

Ovoid egg-shaped, three-dimensional.

Palea flat or terete, moveable appendages attached by a filiform base, found

on tepals in some species ofBulbophyllum. Fig. 8.

Pandurate violin- or fiddle-shaped. Fig. 11.

Panicle a much branched inflorescence.

Paniculate branched, branching (of an inflorescence).

Papilla (pl. papillae) a soft, short, superficial gland or protuberance. Fig. 6.

Papillose covered with papillae.

Parietal placentation placentation when ovules are attached to the walls

of the ovary. Fig. 2, b.Pectinate comb-like; with close, regularly spaced teeth or hairs, often in a

single row, like the teeth of a comb.

Pedicel the stalk of a single flower in an inflorescence. Fig. 3, 8.

Peduncle the stalk bearing an inflorescence or solitary flower. Fig. 3, 5, 6.

Peloric a more or less radially symmetrical mutant of a species which

normally has zygomorphic flowers.

Peltate shield-shaped.

Pendent hanging.

Pendulous hanging.

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Perianth (tepals) a collective term for sepals and petals, together.

Petals in orchids two lateral segments of the inner whorl of perianth (corolla).

Fig. 4, 5, 7, 9.

Petiolate leaf leaf with distinct petiole and leaf blade.

Petiole a leaf stalk. Fig. 6.

Pilose soft hairy.

Pistil the female reproductive organ of a flower, typically consisting of a

stigma, style and ovary.

Placenta that portion of the ovary that bears the ovules.

Placentation the arrangement of ovules and configuration of the placentas

in ovary. Fig. 2, a, b.

Plicate pleated or folded leaves having several to many major longitudinal

veins and folded in each one. Fig. 2, l.

Pollen masses see Pollinium.

Pollinarium(pl. pollinaria) the complete set of pollinia with associated parts,

like viscidium (viscidia) and stipe. Fig. 610.

Pollinium (pl. pollinia) a more or less compact mass of pollen. Fig. 610.

Porate pollinium pollinium with a pore or pores.

Porrect directed outward and forward.

Prostrate lying flat on the ground.

Proteranthous an apical inflorescence produced before the pseudobulbs

and leaves on the same shoot. Fig. 2, g.

Proximal the part nearest the axis, toward the base.Pseudobulb a swollen aerial stem. Fig. 3, 8.

Pseudocopulation a special type of pollination, in which flowers mimic

female insects and are pollinated by the males when they attempt to copulate with the

flowers.

Pseudopollen mealy, farinose, pollen-like deposit (originated from highly

modified papillae) on the lip of some species ofEria andDendrobium.

Pubescent softly hairy or downy.

Pulvinate cushion-like.

Pustulate see pustulose.

Pustulose (pustulate) having small blisters or pustules.

Pyriform pear-shaped. Fig. 11.Quaquaversal directed or bending in every direction.

Raceme an unbranched, elongate, indeterminate inflorescence with

pedicellate flowers maturing from the base upwards.

Racemose having flowers in racemes.

Rachis the axis of an inflorescence. Fig. 3, 8, 9.

Recurved curved backwards or downwards.

Reflexed abruptly bent or turned downwards or backwards.

Reniform kidney-shaped. Fig. 11.

Resupinate flower the flower with resupinate ovary having the lip turned down.

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Fig. 11. Main types of shapes and margins of orchid leaf and tepal.

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Fig. 12. Main types of apices and bases of orchid leaf and tepal.

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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 ofCheirostylis.

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.

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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 ofthe 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 ofPaphiopedilum. Fig. 5.

Tegula see stipe.

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Tepal index see index.

Tepals see perianth.

Terete cylindric and usually tapering. Fig. 11.

Terminal the end or apex.

Terrestrial ground dwelling plants, usually with creeping rhizome, ground

pseudobulbs or subterranean tuber (root-stem tuberoids). Fig. 1, a.

Tessellate chequered.

Theca (pl. thecae) a pollen sac or cell of the anther. Fig. 7.

Tomentose densely covered with matted woolly or short hairs.

Triangular three-angled, with an index more than 0.85. Fig. 11.

Trichome a hair or hair-like outgrowth of the epidermis. Fig. 9.

Tridentate three-toothed. Fig. 12.

Truncate ending abruptly, as though broken off. Fig. 12.

Trunk epiphytes see bark epiphytes. Fig. 1, f.

Tuber in orchids see tuberoid.

Tuberoid (root-stem tuberoid) a thickened, more or less modified tuber-like

root with a bud and some anatomical stem structures at the base. Fig. 7.

Umbo a small blunt or rounded protuberance, as on the ventral surface of

staminode in some species ofPaphiopedilum. Fig. 5.

Umbonate having a central, projecting point, like the boss of a shield.

Uncinnate hook-shaped.

Undulate waved.

Unguiculate contracted at the base into a claw. Fig. 12.Urceolate pitcher-, vase- or urn-shaped.

Velamen one or more layers of hygroscopic spongy dead cells on the outside

of a root.

Ventral refers to the front or inward surface of an organ in relation to the

axis; see adaxial.

Ventricose swollen or inflated on one side.

Verrucose warty.

Verticillate whorled.

Villose with long, weak hairs.

Vine see liana.

Viscidiophore an elongated prolongation of the base of theca (anther canal)and rostellum side lobes bearing at the apex viscidium. Fig. 7.

Viscidium(pl. viscidia) a viscid, usually disc-like part of the rostellum joined

to the pollinium, enabling it to adhere to an insect-pollinator body. Fig. 6-10.

Whorl a circle of three or more leaves or leaf-like organs attached at the

same level on a stem.

Zygomorphic bilaterally symmetrical.

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35Turczaninowia 2008, 11(1) : 5168

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 monograph 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.

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San Diego County Orchid Society Conservation Committee. Explo-

ration of endangered calcium dependent Paphiopedilum species in Northern

Vietnam, 19992000 years.

Fauna & Flora International (Vietnam Program). The distribution of

Paphiopedilum vietnamense and its current status in the wild, 2000; Communi-

ty-based Conservation of the Hoang Lien Mountain Ecosystem, Vietnam, Flora

and vegetation survey of Van Ban district, Lao Cai province of northern Viet-

nam, 2002; Preliminary survey of orchids and gymnosperms in Trung Khanh

district, Cao Bang province northern Vietnam, 2004;

Fauna & Flora International Vietnam Conservation Support Program

and the Ministry of Agriculture and Rural Development, Vietnam. Pu Lu-

ong Cuc Phuong Limestone Landscape Conservation Project, Preliminary bota-

nical survey of primary vegetation in Pu Luong nature reserve, 2003.

Fauna & Flora International (Vietnam Program) & Counterpart In-

ternational. Preliminary survey of Orchids (Orchidaceae) in Phong Nha Ke

Bang National Park, 2005.

WWF Indochina Programme. Green Corridor Project in Thua Thien Hue

Province, VN085301. Lowland flora and vegetation. Preliminary survey, 2005.

Basic Research program in Life Sciences of Viet Nam, # 611001.

Threatened conifers and cycads of Vietnam, 20002002.

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 forphotographs used in Fig. 24, c; 31, f; 36, d; 43, g; 44, d and Dr. H. Koopowiz for

photograph used in Fig. 26, i.

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37Turczaninowia 2008, 11(1) : 5168

KEY FOR IDENTIFICATION OF ORCHID GENERA

IN THE FLORA OF VIETNAM

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 ....................................................Paphiopedilum4. 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 slightlyincumbent; 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

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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 .............. ........ ........ .... 1212. 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 ..........................................................................................................................Macodes16. 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

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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, golden or reddish net 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 .............................................................................Rhomboda26 (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 sepalsovate to elliptic, subsimilar; pollinia with caudicles and basal viscidia ............. ........ .. 29

29. One or two stigmas distinctly concave ........................................................................... 30

Stigmas convex or stalked, rarely hardly concave ........................................................3230. 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

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forward from the column base ...............................................................................Habenaria

Stigma-lobes not freely extending in front of column ................................................... 35

35. Thecas at their base extending from column base; rostellum forming a broad band

above the coherent, hardly prominent stigma-lobes .............................................Pecteilis

Neither stigma-lobes nor thecas extending in front of column base ............................ 36

36. Lip ligulate, undivided, at the base with long narrow spur not dilated toward apex;

stigma sessile, with a single hardly concave surface ................................. ......Platanthera

Lip usually 3-lobed; spur if present short saccate or globular; stigma-lobes separate

or hardly coherent .............................................................................................................. 37

37. Lip without spur; 2 large stelidia rise above anther on lateral sides of the column .........

....................................................................................................................................Herminium

Lip with short, but distinct spur at the base; stelidia small insignificant, sometimes

hardly visible ..............................................................................................................Peristylus38 (7). Creeping epiphytic or lithophytic vines with fleshy succulent green stem to 15 m

long; pollinia very soft and mealy ................................................................................ Vanilla

Stem more or less short, not liana-like; plants sometimes with creeping plagiotropic

woody rhizome up to 23 m long; pollinia mealy to hard ................ .............................. 39

39. Leaves not articulate, circular, broadly cordate or fan-shaped usually appear after

flowering; plants with globular underground tuber; pollinia soft, sectile ........... ..Nervilia

Leaves usually articulate, elliptic, ovate and lanceolate to linear; flowering plants

commonly have leaves, rarely plants develop flowers before the leaves appear; plants

rarely have underground tubers; pollinia mealy to hard, sometimes with caudicles ........ 40

40. Pollinia rather hard, 48, with or without caudicles; plants commonly epiphytic with

or without pseudobulbs .................................................................................................... 41

Pollinia rather soft, 8, rarely 4, with distinct, but sometimes quite small caudicles;plants commonly terrestrial, often with short vertical stem, plicate leaves and lateral

inflorescence ........................................................................................................................ 43

41. Column not distinctly hooded or winged at the apex; plants commonly with

pseudobulbs of several internodes; inflorescence terminal or lateral; pollinia 4-8 with

or without caudicles ........................................................................................................... 42

Column hooded or winged at the apex around anther; plants with pseudobulbs of 1

internode; inflorescence terminal (sometimes superficially appearing as lateral); pollinia

4, hard, with small caudicles ............................................................................................. 59

42. Flowers with a short or no columnfoot; leaves conduplicate or plicate ...................... 64

Flowers with a prominent columnfoot; leaves always conduplicate ........................... 67

43 (40). Plants with tall erect stem bearing normal numerous distichous leaves ..................... 44

Stem creeping, ascending, sometime pseudobulbous, short and flesh, with few sub-

radical leaves ....................................................................................................................... 48

44. Inflorescence terminal ........................................................................................................ 45

Inflorescence distinctly lateral .......................................................................................... 46

45. Flowers white, 812 cm across, lip entire or hardly 3-lobed, white with yellow-orange

keels; flower bracts large, thin, deciduous, 34 cm long; leaves deciduous, thin, with

blue-green waxy surface; lithophytic or epiphytic plant .......................................... Thunia

Sepals and petals white, with pink-purple tint, flowers 38 cm across; lip distinctly 3-

lobed, pink to purple, commonly with yellow center; flower bracts small, insignificant

persistent, usually less than 5 mm long; leaves persistent, rigid, not waxy; terrestrial

or lithophytic plant ....................................................................................................Arundina

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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,

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coriaceous, not deciduous; inflorescence usually few to many-flowered; lip not

convolute around the column ............................................................................... Coelogyne

62 (59). Flowers large, tepals about 4 cm long; sepals with saccate, concave base; lip funnel

shaped, side lobes large, folded around the column ..............................................Neogyna

Flowers small or of medium size, tepals less than 2 cm long; sepals at the base more o

less flat; lip side lobes small, not folded around the column ....... .......... ....................... 63

63. Column long, narrow, dilated at the apex; lip distinctly divided into a deeply concave

hypochile with two high lateral lobes and a flat entire epichile, strongly reflexed

down; epichile and hypochile separated with transversal bilobed band; floral bracts

deciduous .................................................................................................................. Otochilus

Column short, thick, not distinctly dilated toward the apex; lip not distinctly divided

into hypochile and epichile; lip have no transversal bands; floral bracts deciduous or

persistent ...................................................................................................................Pholidota64 (42). Leaves distichous, laterally compressed, fleshy ............ ................................. .............. 65

Leaves dorsiventral, flat and thin .................................................................................... 66

65. Leafy stems close together .......................................................................................Oberonia

Leafy stems spaced on creeping rhizome .....................................................Hippeophyllum

66. Column long, slender, commonly longer than broad; flowers resupinate, very rare not

resupinate, lip turned down, very rare upwards, narrowing to the base, without auricles

...........................................................................................................................................Liparis

Column short, thick, commonly as long as broad; flowers always not resupinate, lip

turned upwards, with broad auriculate base ............................................................. Malaxis

67 (42). Pollinia 4 ............................................................................................................................... 68

Pollinia 6 or 8 ...................................................................................................................... 76

68. Shoots clustering into dense tufts, stems slender, not pseudobulbous, commonlyless than 10 cm tall, with many close distichous leaves usually less than 1 cm long;

inflorescence terminal, less than 1 cm long, few-flowered with small campanulate

flowers 23 mm across; pollinia clavate ..............................................................Podochilus

Shoots commonly not clustering into dense tufts, stems commonly swollen (at least

in some internodes), pseudobulbous, rare slender, reed-like, normally taller than 10 cm;

or short pseudobulbous, consist of one swollen fleshy internode, distant on creeping

woody rhizome; leaves rare less than 1 cm long; inflorescence lateral or sub-terminal

commonly longer than 1 cm, 1- to many-flowered; flowers campanulate to well-opening,

commonly larger than 3 mm across; pollinia not clavate ............................................... 69

69. Pollinia without stipes ........................................................................................................ 70

Pollinia with stipes .............................................................................................................. 74

70. The non rhizomatous part of sympodial shoots consisting of several to many

internodes, usually fleshy, pseudobulbous (rare thin, reed-like), clustering on short

rhizome .................................................................................................................................. 71

The non rhizomatous part of sympodial shoots fleshy, pseudobulbous (sometimes

small indistinct), consisting of 12 internodes, commonly distant on more or less long

creeping rhizome .................................................................................................................. 72

71. Basal part of non rhizomatous sympodial shoot thin, woody, rod-like, often branching

with fleshy apical internodes forming 1-leaved pseudobulbs; leaves not sheathing at

the base; inflorescence 1(2)-flowered; flowers lasting few morning hours ............Flickingeria

The non rhizomatous sympodial shoot not distinctly differentiated into thin and

fleshy portions, or have fleshy base of few swollen internodes, rarely branching,

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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 columnapex; 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; stemnot 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

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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 denselyhaired 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

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96. Raceme nodding down ............................................................................................Geodorum

Raceme normally straight and erect ................................................................................ 97

97. Flowers newer open widely; tepals narrowly lanceolate, free; column narrow, long

and straight without columnfoot; anther with 4 pollinia .....................................Cremastra

Flowers normally widely opening; sepals and petals broadly-lanceolate to broadly-

ovate; lateral sepals often connate at the base forming small or large spur-like menthum;

column with short, but distinct columnfoot; anther with 2 pollinia cleft at the base ......

....................................................................................................................................... Eulophia

98 (85). Anther with 4 more or less equal globular pollinia free from each other ....... ............ ... 99

Anther with 2, sometimes more or less divided pollinia .............................................. 103

99. Very small, normally leafless epiphytes with flat ribbon-like green roots; stem usually

13 mm tall; flowers lasting few hours ........................................................... Taeniophyllum

Plants with green leaves; stem much taller than 3 mm; flowers lasting few days ..... 100100. Small canopy epiphytes; numerous leaves less than 2 cm long; inflorescence very

short, 24 mm long; flowers less than 5 mm across, subsessile .......... ......... .......... .... 101

Large terrestrial, lithophytic or epiphytic plants; leaves few, much longer than 2 cm;

inflorescence 1030 cm long; flowers more than 5 mm across .................................... 102

101. Leaves laterally compressed, equitant, closely placed along stem; stems 48(12) cm

tall; inflorescence of 2 white flowers .................................................................. Microsaccus

Leaves dorsiventral, sometimes fleshy, succulent, more or less distant; stems 10

30 cm long; inflorescence of 14 green flowers ..................................................Adenoncos

102. Terrestrial or lithophytic rosulate plants; stem 25 cm long; all leaves radical, close

each other; inflorescence erect, stout; flowers 1.52.5 cm across ...........................Doritis

Epiphytic plants with long pendent stem to 1 m long; leaves distant on 36 cm;

inflorescence erect to pendulous, slender; flowers 46.5 cm across ........ ........Esmeralda103 (98). Anther with 2 pollen-masses each completely divided in more or less unequal semi-

globular free halves ............................................................................................................ 104

Pollen-masses more integrated ......................................................................................... 128

104. Column without columnfoot ............................................................................................. 105

Column with columnfoot, sometimes small but always quite distinct ....................... 12

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