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FOC | Family List | FOC Vol. 25

1. Orchidaceae

兰科 lan ke

Authors: Xinqi Chen, Zhongjian Liu, Guanghua Zhu, Kai-yung Lang, Zhanhe Ji, Yi-Bo Luo, Xiaohua Jin, Phillip J. Cribb, Jeffrey J. Wood, Stephan W. Gale, Paul Ormerod, Jaap J. Vermeulen, Howard P. Wood, Dudley Clayton & Alexandra Bell

Hemipilia flabellata

Credit: Harvard University Herbaria

Perennial, but sometimes short-lived, terrestrial, epiphytic, or lithophytic, autotrophic or rarely mycotrophic herbs (or rarely scrambling vines), with rhizomes, tubers, or rootstocks with mycorrhizal fungi in roots. Stems either sympodial or monopodial, usually leafy, but leaves sometimes reduced to bractlike scales, 1 or more internodes at base often swollen to form a "pseudobulb"; epiphytic species with aerial, photosynthesizing adventitious roots, often bearing 1 or more layers of dead cells (velamen). Leaves 1 to many, alternate or occasionally opposite, often distichous, sometimes terete or canaliculate, glabrous or very rarely hairy, frequently fleshy or leathery, base almost always sheathing, sometimes articulated, sometimes forming a false petiole, margin entire, apex often emarginate. Inflorescence basal, lateral, or terminal, erect to pendulous, racemose, spicate, subumbellate, or paniculate, 1- to many flowered, flowers rarely secund or distichously arranged. Flowers small to large, often quite showy, usually zygomorphic, very rarely ± actinomorphic, bisexual [very rarely monoecious and polymorphic], sessile or pedicellate, most often resupinate with pedicel and ovary twisted through 180°, occasionally not twisted or twisted through 360°. Ovary inferior, 1-locular, placentation parietal (or rarely 3-locular and placentation axile). Sepals usually free but sometimes variously adnate, median (dorsal) one often dissimilar to laterals, laterals sometimes adnate to a column foot to form a saccate, conic, or spurlike mentum. Petals free or rarely partly adnate to sepals, similar to sepals or not, often showy; lip entire, variously lobed or 2- or 3-partite, ornamented or not with calli, ridges, hair cushions, or crests, with or without a basal spur or nectary, margins entire to laciniate. Column short to long, with or without a basal foot, occasionally winged or with lobes or arms at apex or ventrally; anther mostly 1, less often 2 or 3, terminal or ventral on column, caplike or opening by longitudinal slits; pollen usually forming distinct pollinia, less often loose, pollinia 2, 4, 6, or 8, mealy, waxy, or horny, sectile or not, sessile or attached by stalks (caudicles or stipes) to 1 or 2 sticky viscidia; stigma 3-lobed, mid-lobe often modified to form a rostellum, other lobes either sunken on ventral surface of column behind anther or with 2 lobes porrect. Fruit a capsule, rarely berrylike, usually opening laterally by 3 or 6 slits. Seeds very numerous, dustlike, lacking endosperm, rarely winged.

About 800 genera and ca. 25,000 species (some estimates as high as 30,000 species): worldwide, except for Antarctica, most numerous in the humid tropics and subtropics; 194 genera (11 endemic, one introduced) and 1,388 species (491 endemic, one introduced) in five subfamilies in China.

Recent analyses of orchids incorporating data from DNA analyses have confirmed many aspects of the established classifications but have also provided some surprises for orchid taxonomists. First of all, the results have upheld the monophyly (evolutionary integrity, i.e., the group includes all the taxa derived from an ancestral species) of the orchid family, including the apostasioids and cypripedioids. They also suggest strongly that the orchids are an ancient group that evolved in the great southern continent of Gondwanaland before it split up to form the southern continents of Australia, Africa, and South America, the island of Madagascar, and the subcontinent of India. The subfamilies Apostasioideae, Cypripedioideae, and Orchidoideae (sensu Dressler, Phylogeny Classific. Orchid Fam. 1993) are all monophyletic. However, recent work clearly shows that Vanilla and its relatives form a separate and ancient clade (an evolutionary lineage including all the taxa derived from a single ancestral one) that deserves recognition as the subfamily Vanilloideae, that the Spiranthoideae nest within a more broadly defined Orchidoideae, and that Vandoideae are a specialized clade within a more broadly defined Epidendroideae.

A detailed new classification of the orchid family is currently being produced under the title Genera Orchidacearum, of which four of the six volumes have been published and a fifth is near completion (Pridgeon et al., Gen. Orchid. 1-4(1). 1999-2005). Even when this work is completed, such is the speed with which new information and techniques are being developed and published, it will almost certainly require revision. However, we now have the broad bones of a more robust and predictive classification of the family that will be more satisfactory than the presently widely used systems that are based mainly upon morphological characters.

The classification of the family is currently the subject of some debate, particularly the circumscription and the placement of certain tribes, subtribes, and genera. The classification of Chase et al. (in Dixon et al., Orchid Conservation, 69-89. 2003), elaborated in Pridgeon et al. (loc. cit.), which is strongly supported by recent molecular, embryological, and morphological analyses, is followed here. They recognize five subfamilies: Apostasioideae, Cypripedioideae, Vanilloideae, Orchidoideae, and Epidendroideae.

Lang Kaiyong, Chen Singchi, Luo Yibo & Zhu Guanghua. 1999. Orchidaceae (1). In: Lang Kaiyong, ed., Fl. Reipubl. Popularis Sin. 17: 1-499; Chen Singchi, Tsi Zhanhuo, Lang Kaiyong & Zhu Guanghua. 1999. Orchidaceae (2). In: Chen Singchi, ed., Fl. Reipubl. Popularis Sin. 18: 1-412; Tsi Zhanhuo, Chen Singchi, Luo Yibo & Zhu Guanghua. 1999. Orchidaceae (3). In: Tsi Zhanhuo, ed., Fl. Reipubl. Popularis Sin. 19: 1-437.

Key 5: Subfam. Epidendroideae p.p.: sympodial taxa

1 Lateral sepals connate into a synsepal; inflorescence branched.   92 Acriopsis (p. 280)
+ Lateral sepals free or forming a tube with dorsal sepal, if connate into a synsepal; inflorescence unbranched   (2)
2 (1) Pollinia 2   (3)
+ Pollinia 4-8   (8)
3 (2) Stems each with a single leaf   (4)
+ Stems each with 2 to many leaves   (6)
4 (3) Lip clawed at base; mentum spurlike, cylindric, 4-6 mm.   104 Collabium (p. 311)
+ Lip not clawed at base; mentum not as above   (5)
5 (4) Lip 3-lobed; mentum distinct, conic, ca. 2 mm.   105 Chrysoglossum (p. 313)
+ Lip unlobed; mentum indistinct.   106 Diglyphosa (p. 314)
6 (3) Lip not saccate or spurred at base; leaves with neither long petiole nor pseudostem at base; if mycotrophic, without column foot.   91 Cymbidium (p. 260)
+ Lip saccate or spurred at base; leaves long petiolate; petioles often forming a pseudostem; if mycotrophic, column foot conspicuous   (7)
7 (6) Inflorescence erect; anther cap with 2 dark projections; lip conspicuously 3-lobed.   89 Eulophia (p. 253)
+ Inflorescence nodding; anther cap without projections as above; lip often unlobed or inconspicuously 3-lobed.   90 Geodorum (p. 258)
8 (2) Pollinia 8   (9)
+ Pollinia 4-6   (48)
9 (8) Inflorescence globose, densely flowered; sepals 4-5 mm.   134 Agrostophyllum (p. 362)
+ Inflorescence racemose or reduced to a single flower; sepals (8-)10-70 mm   (10)
10 (9) Inflorescences and often stems, leaves, and leaf sheaths ± covered with reddish brown, or rarely white, hispid hairs; leaves never fleshy and subterete.   128 Trichotosia (p. 357)
+ Inflorescences, stems, and leaves glabrous or rarely bearing white or black hairs; leaves sometimes fleshy and subterete   (11)
11 (10) Pollinia connected by a common stipe to viscidium   (12)
+ Pollinia often sessile, directly attached to viscidium or sticky substance (sometimes viscidium and sticky substance absent), rarely each pollinium with a separate stipe   (13)
12 (11) Column foot absent; mentum absent; anther beaked at apex.   137 Thelasis (p. 365)
+ Column foot conspicuous; mentum present; anther obtuse at apex.   138 Phreatia (p. 366)
13 (11) Column without a conspicuous foot   (14)
+ Column with a conspicuous foot   (26)
14 (13) Pseudobulbs petiole-like, slender, 1.5-2.5 mm in diam   (15)
+ Pseudobulbs absent or much thicker, never petiole-like   (16)
15 (14) Inflorescence racemose, with several flowers; flowers not resupinate; lip superior, base shortly spurred.   93 Nephelaphyllum (p. 280)
+ Inflorescence reduced to a single flower; flower resupinate; lip inferior, spurless.   96 Hancockia (p. 286)
16 (14) Pseudobulbs subglobose to ovoid-globose, rarely ovoid-conic, with 1-5 terminal leaves; viscidium triangular.   98 Spathoglottis (p. 287)
+ Pseudobulbs cylindric to conic, very rarely subglobose, sometimes absent or replaced by long stems, with several to many basal or lateral leaves; viscidium absent or if present then not triangular   (17)
17 (16) Stems 1-leaved; leaves terete or dorsiventrally flattened; lip mid-lobe swollen, bulbous.   132 Ceratostylis (p. 360)
+ Stems few to many leaved; leaves and lip without above combination of characters   (18)
18 (17) Stems short, entirely enclosed by imbricate leaf sheaths; inflorescence a densely flowered raceme with small bracts.   138 Phreatia (p. 366)
+ Stems elongate, leafy throughout entire length   (19)
19 (18) Inflorescence axillary, few flowered, glabrous.   123 Cylindrolobus (p. 349)
+ Inflorescence terminal or subterminal, few to densely many flowered, glabrous to densely stellate-hairy   (20)
20 (19) Inflorescence densely covered with short stellate hairs, usually densely many flowered; lip callus bipartite, farinose, with a powdery median ridge that usually ends in a protruding globose apical callus.   122 Mycaranthes (p. 348)
+ Inflorescence glabrous or nearly so, few to many flowered; lip without above combination of characters   (21)
21 (20) Stem terete, with more than 10 leaves along lower to upper part   (22)
+ Stem, if present, with 2-6(-8) leaves above middle   (23)
22 (21) Leaves not deciduous after anthesis; lip spurless.   107 Arundina (p. 314)
+ Leaves deciduous after anthesis; lip spurred at base.   108 Thunia (p. 315)
23 (21) Stigma usually subterminal; lip neither spurred nor saccate at base, vesiculate on mid-lobe or disk.   99 Cephalantheropsis (p. 288)
+ Stigma lateral; lip often spurred or saccate, very rarely spurless, not vesiculate on mid-lobe or disk   (24)
24 (23) Pseudobulbs subglobose to ovoid-globose, rarely ovoid-conic, with 1-5 terminal leaves; viscidium triangular.   98 Spathoglottis (p. 287)
+ Pseudobulbs cylindric to conic, very rarely subglobose, sometimes absent or replaced by long stems, with several to many basal or lateral leaves; viscidium absent or if present then not triangular   (25)
25 (24) Plants usually rather tall, with conic, ovoid, or subcylindric, long pseudobulbs or elongate stem; leaves sparsely growing on upper part of stem or toward apex of pseudobulb; lip often completely separate from column wings.   100 Phaius (p. 290)
+ Plants smaller, without or with small, mostly ovoid pseudobulbs; leaves subbasal; lip often ± connate along basal margins with column wings.   101 Calanthe (p. 292)
26 (13) Scape or inflorescence arising from upper part to apex of stem or pseudobulbs   (27)
+ Scape or inflorescence arising from middle to base of pseudobulbs or from rhizome   (38)
27 (26) Sepals connate into a cylindric or nearly urceolate tube   (28)
+ Sepals free or only lateral sepals connate at base to column, never forming a tube   (29)
28 (27) Inflorescence 4-10 cm, with 10-40 flowers; leaves 5-40 cm; pseudobulb not reticulate.   133 Cryptochilus (p. 361)
+ Inflorescence very short, with 1 or 2 flowers; leaves 1.5-2.5 cm; pseudobulb surface reticulate.   131 Porpax (p. 360)
29 (27) Stem with 1 distinct internode only   (30)
+ Stem usually with several internodes   (32)
30 (29) Leaves convolute; pseudobulbs conic, 2-leaved; inflorescence many flowered; flowers stellate.   119 Eria (p. 343)
+ Leaves conduplicate, sometimes terete; inflorescence 1-6-flowered; flowers not stellate   (31)
31 (30) Sepals densely tomentose abaxially.   120 Campanulorchis (p. 346)
+ Sepals glabrous abaxially.   121 Conchidium (p. 346)
32 (29) Leaves terete, fleshy; inflorescence synanthous, 1-flowered; flower relatively large, outer surface of sepals woolly.   122 Mycaranthes (p. 348)
+ Leaves dorsiventrally flattened   (33)
33 (32) Column with 2 erect, armlike appendages at apex; stem not swollen to form a pseudobulb; leaf 1.   132 Ceratostylis (p. 360)
+ Column without armlike appendages at apex; stem often swollen to form a pseudobulb; leaves 2 to many in taxa lacking pseudobulb   (34)
34 (33) Lip convex, entire, articulate to column foot and mobile; inflorescence never bottle-brush-like; column foot at right angle to column, bearing a fleshy cushion.   130 Callostylis (p. 359)
+ Lip not convex, 3-lobed or obscurely 3-lobed, if entire, then inflorescence bottle-brush-like, fixed to apex of column foot, lacking a fleshy cushion   (35)
35 (34) Floral bracts large, ca. 8 cm, bright orange; inflorescence bearing a few medium-sized resupinate flowers; rhizome stout; pseudobulbs short.   124 Dendrolirium (p. 350)
+ Floral bracts smaller, not bright orange   (36)
36 (35) Inflorescence densely flowered, bottle-brush-like; flowers small, not resupinate or ovary only slightly twisted.   125 Aeridostachya (p. 351)
+ Inflorescence not as above; flowers resupinate or not   (37)
37 (36) Pseudobulbs usually less than 1/4 as long as leaves, borne sequentially on a stout rhizome; leaves 2 or 3, apical or subapical on pseudobulb.   126 Bryobium (p. 352)
+ Pseudobulbs usually 1/2 or more as long as leaves, clustered, not noticeably arranged along rhizome; leaves 2-6 along upper part of stem.   127 Pinalia (p. 352)
38 (26) Pseudobulbs noded in middle; sepals connate into a tube; column foot conspicuously longer than column, curved upward.   102 Acanthephippium (p. 309)
+ Pseudobulbs not noded at least in middle; sepals completely free; column foot usually shorter than or as long as column, spreading horizontally   (39)
39 (38) Plants leafless at anthesis, with neither pseudobulb nor conspicuous stem; rhizome fleshy, usually geniculate.   97 Pachystoma (p. 286)
+ Plants with leaves at anthesis   (40)
40 (39) Leaf 1, petiolate or with a petiole-like pseudobulb at base; petiole similar to pseudobulb   (41)
+ Leaves 2 to many (Spathoglottis occasionally with 1 leaf), petiolate; petiole quite different from pseudobulb, sometimes overlapping to form a pseudostem   (44)
41 (40) Base of leaf blade often cuneate (only T. longiscapa and T. emeiensis subrounded); petiole distinguishable from pseudobulb.   94 Tainia (p. 281)
+ Base of leaf blade cordate or subrounded; petiole pseudobulb-like   (42)
42 (41) Flowers not resupinate, with lip at top; spur short.   93 Nephelaphyllum (p. 280)
+ Flowers resupinate, with lip at bottom, spurless or long spurred   (43)
43 (42) Lip with a slender spur; column without foot; lateral sepals not forming a mentum at base.   96 Hancockia (p. 286)
+ Lip spurless; column with a long and curved foot; lateral sepals adnate to column foot forming a broad mentum at base.   94 Tainia (p. 281)
44 (40) Lip movable, on a long column foot.   95 Eriodes (p. 285)
+ Lip immovable; column without a foot (except Calanthe labrosa)   (45)
45 (44) Leaves linear-lanceolate or lanceolate; pseudobulb ± globose; mid-lobe of lip with a claw and 2 thickened appendages.   98 Spathoglottis (p. 287)
+ Leaves elliptic or elliptic-lanceolate, if linear or linear-lanceolate, without globose pseudobulb; mid-lobe of lip without a claw   (46)
46 (45) Lip with neither spur nor sac; mid-lobe with many vesiculate appendages.   99 Cephalantheropsis (p. 288)
+ Lip often spurred, rarely spurless; mid-lobe without vesiculate appendages   (47)
47 (46) Plants rather small; leaves subbasal; lip ± connate at base with lateral wings of column to form a tube (except C. labrosa and C. actinomorpha); column often short.   101 Calanthe (p. 292)
+ Plants much taller; leaves sparsely growing on a long stemlike pseudobulb or densely growing toward apex of a large pseudobulb; lip often completely separate from column wings; column long and stout.   100 Phaius (p. 290)
48 (8) Sepals partly connate at base into a calyx tube and almost at right angle to ovary.   103 Anthogonium (p. 311)
+ Sepals free and/or not at right angle to ovary   (49)
49 (48) Column with a conspicuous foot; mentum clearly visible   (50)
+ Column without a conspicuous foot; mentum absent   (60)
50 (49) Inflorescence arising from base of pseudobulbs or from rhizome   (51)
+ Inflorescence arising from upper part of stem or pseudobulbs   (54)
51 (50) Pollinarium with neither viscidium nor stipe.   142 Bulbophyllum (p. 404)
+ Pollinarium with both viscidium and stipe   (52)
52 (51) Lateral sepals densely hispid adaxially.   143 Monomeria (p. 440)
+ Lateral sepals glabrous   (53)
53 (52) Pollinarium with a single, undivided stipe with a common viscidium.   142 Bulbophyllum (p. 404)
+ Pollinarium either with a single, y-shaped stipe with a single viscidium, or with 2 stipes each with a viscidium.   144 Sunipia (p. 440)
54 (50) Pollinarium with neither caudicle nor viscidium; stems fleshy or with internodes enlarged to form pseudobulbs, sometimes bamboolike or bamboo-shoot-like toward end, or covered completely by fleshy bases of laterally compressed leaves   (55)
+ Pollinarium with both caudicle and viscidium; stems without above combination of characters, occasionally cormlike at base   (57)
55 (54) Plants with many 1-noded pseudobulbs along a rhizome.   141 Epigeneium (p. 400)
+ Plants with 1- to many-noded or pseudobulb-like stems   (56)
56 (55) Stems not superposed; either (1) rhizomatous, (2) erect and many noded, (3) erect and 1-noded or several noded from a many-noded rhizome, or (4) rhizome absent, new stems of many nodes arising from base of old ones; leaves 1 to many; flowers long-lived or ephemeral.   139 Dendrobium (p. 367)
+ Stems superposed, non-rhizomatous part of shoot consisting of several quite long thin internodes, uppermost pseudobulbous and 1-leaved; flowers always ephemeral.   140 Flickingeria (p. 397)
57 (54) Leaves several, subbasal; stem less than 1 cm.   118 Polystachya (p. 342)
+ Leaves many, densely and distichously arranged throughout stem; stem usually more than 5 cm   (58)
58 (57) Leaves equitant, shoots iridiform; lip with a conic subapical wart below.   129 Oxystophyllum (p. 358)
+ Leaves not equitant, often twisted at base so as to be in one plane; lip without such a wart   (59)
59 (58) Pollinia 6.   135 Appendicula (p. 363)
+ Pollinia 4.   136 Podochilus (p. 365)
60 (49) Plants with a long stem; leaves cauline.   108 Thunia (p. 315)
+ Plants without a long stem; leaves basal or at apex of pseudobulbs   (61)
61 (60) Leaves laterally compressed or sometimes cylindric.   81 Oberonia (p. 236)
+ Leaves flat, not as above   (62)
62 (61) Terrestrial plants, without green naked pseudobulbs   (63)
+ Epiphytic plants, with green naked pseudobulbs   (73)
63 (62) Plants without underground pseudobulbs, sometimes with stem fleshy or stemlike pseudobulbs basally; pollinarium without caudicle, stipe, or viscidium   (64)
+ Plants with underground pseudobulbs; pollinarium with conspicuous stipe or viscidium (except Tipularia)   (68)
64 (63) Column rather long, arching; flowers resupinate.   75 Liparis (p. 211)
+ Column often very short, erect; flowers often not resupinate, with lip at top; always terrestrial   (65)
65 (64) Leaf 1 or 2, lacking prominent veins.   77 Malaxis (p. 229)
+ Leaves 2 or more, with prominent veins   (66)
66 (65) Column lacking fingerlike projections on each side of anther; anther connective broad with locules well separated; lip with lateral lobes enfolding column.   80 Oberonioides (p. 235)
+ Column with fingerlike projections on either side; anther connective narrow   (67)
67 (66) Lip entire to obscurely lobed, often with a denticulate margin, without transverse callus at base.   78 Crepidium (p. 229)
+ Lip prominently 3-lobed, with a transverse callus at base.   79 Dienia (p. 234)
68 (63) Plants with a single flower   (69)
+ Plants with many flowers   (70)
69 (68) Sepals shorter than 2 cm; lip with a horizontally spreading sac.   86 Calypso (p. 251)
+ Sepals longer than 2.5 cm; lip with an incurved spur.   87 Changnienia (p. 252)
70 (68) Lip with a cylindric spur at base; spur conspicuously longer than pedicel and ovary.   85 Tipularia (p. 250)
+ Lip spurless or with a spur conspicuously shorter than pedicel and ovary   (71)
71 (70) Flowers pendulous; sepals 1.7-3 cm.   84 Cremastra (p. 249)
+ Flowers not pendulous; sepals 1.5-11 mm   (72)
72 (71) Lip saccate or shortly spurred at base; pollinarium without a distinct stipe.   85 Tipularia (p. 250)
+ Lip neither saccate nor shortly spurred at base; pollinarium with a slender stipe.   83 Oreorchis (p. 245)
73 (62) Leaves membranous or papery; lip unlobed, without concave or saccate base   (74)
+ Leaves thickly leathery; lip, if unlobed, with concave or saccate base   (75)
74 (73) Petals not Y-shaped; pollinia 4, without distinct caudicles.   75 Liparis (p. 211)
+ Petals deeply 2-lobed, ± Y-shaped; pollinia 2, each with caudicle.   76 Ypsilorchis (p. 228)
75 (73) Sepals concave and saccate at base.   115 Neogyna (p. 341)
+ Sepals not concave or saccate at base   (76)
76 (75) Lip spurred   (77)
+ Lip spurless, sometimes saccate at base   (78)
77 (76) Pseudobulbs each with 2 leaves at apex; flowers many, on pendulous raceme; spur curved upward.   116 Bulleyia (p. 341)
+ Pseudobulbs each with 1 leaf at apex; flower solitary, not pendulous; spur straight.   117 Ischnogyne (p. 342)
78 (76) Lip concave-saccate at base   (79)
+ Lip not or only slightly concave, but never saccate, at base   (80)
79 (78) Column thick and short, usually shorter than lip.   113 Pholidota (p. 335)
+ Column slender, usually nearly as long as lip.   114 Otochilus (p. 339)
80 (78) Lip sigmoidally curved at base.   111 Panisea (p. 333)
+ Lip not sigmoidally curved at base   (81)
81 (80) Raceme with 20-30 flowers; flowers ca. 1 cm in diam.; column with 2 armlike appendages on both sides.   112 Dendrochilum (p. 334)
+ Raceme often with several flowers or reduced to a single flower; flowers more than 3 cm in diam.; column without armlike appendages   (82)
82 (81) Plants with persistent leaves alive for more than 1 year, present at anthesis; flowers often more than 2, white, yellow, green, or brown, lacking reddish spots on lip.   109 Coelogyne (p. 315)
+ Leaves annually deciduous, absent or very young at anthesis; flower solitary, rarely 2, usually pink to purple, rarely yellow or white, with reddish spots on lip.   110 Pleione (p. 325)

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