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SUBFAMILY APOSTASIOIDEAE REICHENB.

1841, Repert.Herb.: 56 (sub "Apostasieae"). - Fam. Apostasiaceae Lindl., 1833, Nixus Pl. : 22 (sub "Apostasieae").

Type: Apostasia Blume.

2(2) genera and 15(5) species. Tropical and subtropical Asia to N.Australia and New Guinea .

Neuwiedia Blume,

1834, in Hoeven et De Vriese, Tijdschr.Nat.Geschied. 1: 140; id., 1834, Ann. Sci. Nat. (Paris) 2, 2: 93; De Vogel, 1969, Blumea 17, 2: 313-350; Chen et Lang, 1986, Acta Phytotax.Sin. 24, 5: 346-348.

Type: N. veratrifolia Blume.

Sympodial terrestrial herbs with erect semi-woody unbranched stem, often with few aerial roots at the base. Leaves numerous, spirally arranged, plicate, the upper becoming bract-like. Inflorescence sub-terminal, erect, many-flowered, dense raceme. Flowers pedicellate, resupinate, not opening widely, campanulate, hardly zygomorphic. Tepals free, sub-similar, lanceolate to narrowly obovate, petals are thinner in texture, the lip a little broader. Column straight, with 3 fertile stamens, filaments basally jointed to the style, anthers free. Fruit dry capsule or juicy berry. Seeds with hard alveolate testa.

6-8(1) species. Mainland Southeast Asia to SW. Pacific islands.

1. Neuwiedia zollingeri Reichenb.f.,

1857, Bonplandia 5: 58.

Described from Java (“s.l.”). Type (“ Zollinger 2808 ” ) – W.

Distribution. Mainland Southeast Asia, Java, Sumatra, Kalimantan .

Key to varieties

1. Inflorescence glabrous or with very few scattered hairs . . . . 1. N. zollingeri var. annamensis

-. Inflorescence densely hairy . . . . 2.

2. Flowers brightly yellow; ripe fruit white . . . . 2. N. zollingeri var. javanica

-. Flowers pure white with yellow central spot on lip; ripe fruit orange to red . . . 3. N. zollingeri var. balansae

1a. N. zollingeri var. annamensis (Gagnep.) Aver., comb. nov.

– N. annamensis Gagnep., 1933, Bull. Soc. Bot. France 80: 350; Seidenf., 1992, Orch. Indochina: 11; Aver., 1994, Identif. Guide Viet. Orch.: 11; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 46.

Described from S. Vietnam (“ Annam : Nui-han-heo, pres Nhatrang...”). Type (“ Poilane 4937 ” ) – P.

Plant up to 40 cm tall, glabrous or with few scattered hairs. Leaves numerous, thin, close together below inflorescence, petiolate; leaf blade narrowly elliptic to elliptic, acute, 18-50 by 4- 6 cm . Inflorescence 6- 15 cm long, bearing 30-60 flowers. Floral bracts triangular, acuminate, 1-2 by 0.3- 0.5 cm , as long as flowers. Flowers yellow. Tepals lanceolate to narrowly obovate, 15-23 by 4- 8 mm , obtuse, ending below apex in short mucro; petals and lip slightly shorter and wider, dorsally with low keel. Stamens 1.2- 1.8 cm long, longer than style; anthers 6- 8 mm long. Style straight, 1- 1.6 cm long with roundish stigma. Fruit white juicy berry, 5- 8 mm across.

Ecology. Shady broad-leaved evergreen lowland and submontane forests on granite and gneiss. 900 m . Fl. May. Very rare (DD).

Distribution. Vietnam (Khanh Hoa).

Studied specimens. Nui Han Heo, Nhatarang, Poilane, 4937 (P), Poilane, 6180 (P).

Notes. Closely allied to type variety, from which distinctly differs in large flowers.

1b. N. zollingeri var. javanica (J.J.Smith) de Vogel,

1969, Blumea, 17, 2: 329. – N. javanica J.J.Smith., 1914, Bull. Jard. Bot. Buitenzorg 2, 14: 5.

Described from Java (“…Artana…”). Lectotype (“ J.J.Smith s.n. a. 1900” ) – BO.

Plants up to 40 cm tall, all parts more or less densely haired. Flowers bright yellow. Dorsal keel on petals and lip densely hairy. Ripe fruit white juicy berry. Fig. 14, a-d.

Ecology. Shady broad-leaved evergreen submontane forests on granite and sandstone. 600- 950 m . Fl. March. Rare (LR).

Distribution. Vietnam (Dak Lak, Ninh Thuan). Java, Sumatra, Bali .

Studied specimens. Ninh Thuan, Nui Chua mt HLF 4415 (HN, LE); Dak Lak, Chu Yang Sin mt, HLF 5479 (HN, LE).

Notes. Ripe berries of this variety spicy-fragrant, sweet and edible.

1c. N. zollingeri var. balansae (Gagnep.) Aver., comb. nov.

– N. balansae Gagnep., 1933, Bull. Soc. Bot. France 80: 350. - N. griffithii auct. non Reichenb.f.: Seidenf., 1992, l .c.: 11; Aver., 1994, Identif. Guide Viet. Orch.: 12; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 46. - N. inae auct. non de Vogel: Seidenf., 1992, l .c.: 13; Aver., 1994, l .c.: 12; Aver. et Averyanova, 2003, l .c.: 46.

Described from N. Vietnam ("Tonkin: mt. Bavi , …"). Lectotype (" n° 2000 ( Balansa )") – P.

Plants up to 40 cm tall, all parts more or less densely. Flowers pure white, lip with yellow spot in center. Dorsal keel on petals and lip densely hairy. Ripe fruit orange to red juicy berry. Fig. 13, a ; 14, e-g.

Ecology. Shady broad-leaved, evergreen lowland and submontane forests of sandstone, granite and rocky limestone. 450- 1250 m . Fl. May - June. Rare (LR).

Distribution. Vietnam (Dak Nong, Khanh Hoa, Kon Tum; Ninh Binh, Phu Tho, Quang Ninh).

Studied specimens. Cam Pha, Cong Hoa, Quang Ninh, Tran s.n. (LE, UHN); Ninh Binh, Cuc Phuong, Duc B 356 (HN), CPNP 5116 (HN), N.M.Cuong, H.L.Quyen, NMC 309 (HN), DDS 11060 (HN), Averyanov et al., HAL 1686 (HN, LE), Duc B 356 (HN), N.T.Hiep, NTH 3056 (HN); Dak Nong Prov., Ta Dung mt, HLF 5567 (HN, LE); Mt. Bavi, Ha Son Binh 800 m , Balansa, 2000 ; Song Cao near Nhatrang 800 m , Evrard, 508 (AMES) & Poilane, 3408 (P); Dakto, Gialai-Kontum, Dao, 230 (HN).

Notes. Closely allied to N. zollingeri var. singapureana (Backer) de Vogel, from which differs in large pure white flowers. Specimens from Cuc Phuong national park occasionally have flowers with 4 fertile stamens.

Apostasia Blume,

1825, Bijdr. 6, fig. 39; id. 1825, ibid. 8: 423; De Vogel, 1969, Blumea 17, 2: 313-350; Chen et Lang, 1986, Acta Phytotax. Sin. 24, 5: 348-351.

Type: A. odorata Blume.

Sympodial terrestrial herbs with erect semi-woody commonly unbranched stem, with few aerial roots at the base. Leaves few to numerous, spirally arranged, plicate, with acute to filiform tip. Inflorescence sub-terminal, erect to arching and pendent, usually branched, many-flowered. Flowers, sessile, not resupinate, opening widely, hardly zygomorphic. Tepals free, sub-similar, broadly lanceolate, more or less recurved, with dorsal keel extending apically into acute beak, the lip a little larger and wider. Column straight or curved, usually with 2 fertile stamens, often with 1 staminode, filaments short, basally jointed to the style; anthers free; stigma 3-lobed or suborbicular. Fruit dry cylindrical capsule. Seeds with black hard alveolate testa.

7(4) species. Tropical and subtropical Asia to N. Australia and New Guinea .

Key to species

1. Plants 5- 12 cm tall; leaves petiolate, leaf blade ovate to elliptic, 1.5-2.5 cm long, 0.5-1 cm wide . . . . . 3. A . nipponica

-. Plants 20- 100 cm tall; leaves sessile, linear to lanceolate, 7- 35 cm long, 0.3- 2.2 cm wide . . . . . 2.

2. Inflorescence at the base with bunch of numerous sterile bracts; flowers white or yellow; tepals strongly reflexed right back; staminode absent . . . . . . . . 4. A . nuda

-. Inflorescence without bunch of sterile bracts at the base; flowers yellow; tepals reflexed back or straight; staminode present, joined most of the way to style. . . . . . 3

3. Tepals less than 6 mm long; capsule less than 1.8 cm long; column terete, without wings below staminode; leaves linear to narrowly lanceolate, commonly to 1.5 cm wide . . . . 1. A . wallichii

-. Tepals more than 6 mm long; capsule longer than 1.8 cm ; column more or less flattened, with 2 wings below staminode; leaves narrowly lanceolate to narrowly elliptic, commonly 1.5-2 cm wide . . . . 2. A . odorata

1. A . wallichii R.Br.,

1830, in Wall. Pl. As. Rar. 1: 75; Seidenf., 1992, Orch. Indochina: 13; Aver., 1994, Identif. Guide Viet. Orch.: 13; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 11.

Described from Nepal ("in valle Nagaliae minora, Noakote dicta"). Lectotype (" Wallich, 4448 ") - K (holo), E (iso).

Plant 15- 100 cm tall. Leaves numerous, close, linear to narrowly lanceolate, 12-35 by 3- 15 mm . Inflorescence, simple to many branched, many flowered. Flowers pale yellow. Tepals 4-5 by 0.6- 1.2 mm . Column straight to strongly curved at base, cylindrical, without wings. Stamens 2, 3- 4 mm long. Staminode entirely adnate to column or with free obtuse tip. Ovary 1- 1.7 cm long, 0.8- 1 mm in diam. Capsule 1- 1.8 cm long, 1.4- 2 mm wide. Fig. 15, a , b.

Ecology. Shady broad-leaved evergreen and coniferous lowland and submontane forests on shale, sandstone, granite, gneiss and basalt. 200- 1200 m . Fl. March, April. Occasional (LR).

Distribution. Vietnam (Dong Nai, Gia Lai, Khanh Hoa, Kon Tum, Lam Dong, Ninh Thuan, Phu Tho, Quang Binh, Quang Nam, Tay Ninh, Thua Thien – Hue). Tropical Asia from Sri Lanka and Nepal to Philippines , N. Australia and New Guinea .

Studied specimens. Prenh, Dalat, Evrard, 1469, 2314 (P); Lo Go, Tay Ninh, Mueller, 1470 (P); Gian Lau me, Dong Nai, Bienhoa, Pierre, 6583 (P); Mt Tamire, Pierre, 6583 (P); Nhatrang, Poilane, 3808 (P); Ba Na, Danang, Poilane, 7277 (P); Go O to Mo O, Quang Nam 1000 m, Poilane, 31671 (P); Gialai – Kontum, Kbang – Kon Ha Nung, LX-VN 2739 (LE); Khanh Hoa, Khanh Son, Hon Giao mt., VH 4322 (HN, LE); Kon Tum, Kon Plong, DKH 4690 (HN, LE); Kontum, Kon Plong, Mang La, VH 5185 (HN, LE); Kontum, Kon Plong, Mang La, VH 5327 (HN, LE); Ninh Thuan, Ninh Hai, Nui Chua mt., HLF 4321 (HN, LE), HLF 4362 (HN, LE); Ninh Thuan, Ninh Hai, Nui Chua mt., HLF 4362 (HN, LE); Thua Thien – Hue, A Luoi, HAL 7968 (HN, LE); Quang Binh, Minh Hoa, HAL 11708 (HN).

2. A . odorata Blume,

1825, Bijdr. 6, fig. 39; id. 1825, ibid. 8: 423; Seidenf., 1992, Orch. Indochina: 13; Aver., 1994, Identif. Guide Viet. Orch.: 13; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 11. – A. thorelii Gagnep., 1933, Bull. Soc. Bot. France 70: 350.

Described from Java ("montis Salak"). Lectotype (" Blume s.n. ") - L.

Plant 30- 80 cm tall. Leaves numerous, close, narrowly lanceolate to lanceolate, 15-25 by 1.5- 2.2 cm . Inflorescence, simple to few branched, many flowered. Flowers bright yellow. Tepals 9-12 by 1.5- 2 mm . Column straight to strongly curved at base, usually laterally flattened, with 2 short longitudinal low to tall quadrangular wings below staminode. Stamens 2, 5- 6 mm long. Staminode adnate to column with free obtuse tip. Ovary 10- 18 mm long, 1- 1.2 mm wide. Capsule 1.8-2.5 cm long, 1.5- 2.5 mm wide. Fig. 13, b-d; 15, c-e.

Ecology. Shady broad-leaved evergreen and coniferous lowland, submontane and montane forests on shale, sandstone, granite, gneiss, rhyolite and quartzite. 300- 1600 m . Fl. March - June. Occasional (LR).

Distribution. Vietnam (Dak Lak, Dong Nai, Ha Tinh, Khanh Hoa, Lam Dong, Lao Cai, Phu Tho, Quang Binh, Vinh Phuc). Mainland tropical Asia, Hainan, Java, Sumatra, Kalimantan .

Studied specimens. Bao Loc 1000-1200, Poilane, 22045 (P); Bavi, 1200 m , Chevalier, 37777 (P); Dak Lak, Chu Yang Sin mt., HLF 5481 (HN, LE); Ha Tinh, Huong Son, HAL 1281 (HN, LE), HAL 5337 (HN, LE); Khanh Hoa, Khanh Son, Hon Giao mt., VH 4174 (HN, LE), VH 4378 (HN, LE); Lao Cai, Van Ban, DKH 6626 (HN, LE), DKH 6905 (HN, LE); Lao Cai, Van Ban, HAL 2354 (HN, LE); Vinh Phu, Tam Dao 3, LX-VN 2368 (LE); Quang Binh, Quang Ninh distr., HAL 11519 (HN); Quang Binh, Minh Hoa, HAL 11624 (HN).

Notes. The name A. thorelii is attributed to plants with tall quadrangular column wings and rather broad leaves. Taxonomical status of such marginal plants is still uncertain.

Fig. 13. Neuwiedia zollingeri var. balansae: a – flower. Apostasia odorata: b – flowering
plant, c – flower, d – column; A. nipponica: e – flowering and fruiting plant, f – flower. A. nuda:
g – flowering plant.

Fig. 14. Neuwiedia zollingeri var. javanica: a, b (HLF 4415), c, d (HLF 5479). N. zollingeri
var. balansae: e, f (HAL 1686), g (HLF 5567).

Fig. 15. Apostasia wallichii: a, b (VH 5185). A. odorata: c-e (HAL 5337). A. nipponica: f (HLF
1406), g, h (HAL 7175).

3. A . nipponica Masam.,

1935, Journ. Jap. Bot. 11: 46.

Described from Ryukyu Islands (“Yakusima, Kurio, …”). Type (“Jul.13.1933 (leg. S.Suzuki )”) - ?

Plant 5-10(12) cm tall. Leaves few, distant, distinctly petiolate, petiole 0.5-1 cm long, leaf blade ovate to elliptic, 1.5-2.5 by 0.5-1 cm. Inflorescence, simple or with 1-3 branches bearing 1-4 flowers. Flowers bright yellow. Tepals 3.5-5.5 by 1- 1.4 mm . Column straight to slightly curved at base, without wings or with 2 short longitudinal insignificant wings below staminode. Stamens 2, 2- 3 mm long. Staminode adnate to column with free orbicular tip. Ovary 6- 9 mm long, 0.5- 1.2 mm wide. Capsule 0.8- 1.1 cm long, 1.4- 1.8 mm wide. Fig. 13, e, f; 15, f-h.

Ecology. Shady broad-leaved evergreen lowland and submontane forests on shale, sandstone and granite. 300-1000 m. Fl. April, May. Very rare (LR).

Distribution. Vietnam (Ha Tinh, Thua Thien – Hue ). Ryukyu Islands .

Studied specimens. Ha Tinh, Huong Son, Rao An, No HAL 5157 (HN, LE); Thua Thien – Hue , A Luoi, HAL 7175 (HN, LE); Thua Thien Hue , Bach Ma, HLF 1406 (HN, LE); Thua Thien Hue , Nam Dong, HAL 10928 (HN, LE).

4. A . nuda R.Br.,

1830, in Wall. Pl. As. Rar. 1: 76, tab. 85; Seidenf., 1992, Orch. Indochina: 15; Aver., 1994, Identif. Guide Viet. Orch.: 13; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 11.

Described from Malacca Peninsular ("in montibus insulae Penang "). Lectotype (" Porter s.n. Wallich, 4449 ") - K.

Plant 12- 40 cm tall. Leaves numerous, close, linear to linear lanceolate, 7-20 by 3- 8 mm . Inflorescence, simple to few branched, many flowered with tuft of small narrow sterile bracts at base. Flowers white to yellow. Tepals 3.5-4.5 by 0.6- 0.8 mm . Column strongly curved. Stamens 2, 3.2- 3.5 mm long. Staminode absent. Ovary 8- 10 mm long, 0.8- 1 mm wide. Capsule 1.2- 1.4 cm long, 1 mm wide. Fig. 13, g .

Ecology. Shady coniferous and broad-leaved evergreen lowland and mountain forests on shale, 320- 1400 m . Fl. January, February. Very rare (DD).

Distribution. Vietnam (Dak Lak, Thua Thien – Hue ). Myanmar , Thailand , Malacca Peninsular, Java, Sumatra, Kalimantan .

Studied specimens. Dak Lak, Chu Yang Sin mt., HLF 5438 (HN, LE); Thua Thien – Hue , Nam Dong, HAL 6983 (HN, LE).

Notes. Flowers of this species varies from yellow to white. Flower color of Vietnamese samples unknown.

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