A REVISION OF EPIGYNUM (APOCYNACEAE: APOCYNOIDEAE) DAVID J. MIDDLETON1 Abstract. The genus Epigynum Wight from Southeast Asia is revised. Five species are recognized. A key to the species and descriptions are given. Keywords: Epigynum, Apocynaceae, Apocynoideae, revision, Asia.
This work on Epigynum marks the latest in a series of revisions of Apocynaceae with the aim of completing subfamilies Rauvolfioideae and Apocynoideae (sensu Endress and Bruyns, 2000) for the Flore du Cambodge, du Laos et du Viêtnam and for Flora Malesiana. Epigynum is a small genus in the Apocynaceae, subfamily Apocynoideae, found in Northeastern India, Southern China, Burma, Thailand, Indochina, Peninsular Malaysia, Sumatra, and Borneo. They are woody climbers in evergreen and deciduous forests, often at forest margins, with relatively attractive and fragrant flowers. Although many names have been published, the material available falls into five readily distinguishable species. Pitard (1933) published the new genus Argyronerium with one species, A. odoratum. This is very clearly an Epigynum, and Kerr (1939) made the new combination Epigynum odoratum, here treated as a synonym of E. grif fithianum. Pitard made no attempt to distinguish his genus from Epigynum, and given that no Epigynum species were known from Indochina at that time, it is quite likely he was simply unaware of the older name.
The status of Nouttea has been discussed by Middleton (1994). Xu (1988) included Sindechites chinensis (Merr.) Markgr. & Tsiang in Epigynum, where it had originally been described by Merrill (1923). He did indicate that it was an isolated member of the genus and that its status should be reviewed. Li et al. (1995) and Middleton (1999) both excluded it from Epigynum and placed it in Sindechites. Recent phylogenetic work (Livshultz and Middleton, in prep.) has shown that Sindechites chinensis is not close to the other Epigynum species, but that its relationship with regard to the only other species of Sindechites, S. henryi, is also not clear. For our purposes here, it is clearly not an Epigynum. Xu (1988) also indicated that Epigynum auri tum should be removed from Epigynum. The situation with this species, and its close relative E. graciliflorum, is more problematical. Livshultz and Middleton (in prep.) have not demonstrated that E. auritum forms a monophyletic group with the other Epigynum species, but there is also not enough support for splitting the genus. For the time being I shall keep it in Epigynum, and the issue will be discussed further in the forthcoming paper.
MATERIALS AND METHODS Approximately 160 herbarium collections from continental Asia and Malesia have been studied. These are from the following herbaria: A, AAU, ABD, BKF, BM, BO, BRI, C, CMU, E, F, GH, K, KEP, HN, HNU, K, L, NY, OXF, P, SING, TCD, U, US, Z (Holmgren et al., 1990). All specimens have been seen unless
otherwise indicated. All dimensions given are for dried specimens except for androecium and gynoecium characters, which are from flowers reconstituted by boiling in water. Author citation follows Brummitt and Powell (1992).
I would like to thank the curators and staff of the herbaria that loaned material or hosted me on visits and the curatorial staff of the Harvard University Herbaria for their management of the loans. 1 Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. Current address: Royal Botanic Garden, 20A Inverleith Row, Edinburgh EH3 5LR, Scotland. Email:
[email protected] Harvard Papers in Botany, Vol. 10, No. 1, 2005, pp. 67– 81. © President and Fellows of Harvard College, 2005.
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TAXONOMIC TREATMENT Epigynum Wight, Ic. 4: t.1308. 1848. Benth. & Hook. f., Gen. Pl. 2: 717. 1876; K. Schum. in Engler & Prantl, Nat. Pflanzenfam. 4,2: 178. 1895; Pichon, Mém. Mus. Nat. Hist. Nat. sér. B, Bot. 1: 103. 1950. Type species: Epigynum griffithianum Wight. Synonyms: Legouixia Heurck. & Müll. Arg. in Heurck., Obs. Bot. 145. 1871. Type species: Legouixia amabilis Heurck. & Müll. Arg. Nouettea Pierre, Bull. Mens. Soc. Linn. Par. sér. 2, 1: 29. 1898. Type species: Nouettea cochinchinensis Pierre. Trachelospermum subgen. Lachnocarpus C. K. Schneid. in Sargent, Pl. Wilson. 3: 341. 1916. Type species: Trachelospermum auritum C. K. Schneid. A rg y ronerium Pitard, Fl. Gén. IndoChine. 3: 1234. 1933. Type species: Argyronerium odoratum Pitard. Large woody climbers with all parts bearing white latex. Branchlets weakly lenticellate or not, pubescent at least when young. Leaves opposite; glands in axils of petioles; blade ovate, elliptic or obovate, apex with a sharp tip,
secondary veins arcuate ascending, anastomosing before margin. Inflorescence an umbelliform panicle formed from terminal and axillary cymes; bracts small. Sepals with colleters in a row at base inside, only at edges of sepals at base inside, or without colleters. Corolla in bud a narrow tube, bulging around the anthers, and an ovate head, lobes dextrorse, tomentose to glabrous outside, pubescent or glabrous inside, salverform when open; lobes falcate-obovate, mostly with a sharp projection on one side. Stamens inserted in lower half of tube or almost in the middle of tube; filaments broad, short; anthers narrowly triangular, apex acuminate, base sagittate, sterile at apex and base, attached at base of style head and attached again at middle of anthers to top of style head. Disk of 5 separate lobes or in a ring with a crenate margin; glabrous. Gynoecium of 2 separate semi-inferior carpels united into a common style; ovules numerous; style filiform; style head long cylindrical, with an acuminate apex and a basal collar, collar sometimes very small, 5 ribs between collar and apex. Fruit of paired follicles; fusiform or linear, sometimes strongly curled. Seeds consisting of a flattened grain and a coma directed towards the fruit apex.
KEY TO THE SPECIES 1a. Corolla tube 30–49 mm long, usually somewhat curved, 4.5–6.0 mm wide across anther bulge, tinged pink; stamens inserted at 0.4–0.5 of corolla tube length from base; fruit strongly curled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. Epigynum cochinchinense 1b. Corolla tube 13–38 mm long, usually straight, 1.7–3.0 mm wide across anther bulge, white or cream (unknown in E. graciliflorum); stamens inserted at 0.1–0.3 of corolla tube length from base; fruit straight or only slightly curved outward or inward. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Leaf-blade base rounded to cordate; sepal colleters present; corolla throat glabrous or with few hairs. . . . . . . . 3 2b. Leaf-blade base cuneate to obtuse, or rarely rounded; sepal colleters absent; corolla throat densely villous . . . 4 3a. Sepals with colleters only at the inside basal edges; ovary pubescent . . . . . . . . . . . . . . . . . 1. Epigynum auritum 3b. Sepals with a continuous row of colleters inside; ovary glabrous . . . . . . . . . . . . . . . . 3. Epigynum graciliflorum 4a. 9–14 pairs of secondary veins but always with at least some leaves on plant with >9 pairs of secondary veins; never with tuft of hairs in secondary vein axils with midrib unless all of blade beneath pubescent; ovary densely to sparsely puberulent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. Epigynum ridleyi 4b. 5–9(–11) pairs of secondary veins; often with tuft of hairs in secondary vein axils with midrib or all of blade beneath pubescent; ovary glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. Epigynum griffithianum
1. Epigynum auritum (C. K. Schneid.) Tsiang & P. T. Li, Acta Phytotax.Sin 11: 397. 1973; Tsiang & Li, Fl. Reip. Pop. Sin. 63: 246. 1977; Li et al., Fl. China 16: 187. 1995; Turner, Gard. Bull. Sing. 47: 126. 1997; Middleton, Fl. Thailand 7: 118. 1999. Fig. 1–2.
Basionym: Trachelospermum auritum C. K. Schneid. in Sargent, Pl. Wils. 3: 341. 1916; Woodson, Sunyatsenia 3: 72. 1936. TYPE: CHINA. Yunnan, Szemao, Augustine Henry 12136 (Lectotype: A, designated by Woodson, Sunyatsenia 3: 72. 1936; Isolectotypes: K, NY).
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FIGURE 1. Distribution of Epigynum auritum (C. K. Schneid.) Tsiang & Li.
Synonyms: Epigynum lachnocarpum Pichon, Bull. Mus. Hist. Nat. Hist ser. 2, 20: 192. 1948, non Epigynum curtisii King & Gamble, 1907. Replaced synonym: Trachelospermum curtisii King & Gamble, J. As. Soc. Beng. 74,2: 498. 1907; Ridley, Fl. Mal. Pen. 2: 366. 1923; Woodson, Sunyatsenia 3: 73. 1936; Kerr, Fl. Siam. En. 2: 473. 1939. TYPE: MALAYSIA. Penang, Balick Pulau, May 1886, Charles Curtis 838 (Lectotype: K, designated by Woodson, Sunyatsenia 3: 73. 1936; Isotype: SING, fragment A). Climber, recorded to 16 m high. Branchlets sparsely lenticellate or not; sparsely to densely pubescent, often with a combination of shorter and longer crisped hairs. Leaves: petiole 5–27 mm long, densely brown pubescent; blade 4.1–18.5 × 2.2–11.0 cm, 1.3–2.4 times as long as wide, ovate to broadly elliptic or obovate, apex acuminate or apiculate, base rounded to cordate, sparsely pubescent with crisped hairs above and beneath, more densely pubescent on the midrib above and beneath, 7–10 pairs of secondary veins, tertiary venation irregularly scalariform. Inflorescence of axillary and terminal dichasial cymes, mostly somewhat
umbelliform, lax to somewhat congested, 4–12 cm long, bracts small, mostly deciduous; peduncle 0.4–7.5 cm long, densely tomentose, mostly with a combination of shorter and longer crisped hairs; pedicels 1.5–9.0 mm long, tomentose. Sepals 1.6–3.9 × 0.6–0.9 mm, 2.3–4.8 times as long as wide, oblong to lanceolate, apex acute, sparsely to densely pubescent, with colleters on the inner surface at the lower edges of each sepal. Corolla white, fragrant; tube 13–21 mm long, 1.8–2.3 mm wide, 4.6–10.3 times as long as sepals, 1.2–2.3 times as long as lobes, tomentose outside, often more densely so in upper half, sparsely to densely pubescent in upper part of tube above stamens inside, glabrous or with few hairs in throat; lobes 7.2–17.0 × 4.4–7.2 mm, 1.8–2.5 times as long as wide, obovate, falcate, with a sharp projection on one side, outside densely tomentose on part exposed in bud, otherwise glabrous, inside glabrous or with few hairs at base of lobes. Stamens inserted at 0.2–0.3 of tube length from base; filaments 0.8–1.0 mm long; anthers 4.7–5.4 × 0.6–0.8 mm, 6.0–7.8 times as long as wide. Disk of 5 separate squarish lobes, apex rounded, 0.7–0.9 mm high. Ovary very slightly semi-inferior, 0.6–1.2 mm high, densely pubescent; style 4.0–4.4 mm long;
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FIGURE 2. Epigynum auritum (C. K. Schneid.) Tsiang & P. T. Li.
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style head 2.1–2.5 mm long. Fruit of paired pendulous follicles, rather flattened, linear to fusiform, often curved outward from or inward to each other toward end, sparsely to densely pubescent, 12.8–33.0 cm long, 7.5–16.0 mm wide. Seeds 12–23 × 4–16 mm; coma 31–41 mm long. Distribution: northeastern India, southern China, Burma, Thailand, Laos, and Peninsular Malaysia. It is the most widespread species in the genus. Habitat: it is found in primary and secondary forest, in evergreen and deciduous forest, and on a wide variety of soil types from 100–1200 m. Additional material studied: BURMA: Myitkyina, above Chipwi, Kermode 17371 (K). Taninthayi State: Paungdaw, west of power station, Keenan et al. 1597 (E, K). Tenasserim: eastern Tenasserim, Kerr 21648 (K). CHINA: Yunnan: Ban–Chiou–Chian, Che–li, Wang 79564 (A); Jah–leei, Che–li Hsien, Wang 79185 (A); Maan–tsang, Sheau–meng–yeang, Luh–shuen Hsien, Wang 81055 (A); Nan–Chiao, Wang 77048 (A); Salwm Valley, Forrest 29521 (E); Sheau–meng–yeang, Che–li, Wang 75895 (A); Szemao, S mts., Henry 12136 (A, K), 12761 (A), Henry 12852 (A, K). INDIA: Meghalaya: Khasi Hills, Mann 167 (K). Nagaland: Naga Hills, Dimapur, Koelz 25169 (L). LAOS: Champasak: Patounphone District, near Nong Hin Village, Middleton & Lamxay 260 (A, BKF, E). MALAYSIA: Penang: Balick Pulau, Curtis 838 (K, SING). Perak: Larut, King’s Collector 5194 (K). THAILAND: sine loc.: Put 1620 (BM, K). Unknown province: Puugah, Haniff & Nur 3893 (A, BM, SING). Chanthaburi: Khao Soi Dao, Maxwell s.n. (AAU); Makham, Smitinand 3384 (BKF), Smitinand 3700 (BKF). Chiang Mai: Doi Chiang Dao, Geesink et al. 8239 (L). Chon Buri: Khao Kieo, Sriracha District, Maxwell s.n. (AAU, L). Chumphon: Ban Racha Krude, Larsen et al. 43170 (AAU, P); Kra Buri District, Thungraya Nasak Wildlife Sanctuary. Along river near Khlong Hin Lo Ranger Substation., Middleton et al. 1457 (A, AAU, BKF, E). Kanchanaburi: Sangklaburi District, Tung Yai Naresuan Wildlife Reserve, Ban Saneh Poong Village, Wah Mee Mountain, Maxwell s.n. (A, L). Nakhon Si Thammarat: Suvanakoses 348 (BKF); Khao Luang Foothills, Geesink & Santisuk 5410 (AAU,
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BKF, C, E, K, P); Khiri Wong, trail from village up to Khao Luang area, Van Beusekom et al. 737 (AAU, BKF, C, E, K, L, P); Lan Saka, Sangkhachand 714 (BKF); Tung Song, Rabil 172 (A, ABD, BM, K). Prachuap Khiri Khan: Bang Saphan District, Huay Yang National Park. Kha On Waterfall area, trail towards Burmese border, Middleton et al. 2581 (A, BKF), Middleton et al. 2602 (A, BKF); Hua Hin District, Kaeng Krachan National Park. Pa La U, trail to Chol Na Nat waterfall., Middleton et al. 1073 (A, CMU, BKF, E, K, L, Z). Ranong: Kra Buri District, Thungraya Nasak Wildlife Sanctuary. Near W ildlife Sanctuary headquarters., Middleton et al. 2781 (A, BKF). Surat Thani: Khao Pa Ngan, Put 1244 (BM, K, P), Put 1255 (BM, K); Ko Samui, Put 878 (BM, E, K, OXF, P, TCD); Tha Chang, Sangkhachand 16 (BKF). Trang: Khao Chong, Din 293 (K); Khao Pad Pha near Patthalung, Geesink et al. 7257 (AAU, BKF, K, L, P); Muang Trang, Boonkerd 50 (BKF), Nakkan 293 (BKF), Phusomsaeng 249 (BKF). The position of this species within Epigynum is not without doubt (see introduction). 2. Epigynum cochinchinense (Pierre) D. J. Middleton, Kew Bull., 49: 304. 1994; Middleton, Fl. Thailand 7: 119. 1999. Fig. 3–4. B a s i o n y m: Nouettea cochinchinensis Pierre, Bull. Mens. Soc. Linn. Par. n. s. 1: 30. 1898; Pitard in Lecomte, Fl. Gén. IndoChine 3: 1232. 1933; Kerr, Fl. Siam. En. 2: 471. 1939; L y, ´ Fedd. Rep. 97: 652. 1986. TYPE: VIETNAM. Ben Tre, Jean Baptiste Louis Pierre 4468 (Lectotype: HM, designated by L y´, Feddes Repert. 97: 652. 1986; Isotypes: K, NY, P). Chonemorpha nouettiana Pierre, Bull. Mens. Soc. Linn. Paris sér. 2, 1: 30. 1898, nom. illeg. Climber, recorded to 12 m high. Branchlets weakly lenticellate or not; densely gray or pale brown pubescent. Leaves: petiole 5–17 mm long, densely pubescent; blade 4.4–23.5 × 2.4–10.2 cm, 1.9–3.6 times as long as wide, elliptic to obovate, apex acuminate with a sharp tip, base rounded to cordate, pubescent on midrib beneath, sparsely pubescent or glabrous over rest of blade beneath, mostly sparsely pubescent at base of midrib above, otherwise
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FIGURE 3. Distribution of Epigynum cochinchinense (Pierre) D. J. Middleton (★) and Epigynum graciliflorum (Pierre) Pichon.
glabrous above, 5–13 pairs of secondary veins, tertiary venation irregularly scalariform and reticulate. Inflorescence semi-pendulous, of terminal and sometimes also axillary dichasial cymes, umbelliform, fairly lax, 8.0–12.5 cm long; peduncle 1.6–9.6 cm long, densely pubescent; pedicels 3–8 mm long, densely to sparsely pubescent. Sepals 2.2–4.5 × 1.3–3.4 mm, 1.0–2.2 times as long as wide, thickened at the base, ovate, acute to obtuse, glabrous, with few scattered hairs, sparsely pubescent only on central line or sparsely pubescent all over, colleters in a continuous row at base inside. Corolla white and suffused pink or red, often somewhat curved upward from semi-pendulous inflorescence; tube 30–49 mm long, 4.5–6.0 mm wide, 10.0–13.5 times as long as sepals, 1.1–2.6 times as long as lobes, glabrous to sparsely pubescent outside, glabrous to sparsely puberulent in upper half inside; lobes 13–34 × 9.5–21.5 cm, 1.3–2.1 times as long as wide, falcate-obovate with a somewhat flattened top and a sharp projection on one side, glabrous to sparsely puberulent on exposed part in bud outside, glabrous inside. Stamens inserted at 0.4–0.5 of tube length from base; fil-
aments 1.5–2.0 mm long; anthers 7.2–9.8 × 0.9–1.9 mm, 4.6–8.6 times as long as wide. Disk a ring, connate at base, 5-lobed, 0.8–1.3 mm high. Ovary slightly semi-inferior, 0.8–1.6 mm high, glabrous; style 17–22 mm long; style head 1.9–3.0 mm long. Fruit of paired pendulous follicles, linear, often strongly curled, 31–40 cm long, 6–7 mm wide, sparsely to densely pubescent. Seeds with a few hairs on one side; 17.0–18.6 × 3–6 mm, coma 3.0–3.2 cm long. Distribution: Thailand and Vietnam but given its known distribution is likely also to occur in Burma, Laos and Cambodia. Habitat: primary and secondary forests, particularly at forest margins, from 100–800 m. Additional material studied: THAILAND: sine loc.: van Beusekom et al 3379 (BKF, L). Chanthaburi: Khao Soi Dao, Sangkhachand 495 (BKF); Pong Nam Ron, Smitinand 3518 (BKF). Chon Buri: Chundaten Falls, Sriracha District, Maxwell s.n. (AAU). Kanchanaburi: Erawan National Park, Parnell et al. s.n. (K, TCD). Nakhon Nayok: Muang Nakhon Nayok, Khao Yai National Park, near Pah Gluai Mai, Maxwell s.n. (A). Nan: Tham Pa Tok near Nan,
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FIGURE 4. Epigynum cochinchinense (Pierre) D. J. Middleton.
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Larsen et al. 46422 (L); Tham Pa Tok, 10 km N of Nan, Larsen et al. 44402 (AAU); Tham Tha Toop Forest Park, Middleton et al. 133 (A, BKF). Phetchaburi: Along road to Tor Thip waterfall, Larsen et al. 45060 (AAU); Kaeng Krachan National Park, ca. 8 km past Kha Sam Yod Ranger Station, Parnell et al. s.n. (K, TCD); Kaeng Krachan National Park, road to Khao Phanoen Thung, Newman et al. 1020 (A, BKF, E); Kaeng Krachan National Park, trail from Ban Krang Ranger Substation., Middleton et al. 928 (A, BKF, E). Prachuap Khiri Khan: Hua Hin District, Kaeng Krachan National Park. Pa La U waterfall., Middleton et al. 1059 (A, BKF, E); Hua Hin District, Kaeng Krachan National Park. Pa La U, trail from dam up mountain., Middleton et al. 2304 (A, BKF); Huay Yang, Put 3190 (K, BM, P); Kui Buri National Park, trail from park headquarters, Middleton et al. 1231 (A, BKF, E); Thap Sakae District, Huay Yang National Park. Bua Sawan waterfall area., Middleton et al. 1349 (A, CMU, BKF, E, K, L). Sa Kaeow: Track to base of Khao Takrup, Middleton 209 (A, AAU, BKF). –Saraburi: Muak Lek, Kerr 9061 (A, ABD, BM, E, K, L); Sam Lahn, Maxwell s.n. (AAU). Sukhothai: Maxwell s.n. (AAU); Soke Praruang Falls, Maxwell s.n. (AAU). Tak: Khao Pha Wo, ca. 70 km W of Tak, Murata et al. T16879 (A, BKF). VIETNAM: Ben Tre: Pierre 4468 (K, NY, P). Khanh Hoa: Km 52 de la route 20, Poilane 21298 (AAU, HN, L, P); route col. no. 20 kil 54, Poilane 19689 (P). This species is fairly common and widespread but rather infrequently collected. It is the most distinctive species in the genus, with its large robust pink-tinged flowers and curled fruits. The pedicel lengths given above could perhaps be slightly misleading. In many flowers the pedicel is quite long, with two empty bracts around the middle. In other flowers these bracts have flowers and there are no further bracts on the shorter pedicels. The description given is the length only above the bracts to the sepal bases. If one counts the pedicels as including the length below any empty bracts, they can be up to 17 mm long. 3. Epigynum graciliflorum (Pierre) Pichon, Bull. Mus. Hist. Nat. Paris, ser. 2, 20: 192. 1948. Fig. 3.
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Basionym: Chonemorpha graciliflora Pitard, Fl. Gen. Indo-Chine 3: 1248. 1933. TYPE: LAOS. Louang Prabang, Clovis Thorel 3443 (Lectotype: P, designated here; Isotype: P). Climber. Branchlets sparsely lenticellate; densely hispid when young, becoming more sparsely so with age. Leaves: petiole 5–6 mm long, hispid; blade 5.2–12.0 × 3.7–7.0 cm, 1.5–1.6 times as long as wide, ovate, apex apiculate, base cordate, long crisped hairs above and beneath, denser on midrib, 11–14 pairs of secondary veins, tertiary venation scalariform. Inflorescence of terminal and axillary dichasial cymes, clustered into dense umbelliform heads, 6.5–8.2 cm long; peduncle 1.0–6.5 cm long, densely tomentose, with a combination of shorter and longer crisped hairs; pedicels 0.0–3.5 mm long, densely tomentose. Sepals 2.2–2.7 × 0.9–1.1 mm, 2.4–2.5 times as long as wide, narrowly ovate, apex acute, tomentose outside, pubescent on inner surface on upper half, colleters in a continuous row at base inside. Corolla tube 12.0–12.7 mm long, 1.7–1.8 mm wide, 4.4–5.5 times as long as sepals, 1.3–1.6 times as long as lobes, pubescent in 5 rows down tube outside, very sparsely puberulent in upper half of tube inside, few hairs in throat; lobes 8–9 × 3.1 mm, 2.6–2.9 times as long as wide, falcate, pubescent outside on part exposed in bud, lobes ciliate, glabrous inside or with few hairs at base of lobes. Stamens inserted at 0.2 of tube length from base; anthers 4.3 × 0.7 mm, 6.1 times as long as wide. Disk of 5 separate lobes, 0.5 mm high. Ovary very slightly semi-inferior, 0.5 mm high, glabrous; style 2.5 mm long; style head 1.9 mm long. Fruit unknown. Distribution: Endemic to Laos. Habitat: unknown. Epigynum graciliflorum is similar to Epigynum auritum but differs in the slightly more densely clustered flowers, the denser indumentum, the continuous row of calycine colleters and the glabrous ovary. It is known only from the type collection from Laos. 4. Epigynum griffithianum Wight, Ic. Pl. 4: t.1308. 1848; Kurz, Fl. Burm. 2: 184. 1877; Hook.f., Fl. Br. Ind. 3: 666. 1882; Turner, Gard. Bull. Sing. 47: 126. 1997; Middleton, Fl.
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Thailand 7: 121. 1999. TYPE: BURMA. Mergui, William Griffith s.n. (Lectotype: K [Herb. Wight], designated here; Isotypes: K [Herb. Hookerianum]). Fig. 5–6. Synonyms: Legouixia amabilis Heurck. & Müll.Arg. in Heurck., Obs. Bot. 146. 1871. TYPE: “INDIA ORIENTALI.” William Griffith 227 (Holotype: AWH, n.v.; Isotypes: G, P). Epigynum maingayi Hook.f., Fl. Brit. Ind. 3: 666. 1882; Turner, Gard. Bull. Sing. 47: 126. 1997. TYPE: MALAYSIA. Malacca, Alexander Carroll Maingay KD 1088 (Lectotype: K, designated here; Isotypes: GH, K). Epigynum curtisii King & Gamble, J. As. Soc. Beng. 74,2: 501. 1907; Turner, Gard. Bull. Sing. 47: 126. 1997. TYPE: MALAYSIA. Penang, Road to Penara Bukit, Charles Curtis 2441 (Holotype: SING; Isotypes: K, SING). Epigynum perakense King & Gamble, J. As. Soc. Beng. 74,2: 500. 1907; Ridley, Fl. Mal. Pen. 2: 366. 1923; Kerr, Fl. Siam.
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En. 2: 471. 1939. TYPE: MALAYSIA. Perak, Larut, King’s Collector 1971 (Lectotype: K, designated here). Argyronerium odoratum Pitard, Fl. Gén. Indo-Chine 3: 1234. 1933. Epigynum odoratum (Pitard) Kerr in Craib, Fl. Siam. Enum. 2: 471. 1939. TYPE: CAMBODIA. Kampot, Kep, Geoffray 484 (Lectotype: P, designated here; Isotypes: HN, P). Climber, reported to 20 m high. Branchlets not lenticellate; sparsely to densely brown pubescent, glabrescent with age. Leaves: petiole 2–15 mm long, glabrous to densely pubescent; blade 1.7–16.2 × 0.9–7.0 cm, 1.5–4.2 times as long as wide, elliptic to slightly obovate, apex acuminate with a sharp tip, rarely apiculate, base cuneate to obtuse, glabrous, with few hairs on midrib or sparsely pubescent all over and more densely on midrib above, pubescent in the secondary vein axils to all over beneath, more rarely with only a few hairs on midrib beneath, 5–9(–11) pairs of secondary veins, tertiary venation laxly reticulate. Inflorescence of terminal and axillary umbelliform
FIGURE 5. Distribution of Epigynum griffithianum Wight.
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FIGURE 6. Epigynum griffithianum Wight.
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cymes, 3.0–6.5 cm long; peduncle 0.2–2.9 cm long, densely brown pubescent; pedicels 1.8–5.0 mm long, densely brown pubescent. Sepals 1.2–3.0 × 0.5–1.2 mm, 1.6–3.1 times as long as wide, narrowly ovate, apex acute to acuminate, densely brown pubescent, colleters absent inside. Corolla white or cream, fragrant; tube slightly twisted or not, 15–27 mm long, 1.7–2.8 mm wide, 6.0–11.4 times as long as sepals, 2.0–4.1 times as long as lobes, densely pubescent outside, glabrous at very base and then densely pubescent toward upper half of tube inside, villous in throat; lobes 4–10 × 3–6 mm, 1.3–2.0 times as long as wide, falcate with a sharp projection on one side, densely pubescent on part of lobes exposed in bud outside, glabrous inside. Stamens inserted at 0.1–0.2 of tube length from base; filaments 0.7–1.0 mm long; anthers 3.1–5.1 × 0.6–1.0 mm, 4.4–6.3 times as long as wide. Disk a ring with a slightly to strongly crenate margin, 0.5–0.6 mm high. Ovary semi-inferior, 0.7–1.2 mm high, glabrous; style 1.9–2.7 mm long; style head 1.5–2.5 mm long. Fruit linear or very slightly torulose, 9.8–33.0 cm long, 3–4 mm wide, densely to very sparsely pubescent or glabrous. Seeds 13.5–31.0 × 2.0–2.7 mm; coma 17–34 mm long. Distribution: this is a widespread species known from northeastern India, Burma, Thailand, Cambodia, Vietnam, and Peninsular Malaysia. Habitat: in primary and secondary forests, and even in plantations, particularly at forest margins, from 60–1220 m. Additional material studied: BURMA: Me rgui, Griffith Herb. Wight s.n. (K), Griffith Herb. Hookerianum s.n. (K). CAMBODIA: Kampot: Kep, Geoffray 484 (P). INDIA: West Bengal: near Calcutta, Helfer s.n. (A). MALAYSIA: Sine loc.: Scortechini s.n. (E, P). Kedah: Gerik, Unknown Collector KL 4703 (KEP); Hutan Simpan Sungkop, Bedong, Teo & Pachiapu 1047 (K, L, SING); Sungai Petani, David 183 (P); Yan, Ridley 5542 (SING). Kelantan: Bukit Baka Machang, Teo & Pachiapu 1150 (L, SING); Gua Ninik, Henderson 19616 (SING). Malacca?: Maingay KD 1088 (GH, K). Negri Sembilan: Sg. Ujong, Linsum Estate, Ridley s.n. (SING). Pahang: 5 miles S of Kuala Lipis, Burkill & Haniff 17182 (SING); Jerantut, Burkill & Haniff 16069 (BO,
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BRI, SING); Karak Forest Reserve, Best 13887 (SING); Kota Tongkat, Evans 13046 (K, SING); Taman Negara, Sg. Tahan, Lata Berkoh, Whitmore FRI 20176 (K, SING); Tekam Forest Reserve, Jerantut, Ng FRI 20752 (A, KEP, SING). Penang: Road to Penara Bukit, Curtis 2441 (K, SING). Perak: S c o rtechini 1474 (BO); Kota, Wray 1946 (K, P); Larut, King’s Collector 1971 (K), 2324 (K), Wray 4267 (SING); near Ulu Kerling, King’s Collector 8734 (A, K). Selangor: Sg. Buloh, Sow & Tachun 16878 (SING), Saw 16879 (KEP). Kuala Lumpur, Ridley s.n. (K, SING). THAILAND: Unknown province: Biserat, Jalor, Gwynne Vaughan 609 (K). Chanthaburi: Khao Sabap, Put 902 (A, E, P); Pliew Falls, Larsen 9736 (AAU), Maxwell s.n. (AAU). Chumphon: Ban Racha Krude, Larsen et al. 43166 (AAU, P). Krabi: Khao Phanom Bencha, foothills of S range, Larsen et al. 43302 (AAU, P). Nakhon Sri Thammarat: Sanan 762 (BKF, K). Narathiwat: Sangkhachand 197 (BKF); Waeng, Sangkhachand 867 (K). Phatthalung: 10 km E of Na Thawi, Larsen et al. 41688 (AAU, P); Plai Wan Waterfall, 25 km SW of Phatthalung, Larsen et al. 41591 (AAU, P). Ranong: Ko Chang, K e rr 16619 (BM, K, P). Si Sa Ket: Dongrak Range, Chong Bat Lak, Kantaralak District, Maxwell s.n. (AAU, L). Songkhla: Khao Nam Kang, Na Thawi, S of Chana, Larsen et al. 42841 (AAU, P). Surin: Phengklai 3518 (BKF). Trang: Khao Soi Dao, K e rr 19139 (A, ABD, BM, E, K, L, P). Yan Ta Khao District, Khao Banthat Mountains, Trail to summit of Phu Pha Mek., Middleton et al. 1934 (A, BKF, E); Yandakhao District, Sai Roong Falls, Maxwell s.n. (A). Yala: Betong, Kerr 7444 (A, ABD, BM, E, K, L, SING), 7900 (BM, K, P). VIETNAM: Nghe An: Pu Mat, Pu Bun, Unknown Collector CH–C 229 (HNU). This species is closest to Epigynum ridleyi, from which it differs in, for the most part, having fewer secondary veins, the generally diff e rent leaf pubescence, and the glabrous ovary. Also the corollas tend to be shorter than in E. ridleyi. 5. Epigynum ridleyi King & Gamble, J. As. Soc. Beng. 74,2: 502. 1907; Turner, Gard. Bull. Sing. 47: 126. 1997. TYPE: MALAYSIA. Selangor, Rawang, Henry Nicolas Ridley 7567 (Holotype: SING; Isotype: K). Fig. 7–8.
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FIGURE 7. Distribution of Epigynum ridleyi King & Gamble.
Synonyms: Epigynum forbesii King & Gamble, J. As. Soc. Beng. 74,2: 503. 1907; Maxwell, Gard. Bull. Sing. 35: 192. 1983; Turner, Gard. Bull. Sing. 45: 35. 1993. TYPE: INDONESIA. Sumatra, near Bijin Telok, Henry Ogg Forbes 3236 (Holotype: K). Epigynum borneense Merr., J. Malayan Branch Roy. Asiat. Soc. 1: 27. 1923. TYPE: MALAYSIA. Sabah, Sandakan, Maximo Ramos 1117 (Lectotype: A, designated here; Isotypes: BO, GH, K, P, US). Climber, reported to 8 m high but probably can grow higher. Branchlets sparsely lenticellate or not, densely brown pubescent, becoming less densely so with age. Leaves: petiole 4–7 mm long, sparsely to densely pubescent; blade 2.3–13.0 × 0.6–5.1 cm, 2.1–6.2 times as long as wide, elliptic, apex acuminate with a sharp tip, base cuneate to rounded, 9–14 pairs of secondary veins, tertiary venation subperpendicular to midrib and/or laxly reticulate. Inflorescence of terminal and axillary umbelliform cymes, 3.8–7.0 cm long; peduncle 0.3–2.2 cm long,
densely brown pubescent; pedicels 1.5–4.7 mm long, densely brown pubescent. Sepals 2–5 × 0.8–1.9 mm, 1.8–3.8 times as long as wide, narrowly ovate, apex acuminate, densely brown pubescent, colleters absent inside. Corolla white, reported on one specimen as yellow in throat; tube 15.5–38.0 mm long, 2.2–3.0 mm wide, 6.2–12.9 times as long as sepals, 2.2–4.5 times as long as lobes, densely pubescent outside, glabrous at very base and then sparsely to densely pubescent toward upper half of tube inside, villous in throat; lobes 6–14 × 3.3–8.1 mm, 1.6–2.6 times as long as wide, falcate with a sharp projection on one side, densely pubescent on part of lobes exposed in bud outside, glabrous inside. Stamens inserted at 0.1–0.2 of tube length from base; filaments 0.6–1.2 mm long; anthers 5.5–6.5 × 0.8–1.0 mm, 6.1–7.0 times as long as wide. Disk a ring with a crenate margin, 0.6–1.1 mm high. Ovary semi-inferior, 0.9–1.3 mm high, sparsely to densely pubescent; style 2.5–3.7 mm long; style head 2.0–3.1 mm long. Fruit linear or slightly torulose, 16.5–30.0 cm long, 3.5–5.5 mm wide, densely to sparsely pubescent. Seeds not seen.
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FIGURE 8. Epigynum ridleyi King & Gamble.
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Distribution: Peninsular Malaysia, Sumatra, and Borneo (Sabah, Sarawak, Kalimantan). Habitat: it has been collected from Dipterocarp forest, from swamp forest, and in seriously disturbed and logged areas, from 0 to 300 m. Additional material studied: INDONESIA: KALIMANTAN: Selatan: Banjarmasin, Motley 1232 (K). Tengah: Barito Ulu, waterfall below J. Banbang, Ridsdale PBU 491 (A, BO, E, K, KEP, L). Timur: Wanariset, Ambriansyah & Arifin AA 315 (L), Van Balgooy & Kessler 5979 (A, K, L). SUMATRA: near Bijin Telok, Forbes 3236 a (K). Bangka: G. Mangkol, Kostermans 628 (BO); Sungei Liat, Teysmann s.n. (BO). Jambi, near Bivak Selemboekoe, Posthumus 793 (BO). MALAYSIA: PENINSULAR: Johor: 28 miles S from Kota Tinggi towards Mersing, Teo & Pachiapu 456 (K, SING); 6th mile Mawai Road, Corner s.n. (SING); Sungai Kayu, Kiah
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SFN 32141 (A, BO, K, SING). Selangor: Ginting Bedai, Ridley 7398 (SING); Kanching, Symington 21096 (KEP, SING); Klang Gates, Hume 7160 (SING); Rawang, Ridley 7567 (K, SING); Ulu Klang, Teo & Pachiapu 399 (K, SING). Terengganu: Kemaman, Bukit Kajang, Corner 30214 (K, SING). BORNEO: Labuan:Lobb s.n. (K). Sabah: Keningau District, Ulu Sg. Tinagalan Forest Reserve, Fidilis & Sumbing SAN 113085 (K, SAN); Nabawan District, Syt. Bena wood logged area, Sg. Maadun, Asik Mantor SAN 118824 (E, L, SAN); Ranau, Ulu Sungai Lakimut, Amin Gambating SAN 110459 (SAN); Sandakan, Ramos 1117 (A, BO, GH, K, P, US). Sarawak: Sine loc.: Beccari 1858 (K, P), Hewitt s.n. (BO, K); Kuching, Haviland 3048 (K, P, SING). SINGAPORE: Nee Soon, Maxwell s.n. (AAU, L, SING), Sinclair 6509 (US). It is most similar to Epigynum griffthianum (see there).
INSUFFICIENTLY KNOWN Echites parviflora Roxb., Fl. Ind. 2: 20. 1832; G. Don, Gen. Syst. 4: 75. 1837. TYPE: Drawing 2465 in the Roxburgh collection in Kew library.
Synonym: Epigynum parviflorum (Roxb.) Hook. f., Fl. Brit. Ind. 3: 666. 1882. It is probably an Anodendron but of uncertain species. Epigynum beccarii K.Schum., Nat. Pflanzenfam. 4, 2: 178. 1895. nom. nud.
EXCLUDED SPECIES Epigynum chinense Merr., Philipp. Journ. Sc. 23: 262. 1923.; Xu, Agric. Uni. Wag. Pap. 88-6: 25. 1988. TYPE: CHINA. Hainan, Yik Tsok Mau, Floyd Alonzo McClure 9710 (Lectotype: A, designated here; Isolectotypes: A, BM, E, NY, P, PNH, UC, US, Z).
Synonyms: Sindechites chinensis (Merr.) Markgr. & Tsiang, Sunyatsenia 3: 152. 1936; Kerr, Fl. Siam. En. 2: 471. 1939; Tsiang & P.T. Li. Fl. Reipubl. Popularis Sin. 63: 170. 1977; Li et al., Fl. China 16: 188. 1995; Middleton, Fl. Thailand 7: 135. 1999. Cleghornia chinensis (Merr.) P.T.Li, Guihaia 4: 192. 1984.
LITERATURE CITED BRUMMITT, R. K., AND C. E. POWELL. 1992. Authors of Plant Names. Royal Botanic Garden, Kew. ENDRESS, M. E., AND P. V. BRUYNS. 2000. A revised classification of the Apocynaceae s.l. Botanical Review 66(1): 1–56. HOLMGREN, P. K., N. H. HOLMGREN, AND L. C. BARNETT. 1990. Index Herbariorum. IAPT, New York. KERR, A. F. G. 1939. Apocynaceae. Flora Siamnensis Enumeratio 2: 422–476. LI, P. T., A. J. M. LEEUWENBERG, AND D. J. MIDDLETON. 1995. Apocynaceae. Z. Y. Wu and P. H. Raven, Eds., Flora of China 16: 143–188. Science Press, Beijing.
MERRILL, E. D. 1923. Diagnoses of Hainan plants, II. Philippine Journal of Science 23: 237–268. MIDDLETON, D. J. 1994. A new combination in Epigynum (Apocynaceae). Kew Bulletin 49: 304. ––––––. 1999. Apocynaceae. In T. SANTISUK AND K. LARSEN, EDS., Flora of Thailand 7: 1–153. Royal Forest Department, Bangkok. PITARD, J. 1933. Apocynacées. Flore Génerale de l’Indo-Chine 3: 1087–1262. XU, Z. 1988. A revision of Cleghornia Wight, Sindechites Oliv. and Epigynum chinense Merr. (Apocynaceae). Agricultural University of Wageningen Papers 88-6: 1–35.
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INDEX OF EXSICCATAE The number in parentheses after each collection refers to the number given to each species above. Only those specimens with a clearly identified collector and collection number are listed: Ambriansyah & Arifin AA 315 (5); Amin Gambating SAN110459 (5); Asik Mantor SAN118824 (5). Beccari 1858 (5); Best 13887 (4); Boonkerd 50 (1); Burkill & Haniff 16069 (4), 17182 (4). Corner 30214 (5); Curtis 838 (1); Curtis 2441 (4). David 183 (4); Din 293 (1). Evans 13046 (4). Fidilis & Sumbing SAN113085 (5); Forbes 3236 a (5); Forrest 29521 (1). Geesink & Santisuk 5410 (1); Geesink et al. 7257 (1), 8239 (1); Geoffray 484 (4); Gwynne Vaughan 609 (4). Haniff & Nur 3893 (1); Haviland 3048 (5); Henderson 19616 (4); Henry 12136 (1), 12761 (1), 12852 (1); Hewitt s.n. (5); Hume 7160 (5). Keenan et al. 1597 (1); Kermode 17371 (1); Kerr 7444 (4), 7900 (4), 9061 (2), 16619 (4), 19139 (4), 21648 (1); Kiah SFN 32141 (5); King’s Collector 1971 (4), 2324 (4), 5194 (1), 8734 (4); Koelz 25169 (1); Kostermans 628 (5). Larsen 9736 (4); Larsen et al. 41591 (4), 41688 (4), 42841 (4), 43166 (4), 43170 (1), 43302 (4), 44402 (2), 45060 (2), 46422 (2). Maingay KD 1088 (4); Mann 167 (1); Maxwell 02-337 (2), 71-676 (2), 73-423 (4), 73-460 (2), 74-492 (1), 74-70 (2), 74-824 (2), 75-523 (1), 76-233 (1), 76-570 (4), 82-148 (5), 87-437 (4), 93-606 (1); Middleton 209 (2); Middleton & Lamxay 260 (1); Middleton et al. 133 (2), 928 (2), 1059 (2), 1073 (1), 1231 (2), 1349 (2), 1457 (1), 1934 (4), 2304 (2), 2581 (1), 2602 (1), 2781 (1); Motley 1232 (5); Murata et al. T-16879 (2). Nakkan 293 (1); Newman et al. 1020 (2); Ng FRI 20752 (4). Parnell et al. 95-506 (2), 95-652 (2); Phengklai 3518 (4); Phusomsaeng 249 (1); Pierre 4468 (2); Poilane 19689 (2), 21298 (2); Posthumus 793 (5); Put 878 (1), 902 (4), 1244 (1), 1255 (1), 1620 (1), 3190 (2). Rabil 172 (1); Ramos 1117 (5); Ridley 5542 (4), 7398 (5), 7567 (5); Ridsdale PBU 491 (5). Sanan 762 (4); Sangkhachand 16 (1), 197 (4), 495 (2), 714 (1), 867 (4); Saw 16879 (4); Scortechini 1474 (4); Sinclair 6509 (5); Smitinand 3384 (1), 3518 (2), 3700 (1); Sow Tachun 16878 (4); Suvanakoses 348 (1); Symington 21096 (5). Teo & Pachiapu 399 (5), 456 (5), 1047 (4), 1150 (4); Teysmann s.n. (5); Thorel 3443 (3). Van Balgooy & Kessler 5979 (5); Van Beusekom et al. 737 (1), 3379 (2). Wang 75895 (1), 77048 (1), 79185 (1), 79564 (1), 81055 (1); Whitmore FRI 20176 (4); Wray 1802 (4), 1946 (4), 4267 (4).
INDEX Names in roman script are currently recognized species; those in italics are synonyms. KEY: (au) = Epigynum auritum, (co) = Epigynum cochinchinense, (gf) = Epigynum graciliflorum, (gr) = Epigynum griffithianum, (ri) = Epigynum ridleyi, (ik) = insufficiently known, (ex) = excluded species. Argyronerium odoratum Pitard (gr) Chonemorpha graciliflora Pitard (gf) Cleghornia chinensis (Merr.) P. T. Li (ex) Echites parviflora Roxb. (ik) Epigynum auritum (C.K.Schneid.) Tsiang & Li (au) Epigynum beccarii K. Schum. (ik) Epigynum borneense Merr. (ri) Epigynum chinense Merr. (ex) Epigynum cochinchinense (Pierre) D. J. Middleton (co) Epigynum curtisii King & Gamble (gr) Epigynum forbesii King & Gamble (ri) Epigynum graciliflorum (Pierre) Pichon (gf) Epigynum griffithianum Wight (gr)
Epigynum lachnocarpum Pichon (au) Epigynum maingayi Hook. f. (gr) Epigynum odoratum (Pitard) Kerr (gr) Epigynum parviflorum Hook. f. (ik) Epigynum perakense King & Gamble (gr) Epigynum ridleyi King & Gamble (ri) Nouettea cochinchinensis Pierre (co) Sindechites chinensis (Merr.) Markgr. & Tsiang (ex) Trachelospermum auritum C. K. Schneid. (au) Trachelospermum curtisii King & Gamble (au)
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