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BLUMEA 36 109-121 (1991) taxonomic revision A of the Asiatic genus Pentasacme (Asclepiadaceae) M.A. Rahman Department of Plant & Soil & C.C. Science, University Aberdeen Wilcock of Aberdeen, St. Machar Drive, 2UD, Scotland, U.K. AB9 Summary A four complete revision species. the variation in ognized in the Wall. genus Pentasacme of the Asiatic and distribution,ecology, The features within diagnostic one genus, Wight ex comparative morphology of the genus of which is new, and one is shown species is the given with genus is reduced to a to the key discussed Two sections illustration. by are are and rec- synonymy. Introduction The rheophytic genus Pentasacme is characterized borne on sides of the axis alternating calyx lobes, and beaked pollinia ly Asiatic genus consisting the Khasia Hills of India, through Thailand Wallich with Its (1834). transferred to species question a the genus from the genus in the genus, mark Cynanchum by western Spiladocorys, western Peninsula. The genus from is feminine, Sylhet (Bangladesh) not neuter species, as Hong Kong, Himalaya (1914-1915) described and the species, and Hooker f. and Burma. one new species, most recent Nepal. Mazzetti in 1924, is by never Van Steenis tentative. esquirolii, a been name P. was P. synonym subject (1981), to was of were established by Contr. a Long, (1853) already been added third a noted the presence of described Malay Penins. a monotypic (1923). MacGregor shanense), from Shan Grierson & by state a Another Leveille & Smith (1914) of Burma, Long (1982) used for Chinese elements by from Handel- stauntonii (Decne.) Hand.-Mazz. The taxonomic revision (Rahman, in the absence of & which he himself later treated as pulcherrima by Cynanchum Bot. India (Grierson also described from China brachyantha, to a complete- and ?P. stauntonii, de- (1883) Heterostemma. P. shanensis (= P. addition The treated Ridley (1893) Malay Peninsula, and has been transferred Bhutan and genus has P. Wight's glaucescens i in Fl. synonym of Wallich's Pentasacme caudatum non -Pentasacme in traditionally ?P. originally was Decaisne (1844) from China have by a Himalaya, Nepal, Bhutan, Handel-Mazzetti (1936). Bentham from the petioles, glandular Burma, southern China, and Hong Kong Bangladesh, Malay species the presence of lateral cymes species, although previously eight species distributed from the two non-Pentasacme scribed with to two gender 1982). The the to by the nodes between the with medianattachment to caudicles. It is of four recognized (Airy Shaw, 1973), at revision species 1990a) and, distinction is as noted more or less 110 BLUMEA VOL. No. 36, 1991 1, A critical examination of all the available materialof Pentasacme, in ABD, BM, is four represented by only belongs the to that describing be species and that the material of P. species, as from P. caudatum Wall, Wight. now enable to possess championi Bentham mentioned that "it may Hong Kong, well with those described by Decaisne for materials than I ex of P. caudatum, but the floral characters do variety a mere species same including types, K, K- W, L, and P showed that the genus CAL, CGE, DACB, E, to possibly prove to sufficiently agree of the variations of the judge while unite them, without better me to venture to me not Benth. (1853), Sylhet plant." Some as to long from China and specimens their leaf and habitat morphology indicates that have they scales in all these the strikingly similar to that specimens correct name in family tropical and that these pendulous (not erect) pollinia attached are for these the to do specimens being investigated Asia (Rahman & Wilcock, not be- specimens should specimens In addition, the coronal the staminal column and to is Asclepiadeae. Benth. championi species, this genus, however. Close examination of the flowers of these be transferred from the Marsdenieae la. The identified as P. Hongkong, are 1989a, 1989b, not to part of as the corol- our study of 1990). PENTASACME Pentasacme Fl. Wall, species: Wight, Contr. ex Penins. Malay or branching Trans. Linn. dular within to with the the at long right ser. or at glands II, 3 present the nodes, (1893) on Fl. Brit India 4 Sinic. narrow 63 (1883) 28; Ridley, (1977) 413. — herbs. Stems at the junction of bracts mostly slender, ovate-elliptic, acuminate, petiole southern China acute, glabrous, glan- Corolla campanulate; tube very short; lobes ovate-acumi- apices Corona or oblong-lanceolate, cup-shaped or anthers tip obtuse, glabrous, overlapping reduced to scales, adnate to the corolla fleshy, tips membranous; stigma sessile, slender, terete, glabrous. Seeds with a - The genus is distributed throughout the western and Bhutan, and the Khasia Hills of India, including Hong Kong, and through Himalayan region Bangladesh, Burma, Thailand to the Malay and Peninsula - The genus Pentasacme has a unique ecology within the family Asclepi- adaceae. The species inhabit damp rock ledges in waterfalls, moist banks, and genus tuft of 6). Ecology and Cymes Flowers white, small glabrous. tip and lamina. glabrous; peduncles end. including Nepal beds, Type 332. alternate sides of the axis, mm); pedicels filiform; angles. in bud. Distribution 3 & f., beaked. Pollinia beaked, with median attachment to caudicles, erect, soli- at one (figs. Hook, Reip. Pop. Wight. tary in each anther loculus. Follicles hairs Fl. rheophytic, perennial tube. Staminal column very short; conical Li, lobes ovate-acuminate or ovate-lanceolate, large. Calyx nate than 2 (less ex (1834) 60; & from the base. Leaves linear-lanceolate or umbelliform, solitary very short Soc. sometimes pendulous, nearly base cuneate, subsessile, or BoL India (1923) 383; Tsiang Pentasacme caudatum Wall, Spiladocorys Ridley, Erect 2 are impose rheophytic. The marked effects peculiar on environmental conditions leaf form. The leaves are occupied distinctly stream by this thin and, when M.A. Rahman dry, elliptic ovate-elliptic or moist conditions, while the in perennials erect Revision stream leaves with species beds or found are long Stems — Pentasacme sparsely hairy are species in very linear or linear-lanceolate leaves live as MORPHOLOGY have species stems and branched long cm stems, but those of P. shanensis glabrous the nodes, while the at within the genus, hanging perennials occur as erect or 111 Pentasacme slender and about 30-60 stems are from the base. Two Asiatic waterfalls. COMPARATIVE nearly of Linear and broader leaved forms fragile. very with & C.C. Wilcock: of P. pulcherrima are pubescent all over. Leaves sis they There — long, are distinct are two narrow types cies, P. pulcherrima and P. wallichii, they Inflorescences — of leaves. In P. caudatum and P. shanen1 & linear-lanceolate (figs. or Inflorescences are whereas in 2) interpetiolar, a specific other spe- two 4& elliptic-ovate (figs. are 5). character of asclepiads (Nolan, 1969), and borne on alternate sides of the axis. Nolan showed that the apical dome of and a functional apex unequal accounting associated with of the bifurcates Asclepias syriaca continuing stem not to of the the gives the impression of are inflorescence from being short than 1 cm) one be or are apex slightly are not Nolan, the branching to sympodial systems of and in the fresh clearly non-axillary alternate. Pedicels (less to These bifurcations according expected monopodial The inflorescence, therefore, is Peduncles of P. caudatum to an peduncle. and therefore, organs conform rise giving the other. The bifurcation appears displacement subtending does angiosperms. material for the on very short (1-2 mm). whereas they are longer (1-3 cm) in the other three species. Each peduncle of P. pulcherrima produces 3-5 pedicels, but these Flowers — Flowers vation in bud. 1) than 5 in the other three are more Calyx are 5-merous and lobes free, species. radially symmetrical glandular within at the with imbricate aesti- angles, and of two ovate-acuminate in P. caudatum, and 2) ovate-lanceolate in the other three Corolla white with P. caudatum, and Corona types of — types of lobes: ovate-acuminate with long two The corona are corona corona is is corollin in origin (fig. 2c) variable in largely and P. wallichii fleshy & structure. and in 5). Two cup-shaped the base of the corolla tube, but in the other spe- at represented only by five separate scales borne the lobes. The coronal scales in P. caudatum pulcherrima and apices narrow species (figs. 1, 2,4 present in Pentasacme. In P. shanensis it is with five rounded lobes cies the in the three other oblong-lanceolate types: species. they are are on denticulate (fig. with either obtuse (fig. 4c) the tube between whereas in P. lc), or acute (fig. 5b) tips. Pollinia to — Pollinia are ovate to the caudicles. The variation in ment 5b). of the pollinia to Pollinia with beaks ovate-lanceolate, beaked with median attachment shape the translator at the istic of the genus (Rahman, tips of the arms, pollinia are and shown in corpuscles, and median attachment 1990b). and the attach- figures (figs, le, 2b, to caudicles are 4d & character- 112 BLUMEA VOL. INFRAGENERIC Within the Asclepiadaceae, distinct groups; the species with while describing subdivision of the genus into inclined large two two based on sections. section. The two are in placed a new Section Leaves linear lobes less than Foliis longis ellipticis 10 quam Leaves lobes mm long. — vel mm. the we are section Pentasacme P. pulcherrima readily separable and and elliptic more Typus or Leaves or lobes more Stems long. elliptic Pentasacme latioribus sectionis: long. — more than 15 THE sect. Floribus mm. grandibus; lobes 4-5 than 10 mm mm broad. Flowers mm large; corolla Wight. SPECIES mm broad. Flowers small; corolla 2 elliptic, more than 15 mm broad. Flowers large; corolla 3 long Pedicels of cymes 0.4-0.8 long. Iobis corollae long or ovate sparsely hairy Wight. Wight. cm long. Corolla lobes 6-8 Coronal scales 5, denticulate Stems ex nov. Pentasacme wallichii Type species: TO 15 wallichii linear-lanceolate, less than 13 glabrous. broad. Flowers small; corolla mm Pentasacme caudatum Wall, quam Pentasacme elliptic-ovate, than 10 mm Leaves linear 1. Wight Wallichiana Rahman, elliptico-ovatis, — lobes less than 10 mm b. are Long already suggested species, sections Wall. ex Type species: KEY 2a. other linear-lanceolate, less than 13 or 10 Section 2. b. readily group and described below: PENTASACME la. two one leaf and flower size, and The residual two are in this genus fall in of Pentasacme, have includes P. caudatum and P. shanensis. The P. wallichii, in the Tribe Marsdenieaeof sub- flowers in the other group. Grierson & groups divide the genus into to placed leaves and small flowers in narrow species a new 1991 1, species recognized with broad leaves and species (1982), the No. CLASSIFICATION Pentasacme is The four family Asclepiadoideae. 36, at 1. P. caudatum the nodes. Pedicels of cymes 2-3 Corona mm cup-shaped cm long. Corolla with 5 rounded lobes 2. P. shanensis 3a. Stems b. Stems pubescent. Corolla lobes 20-30 mm long. tips Coronal scales with obtuse 3. glabrous. Corolla lobes 10-13 mm long. P. pulcherrima Coronal scales with acute tips 4. P. wallichii M.A. Rahman Fig. 1. Pentasacme coronal scales; d. caudatum fruits; e. & C.C. Wilcock: Wall, ex Wight, a. Revision Habit; b. pollinarium((Lamont 464, BM). of Asiatic gynostegium Pentasacme and sepals; c. 113 corolla with BLUMEA 114 Fig. ing 2. Pentasacme exserted stigma, shanensis MacGregor corolla and coronal VOL. 36, & Smith, scales, and a. No. 1, Habit; calyx lobes 1991 b. pollinarium; c. open ((Parkinson 1184, K). flower show- M. A. Rahman & C. C. Wilcock: Revision Asiatic of Pentasacme 115 TAXONOMY Section 1. Pentasacme caudatum Wall. Pentasacme 3, Siam. 2 14, caudatum f„ Hook. 627; 1 Wall, ex Wight, Fl. Brit. India 4 96. — Wight ex Contr. Bot. India List Burma championi Benth., J. Sinic. Reip. Pop. Waterfall, Champion A small, erect Trans. Linn. Soc. 2885 63 192 BoL Kew 415. (1977) 5 acute. ly tuse Calyx mm margin denticulate, Pollinia ovate. Gynostegium - (BM). 20207 CHINA. — (BM), Griffith 2885 Sritamarat, ma 192 162 Chantaburi, Ram, erect 169. 3 (1974) 264; Leaves almost glabrous bracts 2x1 ses- both sides, on ovate- mm, above with filiform long (6-8 mm) broad- fleshy, than translator longer single, slender, 5-7.5 Stigma arms. cm Seeds long. both sides. BANGLADESH. Asclep. (K); Haw the to Bangladesh, Burma, Malay same locality, 75a & 75c Peninsula m. southern (fig. 3). Flowering 3839 and 8555 HONG KONG. — Lamont 464 s.n. 6361 (BM); (BM), — fruiting be- Cherra Shan, Mount same alt. 4000 punji, Perak, Sira Rimau, 1989, Yapp (BM); Dan Shah Fan 1344 Maan (BM). locality, Champon, Kerr Kiah 17666 INDIA. — Clarke ft, 16.6.1952, (ABD); (L), Ts'uen, C.B. Econ. Prod. 538 8234A no. Hka, Kaul- Cough, Happy Valley Hance 720 (CAL); Behover, Reporter on MacGregor MacGregor & Smith, Rec. Type: Burma, mm Chem Herb. Cat. BURMA. Tara Walter Ind. 11108 Waterfall 31703 Hill, (BM). — Nakawn (ABD); (K). & Smith — Fig. Bot. Surv. India 6 Southern Shan herb. Stems slender, short, less than 2 — (BM, CGE, K, L); (L); Wallich attached, (BM). Krat, Takum. Put 2892 Smith 597 not (E, isotype). McClure T.T. (CCE),Rupchand Biswas Sylhet, locality no. 72172 ft. J.D.H. & shanensis shanensis (1961) An on Thailand MALAY PENINSULA. Pentasacme Pentasacme Ma- ovate-acuminate,. Corolla lobes (BM); Pahang, Sungai Teku, 20.11.68,Mohmd. Kao Type: the staminal column. Anthers with ob- narrow, Hainan, Canton, (L). Assam, Notneng, THAILAND. 2. mm, narrow exceeding convex and Wallich alt. 2000-4000 — — September. (BM, K); Po-ting, (BM, CAL). Fl. Bangladesh Gough, Happy Valley long. cm, long, capillary; 1 x From the Khasia Hills of India, - Waterfall, Champion Ridley mm 0.8-1 x Common in forest streams, alt. 0-1300 June and Mt, never Corpuscles Representative specimens: 30287 4-8 lobes 2-3 Hong Kong through (BM, CGE, K, K-W) 5562 30-50 cm glabrous, almost sessile. Follicle oblong spathulate, Ecology Khasia Fl. Hainanica Mount the corolla tube between the lobes, Distribution 355 Li, base, China and back & Type: Hong Kong, at ovate or Fung t. 63. to c. (1844) (K-W, holo; E, K, iso). (1893) 322, wide tips. tween Asiatic. Soc. 75 no. Prodr. 8 DC., (1923) 383; Kerr, Coronal scales 5, adnate ovate, obtuse. 2 J. 2 (BM). (1853) 54; Tsiang — herb. Stem slender, ovate-acuminate, apices. Penins. (K). margin hairy; pedicels lanceolate, tips Bot. 3 II, ser. sile, linear-lanceolate, tip acute, lamina 6-9 midrib and Malay (1961) 169; Rahman, lay Peninsula, Perak, waterfall, 1892, Ridley Fl. Decne. in (1834) 60; Fl. Type: Bangladesh, Sylhet, Wallich, Asclep. Spiladocorys angustifolia Ridley, Pentasacme 1. Fig. — (1883) 28; Ridley, (1951) 19; Hundley, (1988) Pantasacme (1914) 101; Hundley, State, MacGregor glabrous, sparsely hairy long, pubescent. 2. Leaves linear, tip at 535 List Bur- (E, iso). the nodes. Petioles very acute, lamina 4.5-12 x 0.3- 116 Fig. BLUMEA 3. Geographic distribution of Pentasacme Gregor & Smith 0.8 glabrous cm, nate except for the as the translator anthers. Follicles Distribution Ecology ing - - not 4-5 1184 Corona entire, Pentasacme alt. 3500 streams mm. mm, Corolla ad- cup-shaped, fleshy, never the exceeding Corpuscles oblong-lanceolate, mm long. Stigma as exserted above of mountains, from 0-1100 m alt. Flower- BURMA. Southern 6239 State, MacGregor 48 535 3000 water, alt. (K); Northern (E); May Myo Dist., ft, 2.7.1925, Parkinson Shan State, Niansam Talis, (K). Grierson & Type: Bhutan, Samdrup Shan (K); Laughing ft, Aug. 1913, English 5846 pulcherrima pulcherrima bracts 2x1 1.5-2 August. ft, 6.7.1912, Lace Pentasacme x fleshy, rounded, Pollinia globose Section 3. long; cm (fig. 3). ft, 28.6.1913, Lace (K); Mardacey, 1800 mm. Staminal column c. 2.5 Common in forest alt. 3500 2-3 seen. Representative specimens: alt. and P. shanensis Mac- Wight (■) lobes ovate-lanceolate, 2-3 1.5-2 x ovate. arms. Burma between June and Plateau, 1991 1, caudatum Wall. ex the base of the corolla tube; lobes 5, staminal column. Anthers long No. margins; pedicels Calyx acute. oblong-lanceolate, to 36, (O). ovate-lanceolate, tips lobes VOL. Wallichiana Grierson & Long, Jonkhar to Notes Long Roy. Deothang road, — Fig. 4. Bot. Gard. Edinb. Grierson & 40 (1982) 123, fig. 3. Long 2137 (E, holo; K, iso). — & C.C. Wilcock: M. A. Rahman Fig. c. 4. Pentasacme corolla with An erect pulcherrima coronal herb. Stems acuminate, lamina 6-13 3-5-flowered; pedicels tuse. Calyx 20-30 x d. scales, Grierson 2-4 1-2 cm, cm small, oblong, tip mm. long; anthers. Follicles not arms. seen. & bracts 1.5 x 2 broadly Gynostegium Asiatic Habit; x mm. to gynostegium b. 117 Pentasacme and calyx lobes; Long 2137, E). mm long. and midrib Coronal scales adnate obtuse. Anthers shorter than translator a. of Petioles 3-7 margin lobes ovate-lanceolate, 2.5 2.5-3.5 Long. pollinarium( Grierson pubescent. x & Revision 1 Leaves pubescent mm, ovate-elliptic, beneath. Cymes ovate-lanceolate, tips ob- Corolla lobes oblong-lanceolate, the corolla tube between the lobes, acute. Pollinia sessile. ovate. Corpuscles Stigma conical, ovate, concealed by BLUMEA 118 Fig. 5. Pentasacme stegium; c. wallichii Wight, a. VOL. Habit; b. pollinarium (Stainton 5476, E). 36, No. corolla cut 1, 1991 open to reveal coronal scales and gyno- M.A. Rahman Fig. 6. C.C. Wilcock: & distribution of Pentasacme Geographic Revision Asiatic of Pentasacme Grierson & pulcherrima Long (O) 119 and P. wallichii Wight (■). Distribution Ecology jungle - - Bhutan and Grierson & Jonkhar Samdrup near Representative specimens: Jonkhar, alt. 550 m, alL 230 m, 4. Pentasacme Pentasacme Williams Wight, & Icon. Select. Plant. Fir. PI. Samdrup & Stainton Wight Nepal 5 (1846) (1982) 87; Rahman, J. A small pendulous herb, acute, erect or Asclep. Petioles 2-4 lamina 5-8 x mm 1.5-2.5 few-flowered; pedicels 1-1.5 Calyx cm, cm 10-13 lobes, small, x 2-3 never mm. x 2137 Hook, f., Decne. Fl. Brit. — 8 km September. above Samdrup NEPAL. Chula Chuli, Soc. in DC., India Prodr. 8 4 Bangladesh 14, (1844) 627; (1883) 28; 2 (1988) De- Hara, Enum 97. Type: — (E). cm Leaves slightly ovate-elliptic, acuminate, tip nerves bracts 2x1 mm. Stems and branches long. mm, Corolla Coronal scales adnate exceeding Deothang road, (E, holo; K, iso). (1834) 60; midrib and 2 shaded steep bank in hot 5. 40-60 long. to a between June and (BM, K). Asiatic 74 long; lobes ovate-lanceolate, 2.5 ceolate, no. Flowering Fig. 87; t. Wallich glabrous. 8499 from species Jonkhar Long — Bangladesh, Sylhet, stout, alt. m Contr. Bot. India lessert. 3 550 Grierson wallichii wallichii collected this at BHUTAN. 21.6.1979, 15.9.1967, Nepal (fig. 6). Long the staminal column, pubescent lobes sub- Cymes linear-lanceolate, tips deeply lobed; to beneath. acute. oblong-lan- the corolla tube between the tip acute. Anthers ovate-acumi- 120 BLUMEA Polliniaovate-lanceolate. nate. minal column about 2 shaped, c. 5.2 cm Distribution mm 36, Corpuscles long, tapering at the No. 1991 1, than translator longer narrow, arms. exserted above anthers. Follicles long. Stigma Throughout - VOL. both ends, slightly stout, smooth. of India, Himalayas western Sta- spindle- and Nepal Bangladesh (fig. 6). Ecology - and Flowering In clumps or Wallich locality not Reid s.n. s.n. (K); — Khar locality W NEPAL. Gad, 8463 (E). Bazar, Bheri (K). alt. 6500 Sylhet, locality INDIA. Western — (E); east Kapkot, W 21342 glaucescens (Decne.) cescens attached, of alt.3500 5476 Stainton Kumaun: 24687 no. & 74 Bengal, near alt. Garua, Winter-Bottom et al. 239 272 (CAL). (BM, E); (E); Barpak, 16.10.88, Stainton TAXA (1844) 627 (1936) 994. = E Sarju Valley, (CAL, K); Stainton Sin. 7 Pentasacme stauntonii Decne., I.e. water. Asclep. (K); Winter-Bottom Decne. in DC., Prodr. 8 Hand.-Mazz., Symb. s.n. ft, Strachey & Kutharpekot, 27.4.1954, ft, 28.6.1966, Wallich Wallich Himalaya, (E), Strachey EXCLUDED Pentasacme not (E), Inayet (Duthie's collector) Stewart Khola, ledges by running August. (CGE, E, K, K-W); Sylhet, Bilkasar 2319 Bim Valley, 8235 3147 attached, not Nepal, no. 3774 ft, 14.8.84, Duthie (E); Jaldi Cat. attached, Griffith 4000-5000 and May BANGLADESH. Representative specimens: (E, holo; K, iso), from rocks of cliff hanging between fruiting = Cynanchum glau- stauntonii (Decne.) Cynanchum Hand.- Mazz., I.e. 7 (1936) 996. Pentasacme esquirolii Ldveild, Fl. Kouy-Tcheon Pentasacme (1924) brachyantha 168 = (1914-15) 3 (1936) esquiroli (Levi.) Tsiang, Sunyatsenia 43 = Heterostemma 189. Hand.-Mazz., Anz. Akad. Wiss. Wien, Math.-Nat. 61 stauntonii (Decne.) Cynanchum Hand.-Mazz., I.e. (1936). ACKNOWLEDGEMENTS We thank Professor the manuscript lowing W, L, and herbaria for and P. The D.M. offering putting project Moore valuable (University their material was funded of suggestions. by at our Reading) and Dr. We are most disposal: ABD, A.R. grateful to BM. for Berg going through the Directors of the fol- CAL, CGE, DACB, E, K, the British Council for which we are gratefully K- indebted. REFERENCES AIRY SHAW, H.K. 1973. J.C. Willis, A Dictionary of flowering plants and ferns, ed. 8. University Press, Cambridge. GRIERSON, A.J.C., worthy plants & D.G. LONG collected in 1982. Notes Bhutan in relating 1979. Notes to the Flora Roy. of Bhutan: V. New Bot. Gard. Edinb. 40: and note- 123-125 (Ascle- piadaceae). NOLAN, J.R. RAHMAN, of 1967. Bifurcation of the stem M.A. 1990a. Taxonomic Aberdeen, Scotland. studies apex in the in Asclepias syriaca. Amer. J. Bot. family Asclepiadaceae. Ph.D. 56: 603-609. Thesis. University M. A. RAHMAN, M.A. The Plant Rahman 1990b. Pollinaria Diversity RAHMAN, M.A., & C.C. Wilcock: of Malesia: of some Revision Asian tropical 1989a Notes on Pentasacme Asclepiadaceae. 83-92. Kluwer Academic & C.C. WILCOCK. Asiatic of Publishers, tropical Asian In: 121 P. Baas et al. (eds.), Dordrecht. Asclepiadaceae. Blumea 34: 99-101. RAHMAN, M.A., Tax. Bot. 13: RAHMAN, M.A., & C.C. WILCOCK. 1989b. Notes on tropical Asian Asclepiadaceae II. J. Econ. 181-185. & C.C. WILCOCK. 1990. Notes on tropical Asian Asclepiadaceae III. Blumea 34: 519-520. STEENIS, C.G.G.J. VAN. 1981. Noordhoff, Alphen a/d Rijn. Asclepiadaceae. Rheophytes of the world: 198-207. Sijthoff &