REINWARDTIA Published by Herbarium Bogoriense, Kebun Raya Indonesia Volume 5, Part 4, p.p. 481-508 (I960) A MONOGRAPH OP THE GENUS NEESIA * Blume (Bombacaceae) SOEPADMO ** SUMMARY 1) Eight species are described: N. altissima, synandra, glabra, koster- mansiana, malayana, pivrpurascens, piluliflora and strigosa. 2) N. kostermansiana is a species new to science. 3) N. glabra and synandra, formerly included in N. altissima are rein- stated as distinct species. 4) The area of distribution of the genus covers Lower Siam, the Malay Peninsula, Sumatra, Java and Borneo, with Borneo as centre. ACKNOWLEDGEMENTS I wish to express my thanks to the Directors/Keepers of the Herbaria of Bogor, Kepong, Kew, Kuching, Leiden, Paris and Singapore, for their cooperation in lending me material for my study. I am very grateful to Prof. Dr. C. G. G. J. Van Steenis (Leiden) and Dr. A. J. G. H. Kostermans (Bogor), who have taken the trouble to prepare the Latin diagnoses, to go through the MSS and who have suggested many emendations and corrections. I have to thank Messrs. Soekirno, Damhuri and Moh. Anwar for pre- paring the drawings. INTRODUCTION The generic name Neesia was founded by Blume in 1828 (Fl. Jav., l.c.), as a new name for the genus Esenbeckia which he had described in 1825 but which was a later homonym; its single species was Ar. altissima (Bl.) Bl. In 1874 Masters added N. synandra from Malacca, and in 1875 a third species, N. strigosa from Borneo. Beccari added three new species from Borneo: N. ambigua, glabra, and purpurascens in 1889, and segregated the specimen P.B. 2037 from N. stri- gosa as N. piluliflora. •Name d after Th. Fr. L. Nees von Esenbeck (1787—1837). ** Assistant Botanist, Herbarium Bogoriense, Bogor. — 481 — REINWARDTIA [VOL. 5 Bakhuizen v.d. Brink Sr. (1924) described one new species, IV. malayana, from Sumatra and the Malay Peninsula; he reduced IV. synandra Mast., N. ambigua Becc. and IV. glabra Becc. to the synonymy of IV. altissima Bl. Furtado (1929) reinstated IV. synandra as a proper species. The genus Neesia is closely allied to Coelostegia, Durio and Kosterman- sia. The alliance of these four genera has been discussed by Soegeng (in Reinwardtia 5 (3): 271—272. 1960). Kostermans in his monograph of Durio (in Communication, Forest Research Institute, Bogor 62: 2. 1958 and in Rein- wardtia 4(3): 361. 1959) suggested the desirability of combining Neesia and Coelostegia, as Neesia differs only from Coelostegia by the presence of prurient hairs in its fruit-valves. Soegeng, on the contrary, advocates to keep the genera separate, until more material should be available and I believe too, that this is the best policy. The differential characters of Neesia and Coelostegia are: Neesia 1. Leaves with distinct, parallel, secon- dary nerves; lower surface glabrous or covered by stellate hairs or rarely sparsely covered by minute long-fim- briate scales. 2. Epicalyx completely enveloping the bud. 3. Calyx lobes neither saccate nor in- duplicate. 4. Corolla hypogynous. 5. Filament topped by one two-celled anther. 6. Ovary superior, covered by hirsute, stellate hairs. 7. Inside of the fruit-valves densely covered by brownish, hirsute, pru- rient hairs. Coelostegia 1. Leaves without distinct, parallel se- condary nerves; lower surface cover- ed by scales. 2. Epicalyx reduced, subtending the calyx. 3. Calyx lobes induplicate, saccate. 4. Corolla subperigynous. 5. Filament topped by three, one-celled anthers. 6. Ovary partly embedded in the recept- acle, covered by peltate scales. 7. Inside of the fruit-valves glabrous. From the specimens examined, I am convinced that each filament is topped by one anther, and that the anthers are two-celled, not one-celled as assumed by Bentham & Hooker f. and Baillon. The ovules are biseriate, not uniseriate as contended by Endlicher and Miquel; The indumentum of the young branchlets and leaves is very variable, it may consist of simple or of stellate hairs, or of fimbriate to long-fimbriate scales. It is, therefore, not advisable to use this character to distinguish between the young specimens, as both types of hairs and scales are often found on the same specimen. In old leaves, however, the indumentum of 196 J] SOEPADMO: A Monograph of the Genus Nccsia 483 the lower surface becomes rather important, as this character is constant in some species. Valuable characters for specific delimitation are the shape of fruit and calyx after anthesis. The name Neesia Blume is conserved against Neesia Sprengel (Com- positae, 1818). N E E S I A Blume (nomen. gen. conserv.) Neesia Blume, PL Java 1: vii, in nota, 1828; in Nov. Act. Caes. Leop. Car. 17 ( l a ) : 75—84. 1835; Endlicher, Gen. PI. 990. 1840; Griffith, Not. PL As (4): 513. 1854; Miquel, PI. Ind. batav. 1(2)2: 206. 1859; Bentham & Hooker f., Gen. PL 1: 213. 1862; Baillon, Hist. PL 4: 159. 1872; Masters in Hooker f., Fl. Brit. Ind. 1: 352. 1874; in 3. Linn. Soc. Bot. 15: 503. 1875; Beccari, Malesia 3: 259. 1889; Boerlage, Handl. Fl. Ned. Ind. 1: 119. 1890; King in J. As. Soc. Bengal 60(2): 55. 1891; Schu- mann in Engler & Prantl, Nat. Pfl. Fam. 3(6) : 68. 1895; Koorders & Valeton in Med. 's Lands Plantentuin Buitenzorg 14: 128. 1895; De Dalla Torre & Harms, Gen. Siphon.: 310. 1901; Backer, Schoolfl. J a v a 133. 1911; Ridley, Fl. Mai. Pen. 1: 265. 1922; Bak- huizen v.d. Brink Sr. in Bull. J a r d . bot. Buitenzorg 3, 6: 220 & 245. 1924; Lemee, Diet, descr. Genres 4: 664. 1932; Burkill, Diet. econ. Prod. Mai. Pen. 2: 1537. 1935; Corner, Wayside Trees Mai. 1: 433. 1951. Esenbeckia Blume, Bijdr. Fl. Ned. Ind., 3de Stuk 118. 1825, non Eseubeckia H.B.K. 1825. Blumea Rchb., Consp. 209. 1828, non Nees 1823. Cotylephora Meissner, Gen. Comm. 28. 1837. TYPE SPECIES — N. altissima (Bl.) Bl. Trees, branchlets stout, glabrous, or covered by stellate hairs or by minute fimbriate scales, leaf-scars protruding. Leaves simple, alternate, crowded at the apex of the branchlets, penninerved; nerves patent, parallel, running out arcuately, impressed on the upper surface and prominent on the lower surface. Petioles cylindrical, thickened at apex and base. Stipules extrapetiolar, caducous or long-persistent. Inflorescences generally cymose, axillary, subtended by caducous bracts, covered by stellate hairs and long- fimbriate scales. Buds conical or ovoid-globose. Epicalyx before anthesis completely covering the bud, at anthesis splitting into 2—5 valvate lobes, after anthesis bell-shaped or campanulate, outside lepidote, inside glabrous or in some species pubescent, soon caducous. Calyx monophyllous, coria- ceous, before anthesis cone-shaped or ovoid-globose, in some species um- bonate; after anthesis with a circular, irregularly crenulate apical orifice, sometimes becoming disk-like, the disk with or without incurved margin, outside lepidote (the scales are larger than those on the epicalyx), inside glabrous or partly pubescent. Corolla hypogynous, calyptriform, consisting 4 8 4 - R E I N W A R D T I A [VOL. 5 of 5 free or rarely agglutinate, obtuse petals; aestivation imbricate, slightly contort; the entire corolla soon caducous; petals alternating with the epi- calyx lobes. Stamens numerous, filaments glabrous, filiform, unequal in length, connate at the base only or forming a more or less well-developed staminal tube for half of their length, the upper free part in 5 bundles, alternate with the petals. Each filament with one two-celled anther. Anthers reniform or subglobular, versatile, cells parallel, separate, longitudinally dehiscent. Ovary ovoid, pentangular, 5-celled, sessile, covered by hirsute, stellate hairs, septs alternating with the petals. Ovules anatropous, 5—12 in each cell, biseriate, placentation axillary, ascendant; style one, short, conical or filiform; stigma capitellate, small, rounded or subpentagonous. Fruit cap- sular, ovoid or ellipsoid, woody, 5-angular, 5-celled, almost completely locn- licidally dehiscent into five valves; valves concave, outside muricate-tessel- late, inside densely covered by light-brown, hirsute, prurient hairs. Seeds subhorizontal, alternate on both side of the septae, ellipsoid, smooth, su in- tended by a reddish-brown, thick caruncle. Cotyledons foliaceous, ovate, base emarginate, penninerved, enveloped by the two flat-convex parts of the endosperm. Radicle short, conical, fleshy, basal. DISTRIBUTION — Lower Siam, the Malay Peninsula, Sumatra, Java and Borneo; the centre is found in Borneo. HABITAT — Confined to humid primary forest, from sea level up to 1800 m altitude; on all kinds of soils. K E Y T O THE S P E C I E S 1. a. Calyx after acthesis disk-like 2 b. Calyx after anthesis not disk-like 7 2. a. Lower surface of adult leaf glabrous 3 b. Lower surface of adult leaf pubescent 4 3. a. Buds conical. Style filiform 5 b. Buds ovoid-globose or obovoid. Style conical 6 4. a. Leaves 6—12 X 3—5 cm. Calyx concave 6. N. purpurascens b. Leaves 35—60 X 16—24 cm. Calyx flat 2. N. synandra 5. a. Stipules 1-—1.5 X 0.5—0.7 cm. Fruit-valves areolate, glaucous. 5. N. malayana b. Stipules 2—6 X 0*5—3 cm. Fruit-valves tuberculate, not glaucous 4. N. kostermansiana 6. a. Epicalyx campanulate, buds ovoid-globose. Stipules 2—4 x 0.2—1 cm . . . 1. N. altissima b. Epicalyx bell-shaped, buds obovoid. Stipules 3—9 X 1.5—3 cm . . 3. N. glabra 7. a. Fruit ovoid, rounded S. N. strigosa b. Fruit ellipsoid, acute 7. N. piluliflora 1961] SOEPADMO: A Monograph of the Genus Neesia 483 NEESIA ALTISSIMA (Bl.) Bl. — F i g . 1—2 . Xcesia altissima Blume in Nov. Act. Caes. Leop. Car. 17(la) : 83, t. 6. 1835; Walpers, Rep. 1: 331. 1842; Hasskarl in Tijdschr. Nat. Gesch. en Phys. 12: 125. 1845; Miquel, Fl. Ind. batav. 1(12) 2: 207. 1859; De Sturler, Beschrijv. Houtsoorten Ned. O. Ind. 41. 1866; Masters in J. Linn. Soc. Bot. 14: 504. 1875; Beccari, Malesia 3: 261. 1889; Schumann in Engler & P r a n t l , Nat. Pfl. Fam. 3(6): 68, f. 36 A—C. 1895; Koorders & Valeton in Med. 's Lands Plantentuin Buitenzorg 14: 129. 1895; Jansso- nius, Mikrogr. Holzes 1: 408, f. 52. 1906; Backer, Schoolfl, J a v a 133. 1911; Bakhuizen v.d. Brink Sr. in Bull. Jard. bot. Buitenzorg 3, 6: 221 & 246. 1924, p.p. (excl. N. glabra Becc. and N. synandra Mast.) ; Den Berger in Med. Proefst. Thee, Java 97: 112. 1926; Hej-ne, Nutt. Plant. Ned. Ind., ed. 2, 2: 1058. 1927; ed. 3, 1: 1058. 1950; Furtado in Gard. Bull. S.S. 4: 424. 1929, p.p. (excl. N. glabra Becc. and N. synandra Mast.); Bur- kill, Diet. econ. Prod. Mai. Pen. 2: 1538. 1935. — Esenbeckia altissima Blume, Bijdr. Xed. Ind., 3de Stuk. 119. 1825. — Thespesia altissima (Bl.) Sprengel, Syst. Veg. 4(2) : 257. 1827. — Cotylephora altissima (Bl.) Meissner, Gen. Comm. 28. 1837. — Blume s.n. (L). Payena nigropitnctata Buick in Ann. J a r d . Buitenzorg 5: 53. 1886. — Teijsmann s.n. (BO). Neesia ambigwa Beccari, Malesia 3: 261. 1889;Merrill in J. Str. Br. Roy. As. Soc., Special Number 377. 1921. — Beccari P.B. 3087 (FI). Tree up to 40 m tall, 120 cm in diam.; bark slightly fissured, cracked, greyish-brown; living bark 0.8 cm, brownish-red. Sapwood dirty white, heart- wood brown. Branchlets lenticellate, glabrous or sparsely covered by fim- briate scales, in young plants densely covered by yellowish hirsute, stellate hairs. Leaves coriaceous, oblong or obovate-oblong, (10—) 35—40 (—50) x (6—) 10—15 (—20) cm, emarginate or truncate (in young plants acute), base gradually attenuate, truncate or cordate, margin sometimes slightly undulate and in young plants with stellate hairs or fimbriate scales; nerves 15—20 pairs; upper surface glabrous or sparsely covered by stellate hairs and fimbriate scales, mainly on the nerves; lower surface light green (fresh), glabrous or sparsely covered by fimbriate scales; nerves glabrous or covered by minute fimbriate scales. Leaves of young specimens with stellate or simple hirsute hairs all over. Petioles 4—11 cm, 0.3—0.5 cm in diam., glabrous or sparsely covered by minute fimbriate scales. Stipules caducous, coriaceous, linear-lanceolate, 2—4 x (0.2—) 0.3—0.5 (—1) cm, base trun- cate, apex acute or slightly emarginate, pale green (fresh); young ones with a dense layer of minute fimbriate scales and stellate hairs on both surfaces, glabrescent. Inflorescence cymose, much-branched, multi-flowered, densely scaly, 2—5 cm long, as a rule in the axils of fallen leaves. Peduncle terete or angular, 1—2 cm, 0.5—1 cm thick; pedicels 1—1.5 cm, 0.3—0.5 cm thick. Bracts caducous, ovate, 0.5—0.8 x 0.3—0.4 cm, acute, base truncate, out- side covered by brownish, minute, long-fimbriate scales, inside stellate- 4 8 6 R E I N W A R D T I A [VOL. 5 tomentose. Epicalyx campanulate, with three concave, ovate obtuse lobes, up to 1 cm and 1 cm in diam.; outside covered by minute brownish fimbriate scales, inside stellate-tomentose. Bud ovoid-globose, truncate, at first convex, later concave and umbonate-mammillate, 1—1.5 cm in diam. Calyx disk-like, 2—3 cm in diam., margin circinnately incurved; inside red-brown (fresh, Blume), glabrous or stellate-tomentose, outside brownish, scaly. Petals free, elliptic-oblong, base truncate, 1.5—1.7 x 0.3—0.5 cm; outside densely cover- ed by yellowish-brown stellate hairs, apical part pink, base dirty white (fresh, Blume); inside glabrous. Stamens ca 25; filaments 0.6—0.8cm, connate at the very base, pale straw-coloured (Blume); anthers reniform, yellowish (Blume), ca 0.1 cm in diameter. Ovary ovoid-conical, 0.8—1 cm, apex pink (Blume), gradually merging into the conical, angular, 0.14 cm, glabrous or sparsely stellate-haired style; stigma capitellate, greenish (Blume); ovules ellipsoid, 0.01—0.04 cm, 0.01—0.02 cm in diam., 10—12 in each locule. Fruit ovoid-globose, 15—20 cm, 10—15 cm in diam., acute, base with a short neck; valves outside muricate-tuberculate. Stalk cylindrical, woody, ca 2—3 cm, 1—1.5 cm in diam. Seeds ellipsoid, compressed, 2—3 x 1—1.2 cm, blackish, glossy, smooth. VERNACULAR NAMES — Bungan (Blume, Java), Bengang or Ki Bengang (Java, sometimes also in Sumatra); Punggai, Durian antu (Su- matra) . DISTRIBUTION — Malay Peninsula, Sumatra, W. & C. Java and Borneo. HABITAT — Primary forest, alt. 100—1800 m.; sometimes quite common (Mt. Pajung, Udjungkulon). USE — Timber not durable. A decoction of the fruit's valves mixed with other diuretic substances is used against gonorrhoea (Burkill). Although, this species is closely related to N. synandra Mast., I believe like Furtado, that it is different, as elucidated below: Ar. synandra N. altissima 1. Adult lower leaf-surface densely stel- 1. Adult lower leaf-surface glabrous late-haired. or sparsely covered by long-fimbriate scales and stellate hairs, mainly on the nerves. 2. Bud slender, rounded. 2. Bud broad, concave, umbonate. 3. Calyx flat, margin erect. 3. Calyx concave, with circinnately in- curved margin. 4. Filaments connate into a tube for 4. Filaments only connate at the very half of their length. base. 5. Fruit obtuse. 5. Fruit acute. 1964] SOEPADMO: A Monograph of the Genus Neesia 48f Bakhuizen had reduced N. ambigua and N. glabra to the synonymy of N. altissima. Of N. ambigua I could only examine a sterile fragment of the type specimen; from this material it is not possible to differentiate it from IV. altissima. According to Beccari the fruit of IV. ambigua matches that of N. altissima. N. glabra is a distinct species. It differs from JV. altissima by its very large, up to 9 cm long stipules, obovoid buds and bell-shaped epicalyx. Hasskarl (1855) misidentified the Teijsmann (s.n.) specimen, collected from Banten, with the vernacular name "Karet Kehalan" (a misspelling of "Karet Kihelang", cf. label), as a Sapotoceous species. Burck accepted Hass- karl's contention and described the specimen as Payena nigropunctata. Koorders & Valeton reduced it to N. altissima. MALAY PENINSULA. P e r a k . S. Krian Estate, ster., S.F.N. 36737 (BO, SING); Perak R., Aug., ster., S.F.N. 963 (SING). SINGAPORE. Seletar Road, near 10th mile, May, ster., Corner s.n. (SING). SUMATRA. A t j e h . Gajo, Lueus, Mt. Agosan, alt. 1800 m, Aug., ster., bb. 32402 (A, BO, BZF, L, SING). W. C o a s t . Old Agam, alt. 1300 m, Pebr., ster., 66.2929 (BO, BZF, L) ; Pajakumbuh, Mt. Sago, alt. 1000 m, May, ster., Maradjo 5 (L) ; Padang Upper Lands, Solok, Supajang, ster., s. coll., s.n. (BO) ; ibid., Lubuk Selasih, alt. 1000 m, April, ster., bb. 5501 (BO, BZF, L, W); B e n k u l u . Redjang Lebong, Apr., ster., 66. 2956 (BO, BZF) ; Redjang, N. slope of Mt. Batu Kaba, alt. 1700 m, ster., Endert 1055 (BO, BZF, L); E a s t C o a s t . Bandarbaru, alt. 850m, Jan., fr., Lorzing 7035 (BO); R i a u . Lingga, Febr., ster., bb. 17263 (BO, BZF); P a 1 e m b a n g, Febr., ster., Bum-man v. Vreeden 195 (BO). JAVA. W. J a v a . Banten. Mt. Karang, Pulosari, Tjiudjan, alt. 1050 m, June, ster., Koorders 1,590 (BO, L) ; Tjamara, Tjiringin, Mt. Pangisisan, alt. 10—200 m, July, fl., Koorders 4592 (BO, L) ; Mt. Hondje, alt. 100 m, Sept., ster., Kostermaiis s.n. (BO) ; Tjitoreh, Muntjang. alt. 500 m, fr., Backer 1841 (BO); Bogor. Leuwiliang, Mt. Tjiputih, alt. 500 m, Aug., fr., Bakhuizen v.d. Brink Sr. 6000 (BO) ; near Tjiampea, Tjiteureup R., alt. 500 m, May, fr., Bakhuizen 5503 (BO, K, L, PNH, US) ; near Nan- gela, S.W. Puraseda, Febr., ster., Bakhuizen 7081 (BO, L) ; Nangela, along Tjiteureup R., March, fl., Bakhuizen 7621 (BO, L) ; ibid., alt. 600 m, Apr., fl., Bakhuizen 5884 (B, BO, BRI, CAL, G, K, L, P, PNH, SING, U, UC); Pasir Sireungit, Estate Bolang, alt. 600 m, June, ster., Bakhuizen 6415 (BO) ; Bolang, alt. 700 m, July, ster., Van Steenis 5037 (BO) ; ibid., May, fr., Docters v. Leeuwen 7902 (BO); Pasir Tjihideung, Estate Bolang, alt. 600 m, June, seedling, fr., fl., Bakhuizen 6392 (BO, L, U); Mt. Salak, alt. 1000 m, May, ster., Koorders 33278 (BO, L); ibid., Zippelius 7018 (BO) ; Mt. Pantjar, alt. 500 m, Dec, ster., Dakkus 61 (BO, K, L, PNH, SING, U, UC, WAG) ; Mt. Kembang, July, ster., Ja 5356 (BO, BZF) ; Sukabumi, Mt. Gede, Pasir Tangkil, alt. 800 m, Febr., ster., Ja 3138 (BO, BZF, PNH); Tjiparaj, alt. 1200—1400 m, Febr., ster., Ja 3169 (BO, BZF); Sanggrawa, Djampang- Kulon, alt. 400 m, July, ster., Koor ders 4593 (BO), et 8205 (BO); Tjiandjur, Takoka, Djampang Wetan, alt. 1100m, March, ster., Koorders 12038 (BO, L), et 12039 (BO) ; et 12040 (BO, L), et 32759 (BO), et 39597 (BO); ibid., March, fl., Koorders 15219 (BO, L) ; ibid., Oct., ster., Koorders 25574 (BO, L) ; ibid., Apr., ster., Koorders 4588 (BO), et 4589 (BO) ; Tjiemas, July, 488 ' R E I N W A R D T I A - [VOL. 5 fr., Kartanah s.n. ( B O ) ; T j i r e n g k a s R., a l t . 9 5 0 m , J u l y , s t e r . , Backer 14952 ( B O ) ; T j a d a s M a l a n g , n e a r T j i d a d a p , T j i b e b e r R., a l t . 1000 m , Oct., s t e r . , w i n c k e l 1682 (BO, L) ; C u l t a in H o r t . Bogor., s u b no. X V I . 19—19a, A p r . , fl., Koorders 15045 ( B O ) ; P r i a n g a n . T a s i k m a l a j a , P e n d j a l u , a l t . 720 m , J u l y , fr., Koorders 47801 ( B O ) . C e n t r a l J a v a . B a n d j a r n e g a r a , P r i n g o m b o , a l t . 700—1000 m , Nov., s t e r . , Koorders 451)5 (BO, L) ; ibid., a l t . 800 m, A u g . , s t e r . , Koorders 33786 ( B O ) ; ibid., a l t . 1000 m, S e p t . , fr., Koorders 21917 ( B O , L) ; ibid., a l t . 1000 m, Nov., s t e r . , Koorders 4594 (BO, L ) ; ibid., a l t . 1000 m, Oct., s t e r . , Koorders 21906 (BO, L) ; ibid., a l t . 1000 m, A p r . , s t e r . , Koorders 27183 (BO, L) ; M t . K a p a l , P a n d a n a r u m , a l t . 700—900 m, Nov., s t e r . , Koorders 11139 (BO, L ) . B O R N E O . S a r a w a k . K u e h i n g , J u n e , fr., Beccuri P.B.3087