Revision of the Genus Pandanus Stickman, Part 15 Malayan Species Described by H. N. Ridley

HAROLD ST. JOHN!

As IN MANY OTHER GENERA Henry N . Ridley , but 2-pleated and in sec t ion M-shaped, the director of the Botanic Gard en, Singapore, made blade sword-shaped, from the base gradually noteworthy contributions to the knowledge of tapering to the approximately 20 em long subu­ the Malayan species of Pandanus. His species late apex which about 10 em down is 1.5 mm were validly published in local journals or in wide, at mid section each side with 42 parallel his books. He described them briefly, usually secondary nerves, the base amplexicaul and un­ giving some details of the stem, leaves, appear­ armed but beginning 7-8 em up the margins ance of the syncarp , length of style, and width with serrations 0.6-0.9 mm long, 1- 2.5 mm of the exposed tip of the drupes. He cited apart, pale ; the midrib below from 21 em up specimens but did not adopt the type method. with serrations 0.3-0.5 mm long, 2-10 mm N one of his species were illustrated. The writer apart, subulate-ripped; at midsection the mar ­ has investigated Ridley's species, so ug h t the gins and midrib below with serrae 1-1.5 mm holotypes or has chosen lectotypes in the Singa­ long, 1-3 mm apart ; near the apex the margins pore herbarium. These specimens were kindly and midrib below with rigid serrulations 0.4-0.6 made available by the present director, Dr. H. mm long, 2-4 mm apart; pistillate inflorescence M. Burkitl. 43 em long, 12-13 mm in diameter, trigonous. : Ridley's species of-Pandanus are nearly all bracred, spicate, bearing in the term inal 20 em good, and these are to be maintained . He placed 5 syncarps which are orange and subequal; syn­ most of his species in either the section Rykia carps 6-7 em long, 5-5.5 em in diameter, ovoid or in A crostigma. As will be noted in the treat­ to broadly ellipsoid, bearing about 160 drupes, ment that follows, additional sections are to be these 21-22 mm long, 3.5- 5 mm wide, 3-4 found among his species. The large majority mm thick, lower ~ oblanceoloid, upper Y3 of Ridley's species are treated in this article; subulate, 4-6-angled, the body sides smooth, and the few remaining ones will be illustrated later with exposed fibers; pileus 10-11 mm and described in subsequent parts when -rheir long, of a lower half narrowly conic-pyramidal , type specimens have been located and studied. the surface smooth , sharply angled, the angles continuing along the stout subulate style form­ SECTION A crostigma ing the upper half, arcuate proximally; stigma 3 mm long, linear, in a distal cleft running al­ Pandanus auramiacus Ridl., Roy. Asiat. Soc., most to the apex; endocarp centering in lower Straits Branch, Jour. 41: 49 (1 904) = Y3 , and 7-8 mm long, the walls pale, 0.3 mm 1903; Fl. Malay Penin. 5: 81, 1925 ; thick ; apical mesocarp a broad cavern 2 mm Martelli, Webbia 4 (2 ): r, 30, figs. lO­ long ; basal mesocarp fibrous up the sides, fleshy 13,1914; (sect. A crostigma ) within. Fig. 172 EXPANDED DESCRIPTION: Shrub to 5 m tall; stem with the leaf scars inconspicuous; prop DIAGNOSIS OF LECTOTYPE: "Large branching roots 60 em long; leaves to 2 m long, 4- 5 em shrub with stems 3.6 m. tall, 5-7.5 em. through"; wide, above clear green, below paler and glau ­ leaves "glaucous green"; the lower bracteal leaf cous; foliage leaves little broadened at base, 1.26 m long, 4.2 em wide, coriaceous, l-ribbed the margins beginning at 4 em up with pr ickles

1 B. P. Bishop Museum, Honolulu 17, Hawaii, 1.6-2 mm long, 2- 5 mm apart, stout subulate, U. S. A. Manuscript received July 19, 196 1. ascending; the midrib below beginning at 9

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FIG. 172. Pandanus aurantiacus Rid!., from lectotype. a, Infructescence, X ~; b, drupe, lateral view, X 1; c, drupe, longitud inal median section, X 1; d, drupe, lateral view, X 4; e, drupe, longitudinal median section, X 4; t. drupe, style, and stigma, apical view, X 4; g, leaf base, lower side, X 1; b, leaf middle, lower side, X 1; i, leaf apex, lower side, X 1. Page 274 : Revision of Pandanus, 15. Malaya-ST. JOHN 331 cm with prickles 3-3.5 mm long, 8-20 mm braeted; syncarp 3 cm long, 2.5 cm in diameter, apart, very stout subulate, strongly reflexed; pis­ subglobose, beari ng about 88 drupes, these tillate scape 15-40 cm long, bearing 3- 9 syn­ 14-15 mm long, 4- 5 mm wide, 3-4 mm thick , carps, these 5-7 cm long; drupes very hard until the body ellipsoid to obovoid , 5-6-angled, 10 ripe when they become bright reddish otange mm long; pileus with the base 3.5-4 mm high, and soft. semiorbicular or slightly narrowe r, rather LECTOTYPE : Singapore, Bukit Timah [Bukit smooth, but with longitudinal ridges and lines; Mandai Road}, 1894, H . N. R [idley}, (SING). style 4- 5 mm long, abruptly subulate, angled, Specimen examined and herein designated as sharply proximally curved, bony; stigma 3-4 lectotype. mm long, elliptic-linear, brown, papillose, ex­ SPECIMENS EXAMINED : Singapore, Pasir Pan­ tending to the tip ; endocarp centering in lower jang, 17 Oct. 1912, 1. H. Burkill & M. R. H en­ VI but extending to the base, the walls 0.1 mm derson 6,802 ( BO, SING). Malaya: Johore, Pulau thick, cartilaginous, stramineous; seed 7.5-8 mm Kukub, 1904, H. N. Ridley (pararype); (SING). long, lance-obovoid; upper mesocarp an oblate Ridley, when publishing this species, cited hemisphere, filled with white medullary mem­ three collections and mentioned Sarawak, but branes; basal mesocarp fibrous up the sides, did not designate a holotype. The most com­ fleshy within, sparse. plete of his original specimens is here chosen LECTOTYPE : Malaya, Kedah, Kedah Peak, as the lectotype. woods, 1893, H. N. R[idley} , (SING ) . Specimen examined and here designated as lectotype! Pandanus collinus Ridl., Mater. Fl. Malay Penin. DISCUSSION: P. collinas belongs in the section 2: 228, 1907; Fl. Malay Penin. 5: 79, A crostigma. It was described by Ridley from 1925; Martelli, Webb ia 4 ( 1) : 10, 1913; three collections from Malaya. The best of these (sect. Acr.ostigma) and the one labeled P. collinus! by Ridley is here chosen as the lectotype. Contrasting dif­ Fig. 173 ferences are given for this in our treatment of DIAGNOSIS OF LECTOTYPE : Shrub 1.2-2.1 m P. alticola. tall; stem at summit 6 mm in diameter, brown­ ish, smooth ; leaves 25-45 cm long, 8-11 mm Pandanus glaucophyllus Ridl., Roy. Asiar, Soc., wide, firm chartaceous, apparently pale green Straits Branch, Jour. 41: 50, (1 904) = on both sides and flat except near the base 1903; (sect. A crostigma ) where 2-pleated, l-ribbed, above unarmed, at Fig. 174 the middle with 10-11 secondary parallel veins in each half, below the tertiary cross veins DIAGNOSIS OF HOLOTYPE: "Stems 3-4 m. tall. barely visible, gradually narrow ed in the last 12-25 mm. in diameter," the internodes brown 10 cm to a subulate tip 15-20 mm long and to yellowish,' smooth , mostly higher than the 0.6-0.8 mm wide, the base amplexicaul, un­ nodes; leaves 52-60 "or more" cm long, near ar m ed, blackish purple, beginning at 2 cm the base about 4 em wide, near the apex 5.7-6.5 from the base the margins with prickles 0.7-1 cm wide, appearing narrowly oblong, rather mm long, 3-7 mm apart, subulate, ascending, flaccid and when dry chartaceous, glaucous, 1­ yellowish but red-tipped; the midrib below un­ ribbed but 2-pleated and in section low M­ armed to well beyond the middl e; at mid­ shaped, the secondary para llel veins conspicuous section the margins with subulate tipped ser­ and 34 of them in each half, the apex abruptly rulations 0.2-0.3 mm long, 1.5- 3 mm apart, contracted to a 6-7 em trigonus subulare caudate appressed ascending, red-tipped; near the apex prolongation which near its base is 1 mm wide, the margins with serrulations 0.1-0.2 mm long, the base of the blade amplexicaul and unarmed, 0,3-0.5 mm apart; the midrib below with ser­ but beginn ing at 4 em up the margins with rulations 0.2-0.3 mm long, 0.5-1.5 mm apart; prickl es 0.5-0.8 mm long, 1-3 mm apart, slen­ pistillate inflorescence terminal, erect, bearing der subulate, pale, ascending or diverging and 1 head; peduncle 8 em long, 3 mm in diame ter, some of them doubles or triples; the midrib , ~j o !

FIG. 173. Pandanus coUinus Rid!., from lectotype. a, Habit and syncarp, X 1; b, drupe, lateral view, X 1; c, drupe, longitudinal median section,X 1; d, drupe, lateral view, X 4; e, drupe, longitudinal median section, X 4; I, drupe, style, and stigma, apical view, X 4; g, style and stigma, distal view, X 10; b, leaf base, lower side, X 1; t, leaf middle, lower side, X 1; j, leaf apex, lower side, X 1. oI (~ c ~ u ..o

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FIG: 174. Pandanus glaucophyllus Ridl ., from holot ype. a, Habit andsyncarp, X Y2; b, drupe, lateral view, X 1; c, drupe; longitudinal median section, X 1; d, drupe, lateral view, X 4; e, drupe; longitudinal median section,X 4; t, drupe, style, and stigma, apical view, X 4; g, leaf base, lower side, X 1; b, midrib, lower side, near base, X 4; i, leaf middle, lower side, X 1; j, leaf margin, near middle, X 4; k, leaf apex, lower side, X L 334 PACIFIC SCIENCE, Vol. XVII, July 1%3 below beginning at 4-9 cm with prickles 0.8-1 above," rooting throughout; leaves 27-37 ern mm long, 0:5-6 mm apart, slender subulare, long, near the tip 32-36 mm broad, near the brownish, ascending, oblique, closely appressed; base 16-20 mm broad, appearing ligulare, but at midsection the margins with prickles 0.5-0.7 actually tapering downwards, thin subcoriaceous, mm long, 1-3 mm apart, slender subulate, as­ dark green above, pale and glaucous beneath, cending flat appressed and occupying recesses 2-pleated and in section low M-shaped but only in the margin; the midrib below glabrous or l -ribbed, the midrib below narrow and raised, with a few prickles like those lower down; at the secondary parallel veins prominent through­ the bend of the apical contraction the margins out, and at midsection 18-21 on a side, the terti­ with prickles 0.6-0.8 mm long, 0.6-1.5 mm ary cross veins faintly visible below near the tip, apart, slender subulate, somewhat ascending or making small squarish meshes, near the apex ab­ divergent; the midrib with prickles 0.6-1 mm ruptly rounded contracted to a 3-6 cm caudate long, 1-2 mm apart, arcuate subulate, ascend­ deltoid subulate tip 0.8 mm wide, the base am­ ing; on the caudate tip the margins and mid­ plexicaul, unarmed for 4-5 ern, then the margins rib below with prickles 0.3-0.7 mm long, with prickles 0.7-1.3 mm long, 2-7 mm apart, 0.3-1.5 mm apart, stout arcuate subulate, as­ slender subulate, straight or slightly curved, cending; pistillate peduncle 8 ern long, 7 mm brownish, ascending; the midrib below unarmed in diameter, erect, bearing a single syncarp 5.7 to near the apex; at midsection the margins with cm long, 4.5 ern in diameter, broadly ellipsoid, similar prickles 0.5-1 mm long, 4-12 mm apart; bearing about 320 drupes, these 12-13 mm on the rounded apical contraction the margins long (or 16-18 mm following the curve of the with prickles 0.5-0.8 mm long, 0.5-2 mm apart, stigma), 5-6 mm wide, 3.5-4.5 mm thick, the stouter subulate, slightly ascending; the midrib body narrowly oblong ellipsoid, 5-6-angled, the below with prickles 0.8-1.4 mm long, 1-7 mm sides smooth; pileus smooth, the base 3-3.5 mm apart, arcuatesubulate, slightly ascending; on long, conical or broadly so, bearing the terminal the caudate tip the margins and midrib below style 5-6.5 mm long, stout subulate, strongly with stout prickles 0.3-0.8 mm long, 0.5-1.5 curved proximally, having a distal cleft nearly its mm apart, subulare, arcuate, ascending; in the whole length in which is the stigma 3-4 mm series of bracteal leaves the first several 2-4 long, linear, brown, papillose, running to the tip; cm long, lance-navicular, entire; pistillate in­ endocarp. pale brownish, centering in the lower florescence I-headed; peduncle 5 cm long, 4-5 Y3, its walls 0.1 mm thick; seeds 7-8 mm long, mm in diameter, 3-angled, few braeted; syncarp oblance-oblong; apical mesocarp a cavern 2-2.5 erect, 3.5 cm long, 3.3 cm in diameter, ovoid­ mm long, invaded by trichomes and hairs; basal . globose, bearing about 150 drupes, these 11-12 mesocarp with fibers up the sides and the center mm long or following the curve 14-1.5 mm fleshy. long, 3.5-5.5 mm wide, 2.5-3.5 mm thick, fusi­ HOLOTYPE: Malaya, "Perak, Thaiping Hills, form, the body 7-8 mm long, oblanceoloid, in dense forest about 2500 feet altitude," above the surface smooth, later fibrous; pileus smooth, tea border, Dec. 1898, H. N. Ridley (SING) . apparently glaucous, 9~10 mm long, the base Holorype examined! oblong-pyramidal and with the apex oblique Pandanus parvus Ridl., Roy. Asiar, Soc., Straits and proximally curved, tapering into a hornlike Branch, Jour. 33: 71, Jan. 1900; Fl. style which is 5-6 mm long, all (except the Malay Penin. 5: 78, 1925; P. flagellifer apical ones) sharply proximally curved, stigma Warb., in Engler's Pflanzenreich IV, 9: 4-5 mm long, lance-linear, dark brown, papil­ 80, Dec. 21, 1900; Martelli, Webbia lose, extending tocJl.e tip; endocarp in lower 4(1): 14,1913; 4(2) : r. 33, f. 7,1914; ~, stramineous, bony, the walls 0.3 mm thick, (sect. Acrostigma) and the lateral walls extending upwards to bound the apical cavity; seed 5 mm long, ob­ Figs. 175, 176 lanceoloid, truncate at tip; apical mesocarp .a . DIAGNOSIS OF HOLOTYPE : "Stem 2.92 ern. cavern filled with white medullary-ascending tall, 12 mm. in diameter, slender, branched hairs; basal mesocarp sparse, fibrous and fleshy. 1cm' 1 b 1c

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FIG. 175. Pandanus parvus Ridl., from holotype. a, Habit and syncarp , X 1; b, drupe, lateral view,X 1; c, drupe, longitudinal median section, X 1; d, drupe, lateral view, X 4; e, drupe, longitud inal median sec­ tion, X 4; t. dru pe and stigma, apical view, X 4; g, leaf base, lower side, X I; b, leaf middle, lower side. X I; i, leaf apex, lower side, X 1. 336 PACIFIC SCIENCE, Vol. XVII, July 1963

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FIG. 176. Pandanus parvus Ridl., from Singapore, cult., 1895, H . N . Ridley . a, Staminate inflorescence, X 1; b, stamens, lateral view, X 10; c, /loral bract, lateral view, X 1.

EXPANDED DESCRIPTION: Pistillate stems DESCRIPTION OF STAMINATE PLANT: Stami­ 0.8-2.92 m tall, 6-12 mm in diameter; prop nate inflorescence 15-18 cm long, bracts white roots up to 50 cm long, 4-5 mm in diameter, and somewhat fleshy; peduncle 3-4 mm in di­ smooth, green to brown, nearly vertical ; leaves ameter, fleshy, soon withering and caducous; 27-50 ern long, 21-48 mm wide, the caudate lowest bracts 8-13 cm long, 26-30 mm wide, apex 3- 9 cm long; peduncle 5-12 cm long; sterile, the body 6-9 cm long, navicular, with syncarp 2.5-4 cm long, 2.5-3.3 cm in diameter, prominent parallel veins, the lower half with globose or ovoid-globose. margins unarmed; near the apex the margins I I Page 280 : Revision of Pandanus, 15. Malaya-8T. JOHN 337 with prickles 0.5-1 mm long, 0.3-1 mm apart, serrulations 0.3-0.5 mm long, 2-3 mm apart; subulate, diverging or subascending; the mid­ the midrib below with similar ones but 2-4 rib with stouter prickles 0.7-1 mm long, 0.5-1 mm apart; peduncle 15 em long, 8-9 mm in mm apart, slightly ascending; · on the caudate diameter, straight, bracted; syncarps 3-4, spicate, tip the margins and midrib below with prickles crowded, 4.5-6.5 em long, 3.5-4.5 em in di­ 0.5-0.9 mm long, 0.5-1 mm apart, arcuate ameter, elliptic-cylindric (rarely subglobose), subulate, ascending; median bracts 3-4.5 em appearing spiny from the numerous curved as­ long, 1.6-2.4 em wide, Ianceolate, the margins cending styles (but a few at the very base and the midrib below towards apex minutely reflexed ), bearing about 430 drupes; drupes prickly; spikes 5-8 and 1.5-2.5 em long, 0.8-1.3 20-22 mm long, 5-6 mm wide, 4-5 mm thick, em in diameter, ellipsoid, dense; stamens very oblance-fusiforrn, compressed, apparently green, numerous, attached singly and directly to the upper Y2 free, the body 13-14 mm long, ob­ axis; filament 0.2 mm long; anther 4.5-5 mm lanceoloid, the apex truncate, pileus at last long, 0.2 mm wide, bearing an apical subulate shedding, the baseserniorbicular, bearing a style prolongation of the connective 0.4-0.7 mm 6-7 mm long, stout subulate, ridge angled, long. gently arcuate towards apex of syncarp; stigma HOLOTYPE : Singapore, Kranji, 1892, H. N. 4-5 mm long, linear, distal, running to the very R [idley], (SING). Holotype examined! apex; endocarp 6-7 mm long, centering in SPECIMENS EXAMINED : Singapore, .Hinterin­ lower Y;, ellipsoid, truncate, stramineous, car­ dien, Dec. 16, 1905, A. Engler 3,862 (B); Singa­ tilaginous, the sides 0.15 mm thick; apical meso­ pore, Bidadari, 1898, H. N. R[idley] 9, 171 carp 3-3.5 mm long, a fat discoid cavern with (FI ); Singapore, cult., first male flowers seen, a few white membranes; basal mesocarp fibrous 1895, H. N. R[idley], (FI, SING); without col­ along the margins, fleshy within. lector, ex Hb. (SING), Forest Iurong, low grow­ EXPANDED DESCRIPTION: Pistillate plants ing plant, Jan. 10, 1889, Flora of Singapore 92 with peduncle 13-15 cm long; syncarps 2-3 (B), the holotype ofp, flagellifer. and 4.5-7 cm long, 3.5-5.5 cm in diameter. Malaya: Johore, Sungei Pelepah Kiri, 26 June LECTOTYPE: Malaya, "Perak, Maxwell's Hill"; 1928, E.]. H. Corner 35,556 (SING ). June 1893, H. N. R[idley], (SING). Lectotype examined! Pandanus perakensis Rid!., Mat. F!. Malay Penin. SPECIMENS EXAMINED : Malaya, Pahang, Ku­ 2: 231, 1907; ri Malay Penin. 5: 81, ala Lipis, 1892, A. S. Machado 11,633 (SING); 1925; (sect . Acrostigma) Kedah, Baling Hill, 21 Nov. 1941, E. ]. H. Corner & ]. C. Nauen (SING). Fig. 177

DIAGNOSIS OF LECTOTYPE : (Probably acau­ Pandanus spinulosus (Rid!.) comb. nov. (sect. lescent}; leaves 1.5-1.8 m long, 2-3.6 em wide, Acrostigma ); P. collinus Ridl, var. spinu­ subcoriaceous, light green, linear, rather ab­ loses Rid!., Fl. Malay Penin. 5: 79, 1925. ruptly narrowed to the 9 cm subulate, trigonous Fig. 178 apex, this 10 em down 6 mm wide, in section the blade M-shaped, at midsection with 39-41 DIAGNOSIS OF HOLOTYPE : Stem slender, just secondary veins in each half; near the base the below the leaves 4 mm in diameter; leaves margins with spines 2-3 mm long, 5-13 mm 67-68 em long, 16 mm wide, subcoriaceous, apart, stout subulate, arcuate, pale, broad based, green above, below paler and apparently some­ ascending; the nearby midrib below with spines what glaucous, with a central furrow over the 2.5-3 mm long, 6-13 mm apart, similar but midrib, 2-pleated, and towards the tip the 2 even thicker subulate and reflexed; at mid­ pleats spinulose above, at midsection with 16-18 section the margins with subulare-ripped serrae parallel secondary veins in each half, these 0.5-1.5 mm long, 4-8 mm apart, the midrib prominent throughout, below the tertiary cross below with prickles 0.5-1 mm long, 8-15 mm veins visible, remote and forming elongate apart, subulate; near the apex the margins with meshes, the blade Iigulate, in outer Y; tapering 1 I 0 10 em,

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FIG. 177. Pandanus peraeensis RidI., from lectotype. a, Infrucrescence, X 1; b, c, drupe, lateral view, X 1; d, drupe, longitudinal median section, X 1; e, drupe, lateral view, X 4; I, drup e, longitudinal median sec­ tion, X 4; g, drupe, style, and stigma, apical view, X 4; b, leaf base, lower side, X 1; i, leaf middle, lower side, X l ; ;, leaf apex, lower side, X 1. ~ o..

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FIG. 178. Pandanus spinulosus (Rid!.) St. John, fr om holotype. a, Infructescence, X 1; b, drupe, lateral view, X 1; c, drupe, longitudinal median section, X 1; d, drupe, lateral view, X 4; e, drupe, longitudinal median section , X 4; f. drupe, style, and stigma, apical view, X 4; g, style and stigma, lateral view, X 10; b, leaf base, lower side, X 1; i, leaf middle, lower side , X 1; i, leaf apex, upp er side. X 1; k, leaf apex, lower side, X I. i 'SCM.

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FIG. 179; Pandanus 'unguiculatus Rid!., from holotype. a. Syncarp, X ' I; b, drupe, lateral 'view, X I;c, drupe, longitu din al median section, X 1; d, drupe, lateral view, X 4; e, dru pe, longitudinal median section, X 4; t. drupe, style, and stigm a, apical view, X 4; g, leaf base, lower side, X 1; b, leaf mid dle, lower side; X I ; i, leaf apex, lower side, X 1. Page 284: Revision of Pandanus; 15. Malaya-ST. JOHN 341 gradually to the subulate, deltoid apex about 15-17 mm long, the pileus base narrowly hemi ­ 3 em long and 1.5 mm wide, the base amplexi­ spheric, gradually tapering into the thick subu­ caul, unarmed, partly dark purple, beginning at late style which is 5-5.5 mm long. 4 em up the margins with prickles 0.7-1.5 mm long, 3-5 mm apart, subulate, ascending at 45 °, Pandanus unguiculatus Ridl., Mat. Fl. Malay the base thickened, pale; the midrib below un­ Penin. 2: 229, 1907; Fl. Malay Penin. armed to beyond the middle; at midsection the 5: 78, 1925; (sect. Acrostigma) margins with prickles 0.7-0.9 mm long, 2-4 Fig. 179 mm apart, subulate, ascending; near the tip the DIAGNOSIS OF HOLOlYPE: "Plant small"; margins and midrib below with subulate ser­ leaves 26-48.5 em long, near the base 2-2.3 . rations 0.4-0.6 mm long, 0.7-1 mm apart; pis­ em wide, near the middle 2.4-2.7 em wide, tillate inflorescence erect, bearing a single syn­ ligulate, but perceptibly tapering both ways carp; peduncle 10 em long, 4-5 mm in di­ from the middle, dark green above, pale green ameter, 3-sided, bracteate; syncarp 4.5 em long, below, thick charraceous, only the midrib thick­ 3.1 em in diameter, oblong-ellipsoid, bearing ened, but the blade pleated and M-shaped in about 120 drupes, these 15-17 mm long, 3-4.5 section, the secondary parallel veins conspicu­ mm ~ide and thick, pale green , fusiform but ous, at midsection 17-19 in each side, 'the ter­ the upper end contracted and more slender, 5-6­ tiary cross veins visible in upper third below, angled, the body 11-12 mm long; pileus 8-9 oblique, forming short, rather oblong meshes; mm long, the base 2.5-3 mm high, narrowly near the apex abruptly contracted to a subulate semiorbicular, with 5-6 sharp angles, these tip more than 20-25 mm long, 0.9 mm wide, mostly single ; style 5~5.0

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FIG. 180. Pandanus stelliger Rid!., from lectotype. H, Drupe, lateral view, X 1; e, drupe, longitudi­ nal median section, X 1; I, g, drupe and stigma, apical view, X 1; b, drupe, lateral view, X 4; i, drupe, longitudinal median section, X 4; j, drupe and stigma, apical view,X 4; I, leaf base, lower side, X 1; m, leaf middle, lower side, X 1; n, leaf apex, lower side,X 1. Page 286 : Revision of Pandanus, 15. Malaya-ST. JOHN 343

EXPANDED DESCRIPTION FROM ALL SPECI­ SECTION Ast erostigma MENS EXAMINED: Stems up to 1.8 m tall, 9-13 mm in diameter, smooth, yellowish, erect or Pandanus stelliger Ridl ., Roy. Asiar, Soc., Straits ascending, with aerial roots; leaves arching out­ Branch, Jour. 41: 49"':50, (1 904) = ward and the tips pendent, glaucous below, the 1903; Fl. Malay Penin. 5: 76,1925; Mar ­ subulate tip 20-46 mm long ; inflorescence erect ; telli, Webbia 4 (2 ) : t.26,f.1-2il, 1914; peduncle 5-6 ern long. (sect. Asterostigma ) HOLOlYPE:Malaya, Selangor, Bukit Kuru, Figs. 180, 181 May 1896, H . N. R[idley] 7,659 (SING) . Holo­ type examined! DESCRIPTION OF LECTOlYPE: Shrub 1.5-3.3 SPECIMENS EXAMINED: Malaya, Pahang, Fra­ m tall, the slender stems pale brown, 2.5 cm in ser's Hill, 19 Aug. 1937, E;>?-J; H . Corner 33,233 diameter, branched, bearing a few mucros; (SING). leaves dark green, 55-60 cm long, near the base

FIG. 181. Pandanus stelliger Rid!., from lectotype, habit and infructescence,X Y2 . 344 PACIFIC SCIENCE, Vol. XVII, July 1963

1.8 cm wide, near the middle 2.8-3.8 cm wide, mm apart ; staminate inflorescence 47 ern long, in section with a broad, central channel, and leafy bracted; lowest bract 47 ern long, 2.2 ern hence M-shaped, the secondary parallel veins at wide, for the lower 10 cm slightly distended, midsection 19-20 in each half, prominent, the sheathing and paler, the rest foliaceous; floral tertiary cross veins visible below, transverse or bracts thin , apparently white, the median one oblique, forming short, somewhat oblong 15 ern long, 4.6 cm wide, lanceolate, on the outer meshes, the actual base amplexicaul and un­ 1;3 the margins and midrib below with serrula­ armed, but beginning at 2-2.5 cm the margins tions 0.1 mm long, 0.2-1 mm apart ; the 7 with prickles 1.5-2.5 mm long, 4-9 mm apart, spikes 3-6 cm long, 12-15 mm in diameter, broad subulate, ascending, pale; the midrib cylindric, dense; fascicle 5-8 mm long, with unarmed except near the tip ; at midsection the 19-22 stamens, the naked common column 3-4 margins with prickles 2.1-2.5 mm long, 7-23 mm long, rhachis 2-2.5 mm long, clavate; free mm apart, similar; near the 3-8 ern apex the filament tips 1-2 mm long, divergent; anthers margins and midrib below with serrulations 1.5-2 mm long, linear-oblong, bearing a pale 0.2-0.3 mm long, 2-6 mm apart, brown; pe­ subulate projection of the connective 0.2-0.3 duncle terminal, 5 cm long, leafy bracred, bear­ mm long. ing 4 syncarps in a dense spike about 10 cm LECTOTYPE: Malaya, "Selangor, on the Tras long; syncarps subequal, 5-6 ern long, 3.5-4.5 Route at the 20th mile (8775) ," 2,000 ft. elev., cm in diameter, ovoid to broadly ellipsoid, bear­ July 1897, H. N. Ridley (SING). Lectotype here ing 64-88 drupes, these 19-21 mm long (in­ designated ; specimen examined! Isotype (FI)! cluding the stigma), 7-10.5 mm wide, 6-8 mm SPECIMENS EXAMIN ED: Malaya: Perak, Larut, thick, oblanceoloid with :an obtusely convex top of mountains, open jun[gle], 2,000 to 3,000 apex, the sides 5-6-angled, smooth, when dried ft. alr., April 1883, H . Kunstle« 4,166 (CAL); brown, the body 15-17 mm long, cuneiform; Haram Parak, June 1884, Scortechini (CAL) . pileus 6-7 mm long, oblate semiorbicular; style Pahang : Gunong Tahan, 7 Sept. 1937, E. J. subterminal, 0.5-1 mm long, vertical except on H. Corner (SING); Sungei Chelia, Chegar Perah, the basal drupes where sharply reflexed distally; 14 Oct. 1927, M. R. Henderson 19,370 (SING). stigmas on the expanded apical surface, 4-7 Penang: Government Hill , Dec. 1895, C. mm in diameter, 4- 7-pointed, horizontal or Curtis (SING) ; Waterfall Hill, 500 ft. alt., June mushroom-shaped, papillose, brown; drupe body '88, L. W ray Jr. 2,22<7 {CAL ) . 14-17 mm long; endocarp centering in lower 1;3, bony, light brown, the inner surface shining, the lateral walls 0.5 mm thick, the apical plug SECTION H ombronia 1-1.3 mm thick; seeds 6-7 mm long, obdeltoid; apical mesocarp with a cavity 6-7 mm long, Pandanus irregularis Ridl., Fl. Malay Penin. 5: with a few white membranes near the margins ; 76-77, 1925; Henderson, M. R., Roy. basal mesocarp sparse, fibrous and fleshy. Asiat. Soc., Malay Branch, Jour . 17: 82, EXPANDED DESCRIPTION OF ALL PISTILLATE 1939; (sect. H ombronia) PLANTS EXAMINED: Plants 1.5-4 m tall; leaves Figs. 182, 183 55-113 ern long, at midsection with 19'-29 sec­ ondary parallel veins in each half; peduncle 5-7 DIAGNOSIS OF HOLOTYPE: "Leaves long, very ern long, bearing 2-5 syncarps, these 4-6 ern broad," 9.3 em wide, thick and leathery, 1­ long; drupes 7-13 mm wide, 6-10 mm thick, ribbed, 2-pleated, in section depressed, M­ bright yellow. shaped, the secondary parallel veins visible be­ DESCRIPTION OF STAMINATE PLANTS: Leaves low, scarcely so above and at midsection 83-92 63-70 cm long, near the base 20-21 mm wide, in each half, the base amplexicaul and unarmed, near the middle 26-29 mm wide, at midsection but beginning at 12 cm up the margins with the parallel secondary veins 22-24 in each half, thorns 4.5 mm long, 1-2 cm apart, heavy deltoid and at midsection the margins with appressed subulate, black-tipped, ascending; the midrib ascending serrulations 0.3-0.4 mm long, 7-12 below with similar reflexed thorns 2-3 ern apart; o 2 em, I I

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FIG. 182 . Pandanus irregularis Ridl ., from holotyp e. a, Infrucrescence, X 113 ; b, drupe, lateral view, X 1; c, drupe, longitudinal median section, X 1; d, drupe and stigmas , apical view, X 1; e, drupe, transverse median section, X 1; t, leaf base, lower side, X 1; g, leaf middle, lower side, X 1. 346 PACIFIC SCIENCE, Vol. XVII, July 1963

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FIG. 183. Pandanus irregulsris Ridl., from holorype, leaf apex, lower side, X 1. on a leaf section cut higher up (exact place 8-9 mm long, 3 mm in diameter, ellipsoid ; api­ unknown) the margins with spines 3-3 .5 mm cal mesocarp cavernous, with strong, longitudi­ long, 25-36 mm apart, heavy subulate, ascend­ nal fibers and soft pith; bssal-mesocarp fibrous ing; the midrib below with scars of fallen spines and fleshy. 3-6 em apart; pistillate inflorescence 30 em EXPANDED DESCRIPTION OF ALL SPECIMENS long, with about 7 syncarps; peduncle 15 mm EX AMINED: Leaf more than 1.4 m long, near in diameter, obtusely trigonous, bracted; syn­ the base 9 em wide, near the middle 12 em carps 6.5-7.5 em long, subglobose, bearing wide, near the apex 10.5 em wide, broad ligu­ about 36-44 drupes , these 1-3- but mostly 2­ late; margins near the base"with thorns 3.5-4.5 celled, 3-3.5 em long, 1.8-2.4 em wide, 1.2-1.5 mm long, 1-2.5 em apart ; midrib below with em thick, "dark green," narrowly obovoid to thorns 3-3.5 mm long, 2-5 em apart, subulate, oblanceoloid, compressed, the apex entire, ob­ reflexed; at midsection the margins with spines tuse, the sides smooth, convex, lateral sutures 4-4.5 mm long, 20-25 mm apart, sharply as­ none; stigmas almost all eroded but 1 basal cending, subulate but the base much thickened; drupe bears a single stigma 5 mm long, lanceo­ the midrib below concealed; the apex apparently late, sharply bent to the horizontal,-the recep­ damaged in growth and without a caudate apex, tive surface distal; endocarp 22 mm long, cen­ rather contracted to 2 broadly obtuse lobes and tering in upper ~ , at midsection filling the deeply emarginate, and even Prickly on the drupe, bony, massive, dark brown but including margins of the cleft, all this, however, appar­ pale fibers, the lateral walls 4- 5 mm thick ; seeds en tly an abnormality; peduncle 30 em long; Page 290: Revision of Pandanus, 15. Malaya-5T. JOHN 347 spike 14 cm long, bearing 4 syncarps; stigmas a subulate apex or compressed and more or less 4 mm long, sharply proximally bent, the recep­ oblong, with the apex sharply bifurcate; stig ­ tive surface usually proximal. mas centripetal, ellipsoid or lanceolare; stami­ HOLOlYPE : Malaya, Kelantan, Sungei Ketah, nate plants unknown. Bam Bau (Bow), 15 Feb. 1924, Mohamed Nur HOLOlYPUS : Palldallus Klossii Rid!., Fed. & Foxworthy 12,104 (SING). Holotype exam­ Malay States Mus., Jour. 6: 190-191, 1915. ined! DISCUSSION : The single known species occurs SPECIMENS EXAMINED: Malaya, Pahang, Gua on the mountains of Pahang in Malaya. It com­ Tipus, the common pandan on the limestone, 17 bines the characteristics of the section Rykia Oct. 1927, M. R. Henderson 19,468 (SING). in its hornlike, simple or bifurcate styles, and DISCUSSION : P. irregularis was placed by Dr. of the section Pandanus, in its many-carpellate, Ridley in the section Rykia. It has, however, the large phalanges. Since these two sections are drupes 1-3-celled, and usually 2-celled, with very distinct and are only remotely related, it the cells in linear arrangement, and the large is best to make the very bizarre P. Klossii the stigmas terminal, but distal and laterally di­ basis of a new section." rected. There is no question but that it must now be placed in the section H ombronia. Pandanas Klossii Ridl. , Fed. Malay States Mus., It is one of the numerous plant species known Jour. 6: 190-191, 1915; Fl. Malay Penin. to occur only on limestone knobs. These pre­ 5: 74, 1925; (sect. Malaya) cipitous pinnacles or narrow ridges rise spec­ tacularly from the rolling uplands of Thailand Figs. 184, 185 and Malaya. Their sides are often precipices of shining, white limestone, sparsely vegetated, but DIAGNOSIS OF HOLOlYPE : Shrub 2-7 m tall; their crests have-a .dark .evergreen, virgin forest stems usually solitary, 7.5 cm in diameter, gray, cover. To a botanist they are very attractive. erect or arcuate; leaves 1.3-1.8 m long, 6.8-7.2 Four species of Pandallus in Malaya are re­ cm wide, thick coriaceous, above dark green, stricted to these calcareous habitats. Besides this below pale green, I-ribbed, but 2-pleated and species published by Ridley, two more are now in section low M-shaped, the longitudinal paral­ being described, and one more has yet to be lel secondary veins 0.5-0.8 mm apart, 49-52 found in fruit. A good account of this localized in each half, evident on both sides as are the flora is given by Dr. M. R. Henderson (Roy. tertiary cross veins that form a network of Asiar, Soc., Malay Branch, Jour. 17: 13-87, pI. meshes, oblong or rhombic, longer than broad, III-XI, 1939). the blade ligulate, and only near the tip rather abruptly narrowing to a 5 cm stout subulate tip, Malaya, SECT. NOV. trigonous and 2-3 mm wide, the very base not collected, near the base the margins with stout Phalangibus pluricarpellatis, stylis rerminali­ thorns' 5-8 mm long, 8-30 mm apart, arcuate bus adscendentibus vel arcuatis corniformatis in­ ascending, black; the midrib below with similar regris vel bifidis, stigmaribus cenrripetalibus el­ but reflexed thorns; at midsection the margins lipsoideis vel lanceolaris; plantis masculis in­ with prickles 3 mm long, 15-50 mm apart, cognitis. subulare, ascending, flat appressed; the midrib Phalanges of several united carpels; styles ter­ below with prickles 2 mm long, 35-50 mm minal, ascending or arching, stout, hornlike with apart, heavy, arcuate, ascending; near the tip

2 The new section can be inserted in the author's key in this Revision of the Genus Pandanus (p. 2) by replacing the 11th line from the bottom by :

L. Stigmas elongate, acerose, or stout hornlike, simple or bifurcate, a. Stigmas stout hornlike, simple or bifurcate __ Malaya a. Stigmas elongate, acerose, I o

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FIG. 184. Pandanus Klossii Rid!., from holotype. a, Phalange, lateral view, X 1; b, phalange, longitudi­ nal median section, X 1; c, phalange, and stigmas, apical view, X 1; 'd, 'carpel apices and stigmas, apical view, X 4; e, style and stigma, oblique view, X 4; t , leaf base, lower side, X 1; g, leaf middle, lower side, X I; b, venation in outer third, lower side, X 4; i, leaf apex, lower side, X 1. Page 292: Revision of Pandanus, 15. Malaya-ST. JOHN 349 the margins and midrib below with serrations 0.3-0.5 mm long, those of the margin 1-1.5 mm apart, those of the midrib below 2-5' mm apart; pistilla te peduncle 15-30 ern long, 12-18 mm in diameter, 3-sided, leafy bracted; syncarp sin­ gle, globose or ellipsoid, about 15 cm in di­ ameter; phalanges numerous and 6-6.5 ern long, 3.6-3.9 cm wide, 2-2.7 ern thick, cuneiform,' 4-6-angled, "showing signs of turning red or orange," the sides smooth , gently convex, free in upper Y:l ,the apex low convex, lateral sutures visible only in upper Y:l; central apical sinuses 1-4 mm deep, straight or arcuate, narrow; car­ pels 9-10 (in a double 16-17 and the phalanges 5.7-6 cm wide), the apices semiorbicular, faintly angled; styles 4-6 mm long, dark, bony, either simp1e and hornlike or flattened and sharply bifid, mostly centripetal; stigmas as long o as the styles, ellipsoid or lanceolate, brown, pap­ .illose, covering the proximal face; proximal sinus none; endocarp in lower Y:l and 2.7 cm long, bony, massive, dark, brown, the lateral walls 2 mm thick, the inner surfaces shining and rugulose; seeds 14-16 mm long, 2-3 mm in diameter, oblong-ellipsoid; upper mesocarp 2-2.5 cm long, of dense pith and a few fibers; basal mesocarp fibrous and fleshy. ADDED DESCRIPTION FROM ALL SPECIMENS EXAMINED: Phalanges about 96; carpels 5-10; syncarp 9-15 cm in diameter. HOLOTYPE: Malaya, Pahang, Gunong Tahan, July 1911, H. N. Ridley 16,037 (SING). Iso­ type (BO). Ridley stated,"Common all over FIG. 185 . Pandanus Klossii Ridl., from hol otype , the Padang. In the more open exposed spots syncarp, X Y:l. the stem is short and erect, about 6 to 8 feet tall; in the woods the stems are long and weaker, Gunong Tahan, 14 Sept. 1937, E. ] . H . Corne-r often falling about at all angles, 20 feet or (SING); Pahang, Gunong Tapis , Kuantan, 4,600 more long. No trace of male flowers could be ft. alt., 14 June 1934, C. F. Symington & Kiah seen anywhere. The plant is very abundant, al­ 28,843 (SING ). . most filling up the woods in some places:' The . DISCUSSION : P. Klossii Ridl., a very unique collection, Ridley 16,037, was made by Ridley species, is here made the holotype for the new previous to his publication of the species in section Malaya. This combines the characters of 1915, was labeled P. Klossii by him, and seems the two sections Pandanus and R ykia. When to have been the only collection in the Singa­ Ridley described the species in 1915 he did not pore herbarium up to that time . In his Flora place it in a section, but later in 1925 in his of the Malay Peninsula (1925) he accepted the Flora of the Malay Peninsula he put it in the species and cited only his collection from Gu­ section Keura, which 'is now called Pandanus. nang Tahan, 'so this one seems acceptable as .As usual he had a good eye for species, but his his holotype. description seems to have been hastily drawn. SPECIMENS EXAMINED : Malaya, Pahang, He stated (1915: 191) "the drupes are separate 350 PACIFIC SCIENCE, Vol. XVII , July 1963 till the fruit is nearly ripe, when from 6 to 7 Pandanus pentodon Ridl., Fl. Malay Penin. 5: become adnate and remain so as the whole fruit 75, 1925; (sect. Multidens) breaks up." The holotypic collection consists of Fig. 186 two sheets, one with overripe fruit, the other with a small, immature syncarp. In both the DIAGNOSIS OF L ECTOTYP E: Stem short, phalanges are of several fused carpels. The car­ branched, up to 2 em in diameter ; leaves said pels are fused from the base to with in 4 mm to attain more than 100 em in length , 11-12 of the apex, and the sides lack sutures.The writer mm wide, coriaceous, dark glossy green, pale sees no evidence on these specimens to confirm green beneath, channeled above the midrib and Ridley's statement that the drupes are separate with 2 lateral pleats, in section low M-shaped, and only tardily connate; on the contrary, all narrowly ligulare, outer halves of leaves missing, the structural evidence indicates that the pha­ at midsection with 17-22 secondary parallel langes are of several carpels united from the veins in each half, the tertiary cross veins barely beginning. visible below near the base, the base amplexi­ Though Ridley did not explain the source caul and unarmed, apparently copper-colored, of his specific epithet, it was obviously given in beginning at 3-5 em up the margins with pale honor of Cecil Boden Kloss, an English zoolo­ serrulations 0.5-0.8 mm long, 1-6 mm apart; gist and to a lesser degree a botanist. In 1903 the midrib unarmed to at least beyond the mid­ and 1907 he was employed by the Singapore dle; at midsection the marg ins with slender ser­ Botanic Garden. He was later director of the rulations 0.3-0.6 mm long, 4- 8 mm apart, the Raffles Museum. upper surface unarmed; the apex tapering to a 10 em trigonous subulate tip which 10,'em Multidens, SECT. NOV. down is 4 mm wide, the margins and midrib below with serrations 0.2-0.3 mm long, 0.7-3 Frutices, foliis angustis, syncarpio solitario el­ mm apart; pistillate peduncle 13 em long, 8 mm lipsoideo, srylo singulo terminali osseoso prox­ in diameter, bracted, bearing a single, erect ime curvato basi cylindrico margine (2-) 3-6c syncarp; the highest peduncular bract 30 or lobaro lobis deltoideis, drupis 1-loculatis. Plantis more em long, at base 2.2 em wide, narrowly masculis cum inflorescentia bracteata l-spicata, lanceolate and tapering noticeably right from staminibus distinctis in axili affixis, filamentis the base, the margins. with serrulations similar 4-plo tam longis quam antheris. to those of the leaves, but on the upper surface Bushes; leaves narrow; syncarp solitary, el­ beginning 10 em up the 2 lateral pleats with lipsoid; style single, terminal, proximally curved, similar serrulations, the lower ones 5-1 3 mm bony, shining, the base cylindric, the margin apart, the upper ones 2- 5 mm apart; partly (2-) 3-6-lobed, the lobes deltoid ; drupes 1­ hiding the fruit are 3 series of floral bracts, 3 celled. Staminate plants with inflorescence brae­ deep, and subequal , 5 em long, 2 em wide, lance­ reate, bearing a solitary terminal spike; stamens elliptic, fleshy, cream-colored; syncarp 5 em separate, attached directly to the axis; filament long, 3 em in diameter, bearing about 400 4 times as long as the anthers. drupes (when young these are cream-colored), HOLOlYPUS: P. pentodon Ridl., Fl. Malay 12-13 mm long, 2-3.5 mm wide and thick, Penin. 5: 75, 1925. Also in the section is P. linear-oblanceoloid, 5-6-angled, the body 10-11 tetrodon Ridl. of Singapore. mm long, the summit rounded ; pileus 2.5-3 Insertion in Key On p. 225 (p.2 of reprint) for the second J, read J. Styles 1 (-2), often cartilaginous or bony, z. Styles-toothlike to broad subulate, entire or bifid; staminate spikes bearing stamens fascicled at apex of column ...... Rykia z. Style with cylindric base, then curving proximally and the margin ( 2- ) 3- 6-lobed with deltoid lobes; staminate spike solitary, bearing separate stamens attached directly to the axis . .__ .__ .__ ._.__ Multidens i 1C;m. .

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FIG. 186. Pandanus pentodon Rid!., from lectotype . a, Infructescence, X 1; b, drupe, lateral view, X 1; c, drupe, longitudinal median section, X 1; d, drupe, lateral view, X 4; e, drupe, longitudinal median sec­ tion, X 4; I, drupe and sryle, apical view, X 1; g, drupe apex and style, apical view, X 10; b, stigma, oblique view, X 10; i, drupe apex, style, and stigma, lateral view, X 10; j, leaf base, lower side, X 1; k, leaf middle, lower side, X 1; I, leaf apex, lower side, X 1; m, staminate inflorescence, X 1; n, stamens and axis, X 10. i i o S em. .

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FIG. 187. Pandanus tetrodon Ridl., from holotype. a, Syncarp, X 1; b, drupe, lateral view, X 1; c, drupe, longitudinal media n section, X 1; d-g, drupe, style, and stigma, apical view, X 1; b, drupe, lateral view, X 4; i, drupe, longitudinal median section, X 4; i. k, drupe, style, and stigma, apical view, X 4; I, stigma, proximal view, X 4; m, leaf base, lower side, X 1; n, leaf middle, lower side, X 1. Page 296 : Revision of Pandanus, 15. Malaya-ST. JOHN 353 mm long, the base pyramidal-serniorbicular, DIAGNOSIS OF HOLOTYPE : "Large bushy smooth, when dried somewhat yellowish; style plant; branches at apex 2.5 cm in diameter"; 1-1.5 mm long, apical, bony, yellowish, shining, leaves well over 97 cm long, 2.6 cm wide at its base short cylindric, then the rest sharply the middle, 2 ern wide near the base, coriaceous, curved proximally and flabellate with the mar­ l-ribbed, 2-pleated, in section low M-shaped, gins (3 -) 4-5 (- 6) dentate; stigma 0.5-1 mm about at the midsection with 25-26 secondary long, obdeltoid to flabellate, papillose, brown, parallel veins in each half, these visible at least on the distal side of the style; endocarp central , below throughout, tertiary cross veins visible in bony, pale brown, the upper lateral walls 1 mm outer half, transverse, forming oblong meshes, thick; seed 5-5.5 mm long; apical mesocarp mostly short, the blade ligulare, the apex not an empty cavern ; basal mesocarp fibrous up the preserved, the base amplexicaul, unarmed, be­ sides, fleshy within. ginning at 5 cm up the margins with prickles STAMINATE PLANT: Herbage not seen; stam­ 1.5-2.5 mm long, 5-10 mm apart, subulate­ inare .inflorescence dense, leafy bracted, the main tipped serrae, pale, with reddish tips ; the mid­ part 14 cm long; lowest floral bract about 30 rib below unarmed for at least lower ~' s ; near cm long, the upp er part green , foliaceous, 10 the midsection the margins with similar serrae, mm wide, with serrulations 0.2 mm long, 2- 5 1.3-1.5 mm long, 6-13 mm apart; leaf apex mm apart, the 10 ern basal part 19-20 mm wide, "acuminate," but not preserved; pistillate in­ folded, firm, apparently pale, with similar or florescence terminal, more or less concealed; pe­ smaller serrulations; median floral bract 7 ern duncle 19 cm long, 7 mm in diameter, 3-sided, long, 3 ern wide, navicular, elliptic, acute, ser­ sinuous, leafy bracted, bearing 1 syncarp, this rulate near the apex on margins and midrib 8.8 cm long, 5 cm in diameter, cylindric with below, firm, apparently pale; spike single, ter­ obtuse ends, bearing about 176 drupes, these minal, 7.5 cm long, 13 mm in diameter, cylin­ 22-24 mm long, 6-10 mm wide, 5-10 mm dric, dense; stamens distinct, attached directly thick, increasing upwards in size and the apical to the axis; filaments 4.2-4.4 mm long, anthers ones definitely the largest, cuneare-oblanceoloid , 0.8 mm long, narrowly oblong, bearing at tips obtuse, 5-6-angled, upper Y3 free, the sides a 0.15-0.2 mm subulate pro longation of the nearly smooth; style 3-4 mm long, bony, dark connective. shining brown, at base cylindric, then quadrate LECTOTYPE: Malaya, Johore, Ulu Kahang, and curved proximally, and all the lateral ones abundant along edges of Kahang river, 250 ft. with an oblique plane to the 2- 3-4-dentate aIt., 1 Ju ne 1923, R. E. Holttum 10,861 (SING) margi ns; stigma 1-2 mm wide or long, rugose, - the pistillate plant here designated as lecto­ of various shapes, filling the cavity proximal type. Mingled on the same sheet is a staminate of the style apex, with 2-4 (- 5) deltoid lobes; plant. endocarp centering in lower %, bony, dark brown; seed 6 mm long, obdeltoid; apical meso- : Pandanus tetrodon Ridl., Roy. Asiar , Soc., Jour. carp 6-7 mm long, a rounded cavity with a few 68 : 13-14, 1915; P. singaporensis Ka­ pale membranes; basal mesocarp fibrous and nehira, Jour. Jap. Bot. 14: 173-177, figs. fleshy. 6-8, 10, 1938, a superfluous name; non EXP ANDED DESCRIPTION OF PISTILLATE Barrotia tetrodon Gaud., Bot. Voy La PLANTS : Stems at apex 1.5-2.5 cm in diameter, Bonite, t, 13, figs. 1-8, 1841, an invalid brown, striate, covered with ascending adventi ­ name; non P. tetrodon Balf. f., Linn. tious rootlets 1-2 cm long; leaves 78 to more Soc. Bot. Jour. 17: 63, 1878, a name than 97 cm long, the tip gradually diminishing not accepted by its author; non P. tetra­ to a rrigonous subulate apex about 10 cm long, don Balf. f. ex Kan ehira, Bot. Mag. To­ and at base 4 mm wide, on the subulate apex kyo 52: 236-239, fig. 70, 1938, a later the margins and midrib below with serrations synonym of P. compresses M artell i 0.3-0.5 mm long, 2-5 mm apart; syncarp sin­ (1905) ; (sect. Multidens ) gle or with a secondary one below and 4.5 cm Fig. 187 long, ovoid. o

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FIG. 188. Pandanus immersus Ridl ., from holotype. a, Syncarp, X 1; b, drupe, lateral view, X 1; c, drupe, longitudinal median section, Xl; d-i, drupe, style, and stigma, apical view, X 1; j, drupe, lateral view, X 4; k, drupe, longitudinal median section, X 4; I, drupe apex and style, apical view, X 4; m, drupe apex, style, and stigma, proximal view, X 4; n, leaf middle, lower side, X 1; 0 , venation of leaf middle, lower side, X 4; p, leaf apex, lower side, X 1. Page 298: Revision of Pandanus, 15. Malaya-5T. JOHN 355

HOLOTYPE: Singapore, on the west coast road median and lower ones about half are entire near Pasir Panjang, Dec. 11, 1911, H. N. Ridley with .a single horn but the other half are bifur­ 15,465 (SING ). Holotype examined! cate and the 0.5-2 mm lobes often are wide­ SPECIMENS EXAMINED : Singapore, Botanic spreading; stigma 3.5-5 mm long; endocarp Gardens , Aroid Rockery, cult., 20 April 1936, (immature) oblanceoloid, nearly as long as the E. ]. H. Corner 30,993 (KEP, SING). body; mesocarp fibrous and fleshy. HOLOTYPE: Malaya, "Selangor, in the Labu SECTION Rykia River, forming dense thickets," May 1891, H . N. R [idley} , (SING). Holotype examined! Pandanus immersus Ridl., Asiat. Soc., Straits Branch, Jour. 41 : 48-49, 1903; Mater. Pandanus [oborensis Martelli, Soc. Bot. Ital., Bul. Fl. Malay Penin. 2: 224, 1907; Fl. Malay 302, 1904; P. muarensis Ridl., Mater. Penin. 5: 76, 1925; Martelli, Webbia Fl. Malay Penin. 2: 226-227, 1907, and 4(1): 18, 1913; (sect. Rykia) Fl. Malay Penin. 5: 77, 1925; (sect. Rykia) Fig. 188 Figs. 189; 190 NOM. VERN.: "mengkuang ayer." DIAGNOSIS OF HOLOTYPE: Aquatic, "the stem NOM. VERN. :"rassau.' rising but little above" the surface of the river ; DIAGNOSIS OF ISOTYPE : Leaves 1.3-1.45 m leaves "many feet long," 10.2 em wide, coria­ long, 4.2 em wide, coriaceous, paler beneath, ceous, green above, pale and glaucous beneath, I-ribbed, 2-pleated, at midsection with 43 sec­ the longitudinal veins strong, visible above and ondary parallel veins in each half, no visible . conspicuous below, abour 1 mm apart, at mid­ cross veins, ligulare, gradually tapering to a section with 70 secondary parallel veins in each 27 em subulate deltoid apex, this 10 em down side, the venation also consisting of tertiary 0.7 mm wide, the base amplexicaul, unarmed, cross veins 1-3 mm apart, these easily visible but beginning at 10-11 em on one side, and above and conspicuous below, the base not pre­ at 18 em on the other the margins with prickles served but the blade l-ribbed, 2-pleated, in sec­ 1.5-2 mm long, 4-11 mm apart, subulate, as­ tion M-shaped, Iigulare, narrowing upwards and cending, dark reddish; the midrib below begin ­ in the upper quarter tapering to a subulate del­ ning at 8 em with prickles 3-3.5 mm long, toid apex about 30 em long, and at a point 12-22 mm apart, stout- subulate, arcuate, re­ about 10 em down from the tip only 1.3 mm flexed; at midsection the margins with prickles wide; at about the midsection the margins with 1-1.5 mm long, 4-14 mm apart, stout subulate, thorns 2.5-4 mm long,9-18 mm apart, subulate arcuate, closely ascending; the midrib below but the base thickened, nearly straight, ascend­ with prickles 1.5-1.8 mm long, 5-45 mm apart, ing, pale except for the very tip which may be stout subulate, arcuate, reflexed or ascending; on reddish; near the tip the margins with serrae the subulate apex the margins with serrae 0.5 0.8-0.9 mm long, 1.5-3 mm apart, stout; the mm long, 1-3 mm apart; those of the midrib midrib below nearly unarmed; pistillate inflores­ below similar but 3-7 mm apart; peduncle 29 cence apparently emersed; peduncle more than mm long, 8 mm in diameter, 3-sided, slightly 8 em long, 15 mm in diameter, 3-sided, brae­ sinuous, leafy bracted; syncarp single, 6 em long, reate: syncarp perhaps a third mature, 9.5 em 3 em in diameter, narrowly ellipsoid (very long, 6 em in diameter, ellipsoid, 3-sided, bear­ young), bearing about 4,096 drupes, these (im­ ing about 1,440 drupes, these 17-19 mm long, mature) 11-12 mm long, 1.5-2.5 mm wide and 5-6 mm wide, 3-4 thick, the body 12-13 mm thick, 5-6-angled, upper ~ free, the body 4.5-5 long, arcuate spatulate, upper Y3 free; cuneate mm long, oblong-ellipsoid, papillose, with thick to the base; the pileus 7-8 mm long, the base longitudinal fibers; style 7 mm long, setiforrn, subsemiorbicular, minutely papillose; the style hard, shining and yellow; stigma linear, proxi­ 4-6 mm long, hornlike, mostly gently curved mal, brown, as long as the style; endocarp and proximally, the apical ones unbranched, of the seed not yet formed, but inside the ovary wall i a cm ,

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FIG. 189. Pandanus [oborensis Martelli, from isotype. a, Syncarp, X 1; b, c, drupe, lateral view, X l ;d, drupe, lateral view, X 4; e, drupe, longitudinal median section, X 4; l, drupe apex and style, apical view, X 4; g, style and stigma, proximal view, X 10; b, leaf base, lower side, X 1; i, leaf middle, lower side, X 1; j, leaf apex, lower side, X 1. o c fi o..

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FIG. 190. Pandanus [oborensis Martelli, from Corner 28,161. a, Staminate inflorescence, X V3 ; b, spike, X 1; C, staminate column , scale, and stamens ,X 10; d, leaf base, lower side,X 1; e, leaf middle, lower side, X 1; I, leaf apex, lower side, X 1. 358 PACIFIC SCIENCE, Vol. XVII, July 1%3

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FIG. 191. Pandanus penangensis Rid!., from holorype. a, In frucrescence, X 1/5; b, drupe, lateral view, X 1; c, drupe, longitudinal median section, X 1; d, drupe apex and style, X 1; e, drupe apex and stigma, p roximal view, X 4; t. leaf base, lower side, X 1; g, leaf middle, lower side, X 1; b, leaf apex, lowe r side. X l. Page 302: Revision of Pandanus, 15. Malaya-ST. JOHN 359 of tissue and heavy longitudinal fibers there is "Stare di Johor. Penis. Malese, no. 11326 Ridl, a single large caviry, with indefinite traces of (H. Becc.)." The specimen above cited is an inner tissue. isotype in the herbarium in Singapore. It bears EXPANDED DESCRIPTION FROM ALL PISTIL­ the data: "Malay Peninsula, Johor, Bukir Wilgu, LATE PLANTS EXAMiNED: Shrub, to 6 m tall, Sungei Pau, 1902, Fox 11,326." In his Materials upper branches 3-4 cm in diameter, dark brown, for a Flora of the Malayan Peninsula 2: 226, forming dense thickets, with a few thornlike 1907, Ridley accepted Martelli's species, and axillary rootlets 4-6 mm long, divergent; leaves cited the single collection, "Muar: Bukit Wilgu, 1-1.8 m long, 2.9-4.1 ern wide; syncarp solitary, Sungei Pauh (W. Fox) 11326." Later, Ridley pendulous, 12-16 ern long, 6-10 cm in diame­ in his Flora of the Malay Peninsula 5: 77, 1925, ter, ellipsoid. treated this plant as P. johorensis sensu Ridl., DESCRIPTION OF STAMINATE PLANTS: Brae­ not of Martelli, and redescribed it as the new teal leaves up to 80 cm long, 3.5 cm wide, coria­ species P. muarensis Ridl . For this he had the ceous, apparently pale, Iigulate, the upper Y3 holotype: "Muar, Sungei Pauh (Fox)." What­ long tapering to a trigonous subulare apex, rhis ever misconception Ridley thought there was, 10 ern down 2 mm wide, the lower Y3 wholly he had no right to rename the species using the unarmed, at midsection with 51 secondary veins same holotype. Certainly the collection Fox in each half, no visible tertiary veins; near the 11326 from Bukit Wilgu, Sungei Pauh, Johor, apex the margins and midrib below with subu­ is the same as [Ridley} 11,326 with the same late serrulations 0.5-0.7 mm long, 2-6 mm locality data. The number is a Singapore Her­ apart; staminate inflorescence 45 ern long, with barium number, as the staff members were re­ 8 spikes, these 15-25 cm long, 12-15 .mm in quired to use this single series of numbers, not diameter, densely flowered but somewhat in­ their own personal collection numbers. The hol­ terrupted; flowers sweet and sickly scented ; fila­ otypic specimen in the herbarium at Firenze was ment column 4 mm long, stout, clavate, bearing not a duplicate distributed from Singapore. an apical parasol-like scale 3-5 mm in diameter, Rather, it is a fragment consisting of about two the apex hispidulous with hairs 0.3-0.4 rom score of the drupes placed in a paper pocket on long; free filament tips 0.5-2 mm long, pendent which in Martelli's hand is written the data. from the lower surface, glabrous or sparsely These fragments were taken from the syncarp hispidulous; anthers 10-16, and 0.8-1.3 mm of the isotypic specimen, then in Calcutta , now long, mostly cylindric, bearing an apical mucro in Firenze. It lacks stem, leaves, inflorescence, 0.2 mm long, formed of a projection of the and syncarp. Nevertheless, the actual holotype connective (description from Corner 28,161). is this fragmentary specimen, Fox (or Ridley) , ISOlYPE: Malaya, Johor, Muar, Bukir WiIigu; once in Calcutta, and now in Firenze. However, Sungei Pauh, 1902, Fox 11,326 (SING). Isotype there seems no doubt but that P. muarensis Ridl. examined! is a typonym and must be placed in the syn­ SPECIMENS EXAMINED: Malaya, Johore, Ma­ onymy of P. johorensis Martelli. The isotype in wai, generally flowering of staminate and pistil­ Singapore has a syncarp with immature drupes late on riverside, flowers visited by bees and that have not matured their endocarp and seed: beetles, May 13, 1934, E. ]. H. Corner 28,161 (SING). . Pandanus penangensis Rid!., Asiat, Soc., Straits DISTRIBUTION : Malaya and Sumatra, form­ Branch, Jour. 41: 50, (1904) = 1903; ing dense thickets in estuaries, forming a zone F!. Malay Penin, 5: 81, 1925; (sect. beginning where Nipa stops, in tidal fresh wa­ Rykia) ter. Plants with similar leaves have been seen Fig. 191 from Borneo. DISCUSSION : P. johorensis Martelli is a mem­ DIAGNOSIS OF HOLOlYPE : "Tree 7 m. tall, ber of the section Rykia. It was briefly pub­ 10 ern, in diameter"; leaves 2 m or more long, lished in 1904 by Martelli as only a few phrases near the base 12 cm wide, when young marbled in a key to certain species. His holotype was: light and dark green , beginning at 15-16 cm 360 PACIFIC SCIENCE, Vol. XVII, July 1963 the margins with prickles 1.3-1.5 mm long, 4- 7 proximally almost to a right angle; stigma 4-5 mrn apart, pale, subulate, flat appressed ascend­ mrn long, lanceoloid, papillose, brown; endocarp ing ; the midrib below sharp salient and with 32-33 mm long, narrow ellipsoid, bony, ebony­ similar ascending prickles, the secondary parallel colored, the inner surface shining, the lateral veins conspicuous and in lower part 69 in each wall 1-1.5 mm thick; seed 15-17 mm long, half, and except at base the tertiary cross veins 4-6 rnrn in diameter, ellipsoid; apical and basal prominent on both sides, transverse or oblique mesocarp fibrous and fleshy. . and forming short, rather oblong meshes, mid­ EXPANDED DESCRIPTION FROM ALL SPECI­ section not seen; at 45 cm down from the apex' MENS EX AM INED: Leaves to 4 m long, the mid­ the blade 11.5 cm wide, the apex abruptly nar­ section 10 cm wide, the margin with prickles rowed to a subulate tip about 5 ( ? ) cm long, the 2.8- 3 mm long, 5-1 4 mm apart, stout, heavy margins with subulate serrae 1.5- 2.5 cm long, based, arcuate subulate, appressed asc en d i ng, 3- 10 mrn apart ; the midrib below with serrae brown; pistillate inflorescence of 4-5 syncarps, 0.8-1 mm long, 7-20 mm apart; drupes 42-47 each with numerous drupes; syncarps 15-20 cm mm long, 7-9 mm wide, 6-7 mm thick, 5-6­ long, 9.5-10 ern in diameter, ellipsoid. angled, the sides smooth, gently curved, prob­ HOLOTYPE: "Malaya, Penang Hill from base ably only the pileus free, the body 37-40 mm to the top, in woods." The specimen is labeled : Iong, d r upes and pilei shedding separately; Malaya, Pulau Penang, Fort Hill ; 1900, H. N . pileus 9- 11 rnm long (o r if measured along Ridley ( SIN G). Holotype examined! the curve of the style 13-16 mrn long ), 10-13 SPECIMENS EXAMINED: Malaya, Pulau Pe­ mm wide, 8-10 mm thick, 6-angled, broad nang, Government Hill, common on the hill ovoid-pyramidal, the tip proximally curved; top, but not often in fruit, 29 July 1907, 1. H . style 5-7 mm long, heavy subulate, curved Burki1l 2,666 (SING ) .