The Genus Rhopaloblaste (Palmae)

r9701 MOORE: GENUSRHOPALOBLASTE 75 The Genus Rhopaloblaste(Palmae) Hanor-rE. Nloone,Jn.*

Three species of attractive palms are time to one or both generaare found to cultivated ln botanical gardens and col- belong to the genus Heterosrythe and lections under the respective names Pty- are so assigned in the list of excluded choraphis &ugtrsta (S. Kurz) Beccari, P. specieson:page 90. The remaining spe' singaporensi.s (Beccari) Beccari, and cies form a homogeneousunit including Rho'p,Ioblaste cerarnica (Miquel) Bur- the type-species of both Ptychoraphis ret (R. hexandra Schef{er). A relation' and Rho'paloblaste. For this genus, the ship betweenthe genera has been appar- earlist name of Rh'opaloblasteis used. ent since 1885 when Beccari established The following characteristics unite the the genus Ptychoraphis, including the speciesand, in combination, set them off two sneciesmentioned above and a third fiom other genera of the subfamily Are- which was later referred to Heterospathe coideaetribe Clinostigmateae.The leaf- by Beccari himself. Ptychotaph,is was sheaths form a distinct, tubular crown- first distinguished fuom Ptychosperrnd shaft, the pinnae have only the midnerve and Rhopal,oblasteonly by the deep s'ul' elevated and prominent above, and the cation of the seed along the hilum, but indument of upper and lower surfaces of in 1886 (Malesia 3: 109) the genuswas the petiole and rachis is distinctive-pel- {urther separated hom Rhopaloblaste by tate, lacerate-fimbriate-marginedinter- "the stamen-filamentsconnate at the base locking scalesbelow, basifixed, twisted, and not free, by the form of the pistillode membranous scalesabove, at least when in tho staminate flowers, by the different young. The short-pedunculate, infra- mode o{ vascular branching in the seed- foliar inflorescence has 2 caducous coat, by the form o{ the embryo, after all bracts of which the upper is enclosed by the seed sulcate along the raPhe within the lower (prophyll) and the usu- [transl.]." Sincethat time, there has been ally twice-branchedbasal branches are no detailed study of the two genera. The characteristically abruptly divaricate most recent separation (Beccari & Pichi' from the rachis immediately above the Sermolli, Webbia Il: 18, 66-68, 1955) insertion of the upper bract, Rhopalo' relies on symmetry of fruit, presence or blaste singaporensisexcepted. The outer absenceof a sulcate hilum, and number sepal in both staminate and pistillate of stamens. flowers develops much before and en- When one examines tlle sPeciesnow folds a large part of the remaining peri- included in Ptych'oraphzsand Rhopalo' anth in bud in a fashion unusual for blastein the light of, new collections from palms. Staminate flowers at anthesis are New Guinea and the Solomon Islands, symmetric or subsymmetric. The fruit and and in the light of a better understanding is often rostrate, lacks fiber-sclereids cells in the mesocarp, dries of the importance of the structure of the tannin smooth, and has an apical stigmatic inflorescenceand histology of the fruit- residue; both endocarpand seedare im- coat in classification, two groups of taxa pressedor sulcate along the linear hilurn. emerge, Several speciesreferred at some Seedlings have two bladeless sheaths followed by a pinnate eophyll or first *From work relating to National Science Foundation grants GB-1354, GB-3528' GB-7758. leaf. 76 PRINCIPES lVol. 14

Those speciesreferred to Heterospffihe nerve and one or more secondarl nerves differ in lacking a well-defined crown- on each side prominent belon'. minutely shaft, in having pinnaewith usually three brown-puncticulateand at least the mid- nervesprominent above,at leastbasally, nerve with prominent dull-broln. basi- in having a long-pedunculateinflores- fixed or medifixed,twisted, membranous cencewith branchesnot abruptly divar- scalesbasally or throughout. icate and with bracts markedly unequal, In{lorescencesborne below the leaves; the upper much exceedingthe lower, and peduncleshort; bracts 2, caducous,the in having fiber-sclereids in the outer lower ancipitous and enclosing the up- layer of the mesocarp. A fuller charac- per, both usually more or less lepidote- terization oI Heterospailrcis to be found tomentoseat least when young; rachis in Principes 13: 99-100, 1969. short to prominent but as long as or longer than the peduncle;basal branches Rhopaloblaste Scheffer, Annales du usually abruptly divaricate-spreadingat Jardin Botanique de Buitenzorg an angle of about 90o to the rachis (in I:137. 1876. R. singaporensis o{ten at an acute Type-species:R. hexandraScheffer angle); bracts subtending the branches (= R. ceramica (Miquel) Bur- often prominent; bracts subtending the ret) triads prominent or not; bracteolessurrounding the pistillate flower subequal PtychorapltisBeccari, Annales du Jardin or unequal, prominent and sepal-like. Botanique de Buitenzorg 2: 90. Flowers borne in triads of two stami- IBB5. nate and a pistillate in the lower portion Type-species: P. singaporensis of or nearly throughout the rachillae; (Beccari) Beccari staminateflorvers symmetric or subsymmetric at anthesisbut in bud the outer Solitary or cespitose,unarmed, monoe- sepal prominent and largely enfolding cious, low to tall palms with stems often the remainder of the perianth; sepals3, enlarged at the base but uniform and broadly imbricate at anthesis,rounded, relatively slenderabove. more or less gibbous and keeled dor- Leaves reduplicately pinnate; sheaths sally; petals3, valvate: stamens6-9, the tubular, forming a crownshaft; petioles filamentsvery briefly connatebasally or short to elongate,rounded below, chan- essentially distinct, strap-shaped,nar- nelled above, rachis rounded below, an- rowed and prominently inflexed at the gled above toward the apex, the sheath, apex in bud, the anthers narrowly ellip- lower surface of petiole and rachis usu- tic in outline, medifixed, only emargi- ally densely lepidote with peltate scales nate apically and basally,the connective having lacerate-fimbriate interlocking prominent the entire length of the anther, margins,the upper surfaceof petiole and dehiscenceby longitudinal lateral slits; rachis usually denselyclothed with basi- pistillode fixed, twisted, entire or lacerate, mem- conic to columnar and more or branous scaleswhich persist about the less angled, the apex briefly three-lobed bases of pinnae and where protected and sometimessomewhat expanded; pis-' elsewhere; pinnae spreading or pendu- tillate flowers broader than high in bud lous, with a distal pulvinus at the base, and with the outer sepal usually enfold- linear, acutely to acuminately and ing the remainder of the perianth as in obliquely bifid-praemorse,with only the the staminate; sepals3, broadly imbri- midnerve elevated,prominent, and often cate, rounded; petals 3, broadly imbri- lepidote near the base above, the mid- cate basally, the short valvate apices 19701 MOORE: GENUS RHOPALOBLASTE

I. Rhopaloblaste elegd,ns.a, portion of rachilla with triads X 1; b, triad with {lowers removed X 2; c, portion of rachilla with paired and solitary staminate flowers X 1; d, scars and bracteoles of paired staminate flowers X 2; e, staminate flowers at anthesis N 2; f, staminate bud X 4; g, staminate bud in vertical section X4; h' staminate sepals t 4; i, staminate petal X 2; j, stamens in 3 views X 2; k, pistillode X2; 1, pistillate bud X 4; m, pistiliate bud in vertical section X 4; n, pistillate sepals N 4; o, pistillate budwith sepals removed X4; p, pistiilate petal X 4; q, pistii and staminodes X4; r, staminode X B; s, fruit t I; t, fruit in vertical section X 1; u, fruit in cross- section )( 1; v, endocarp X I; w, operculum | 2; x, seed in lateral, abaxial and adaxial views X LL From material of Moore & Whitmore 9310 preserved in liquid. erectand scarcelyexceeding the sepalsat cells,with flattenedlongitudinal fibers in anthesisbut the petalsin fruit generally one or usually more than one layer nearly twice as long as the sepals;stami' against the yellowish, fragile endocarp, nodesmostly 6, theseobtuse, more or less this impressedover the hilum and with deltoid, membranous, o{ten united in a round basal operculum; seed brown, pairs or irregularly united or united in a with lightly to deeply impressedhilum membranous, lobed ring; Pistil uni' the length of the adaxial side, vascula- locular, uniovulate, the ovule (in R. ce' ture anastamosed, endosperm deeply ramica) hemianatropous,attached adaxi' ruminate,embryo basal,large. ally (in the ventral angle) and pendulous Seedlingwith 2 bladelesssheaths, then from the top of the locule,broadest in its a pinnate eophyll and successiveleaves lateral axis; stigmas erect to recurved (in R. augusta, R, Brassii, R. singapo- betweenvalvate apices of petalsat anthe' rensis). sis. Chromosomecomplement: not known Fruit orange-yellowto red, ovoid or Distribution: rain forest in Nicobar ellipsoid to subglobose,with apical stig- Islands, southern Malay Peninsula and matic residue; exocarp smooth; meso- Singapore, Molucca Islands, New carp lacking fiber sclereids or tannin Guinea,Solomon Islands. 7B PRINCIPES tVol. 14

A Key to the Species of Rhopaloblaste I. Stemscespitose, slender, to 4 m. high; leaveswith elongatepetiole 60 cm. or more long and spreading pinnae; inflorescencedecurved, short, less than 30 cm. long, simply branched or with the basal branchesonly furcate and borne at an acute angle with the rachis, the approximately 5 branches or rachillae transverselyrugose, glabrescent or coveredwith simpleto stellate,brown or pale trichomes; fruit ovoid, orange-yellowto red, 13*14 mm. long including the prominent rostrum I.5*2mm.long, 9-10 mm. in diam. Malay Peninsula,Singa- pore. ------: - R. singaporensis I. Stemssolitary, mostly more than 10 m. high, 10 cm. in diam.; leaveswith short petioles less than 50 cm. long; inflorescencesspreading, the lower branches divaricate from the rachis at about a 90o angle and once- or twice-branched into glabrous or lepidote rachillae that are not transverselyrugose when dry; fruit red or orange-red. 2. Fruit small, 10 mm. long, 8 mm. in diam.; perianth 4 rnm. long in fruit. New Guinea.------R. Led,ermanniana 2. Fruit larger, 2.A4.5 cm. long, I.0-1.8 cm. in diam.; perianth 6-10 mm. long in fruit. 3. Fruit subglobosewhen fresh, 2.6 cm. high, 2.3 cm. in diam.; staminate flowers subsymmetric, the petals slightly angled, 6-7 mm. long, about three times as long as sepals; lower branches of inflorescencemostly once-branched.Solomon Islands. R. elegans 3. Fruit ellipsoid or ovoid-ellipsoid to obovoid-ellipsoid,distinctly longer than broad; staminateflowers (where known in the mature state) symmetric; lower branchesof the inflorescencemostly twice-branched. 4. Triads subtendedby a prominent, more or less reflexed,liplike bract and with a prominent, elevated,and thickenedupper margin; pinnae pendulous; rachillae ca.3 mm. thick at base; fruit ellipsoid, 2.2-2.6 cm. long including a prominent rostrum 2 mm. high, 1.0-1.4 cm. in diam.; petals6-7 mm. high in fruit; staminodesunited in a membranous 6-lobedring. Nicobar Islands. - - R. augusta 4. Triads subtendedby a low bract and without a prominent elevated and thickenedupper margin; pinnae and fruit various. 5. Pinnae pendulous; staminateflowers 6-7 mm, high; fruit 3.0-3.5 cm. Iong, 1.8 cm. in diam., with perianth I cm. long; rachillae 5-7 rrlm. thick at base,glabrous; staminatepetals not appearing puncticulate; pistillate buds ca. 4 mm. high, 6 mm. wide, with petals scarcely or not evident at staminateanthesis; staminodes more or lessunited in a lobed membranousring. Molucca Islands. R. ceramica 5. Pinnae spreading; staminateflowers 4-4.5 mm. high. 6. Fruit 2.0 cm. long, I.0-I.3 cm. in diam., with perianth 6-7 mm. high; rachillae 2-3 mm. in diam., densely lepidote with pale to red-brown, thickened and often branched trichomes; staminate flowers 4-4.5 rnm. long, the petals appearing minutely puncticulate; pistillate buds ca. 3.5 mm. high and wide with valvate apices of petals evident at staminate anthesis; staminodesmore or lessdistinct. New Guinea.---- R. Brassii 19701 MOORE: GENUS RHOPALOBLASTE 79

6. Fruit 3.5 cm. long, 1.8 cm. in diam., with perianth I cm. high; rachillae 5 mm. in diam. at middle. New Guinea. R. d'yscrita

Rhopaloblaste augusta (S. Kurz) H. oermost: lowermost branches twice' E. Moore,Lr. nou. tra.rched into rachillaeto 36 cm. long, 3 mm. in diam. at base (when dry), these Areca augustaS. Kurz, Journal of Bot- drying lined but not transverselyrugose, any 13: 331,pl. I70. 1875. glabrous except for floccose scales at Ptychoraphis augu ta (S. Kurz) Bec- margins o{ triads; bracts subtendingthe cari, Annales du Jardin Botanique triads very prominent and concealingthe de Buitenzorg2: 90. 1885. buds in young stages,to ca. 2 mm. long rounded, and Solitary, to 30 m. high or more, 30 on inner side, prominent, of the triad cm. in diam. at base,trunk brown or be- decurved,the upper margin (when at matur- coming gray in age,clearly ringed. elevatedand liplike dry) Leavesabout 10, to 4 m. long; sheath ity; bracteolessurrounding the pistillate 2.5 mm. about 60 cm. long, gray-brown lepidote- flowers subequal,imbricate, to pairs tomentose;petiole very short, 10-11 cm. high, bracteoles of the staminate long; rachis denselypale-lepidote below orominent. green- when young with interlocking lacerate- Staminate flowers symmetric, fimbriate, brown-centeredscales but be- ish outside,whitish inside, 6-7 mm. long min- coming merely brown-puncticulate in at anthesis; sepalsca. 3 mm. long, anthe- age, glabrescentabove in age but with utely ciliolate; petalsspreading at floral rusty or dark-brown to pale, twisted and sis, ca. 5 mm. long, attachedto a fila- often lacerate, membranous scalesper- receptacleI mm. long; stamens6, pistillode sisting in the anglesat the baseof pinnae ments and anthers yellow; stamens, or elsewhere when protected; pinnae conic, about half as long as the 4.5-5 90-100 on each side, pendulous, to 70 yellowish: pistillate flowers green, cm. long, 3 cm. wide, glabrous, dark- mm. long when dry, to 6 mm. long when green,and shining above,very minutely fresh; sepals3-4 mm. long, very strongly and brown-puncticulatealong all the nerves imbricate, the outer almost shell-like mar- and the midnerve with dark brown or somewhatridged, all with ciliolate only dull-brown, twisted, basifixed, membra' gins; petals 4-4.5 m:rl'. long, with nous scalesto 3 mm. long below. the short, erect, valvate apices exserted In{lorescencesusually 3-5, ringing the above the sepalsat anthesis,but becom- trunk below the crown; lower bract gray- ing 7 mm. long in fruit; staminodes brown; peduncleca. 5 cm. long; rachis united in a membranous,more or less densely brown floccose-lepidote,ca. 37 3-lobed ring; stigmas white, recurved cm. long with ca. 15 branches,the lower betweenthe apicesof petals. two branches divaricate at about 90o Fruit ellipsoid, orange-red, 2.2-2.6 from the rachis immediatelyabove the cm. long including prominent beak 2 insertion of the upper bract, the third mm. high, 1.0-1.4 cm. in diameter; similarly divaricate abaxially between mesocarpthin, with flat fibers longitudi- the first two, remaining branchesmostly nally appressedagainst the fragile endo' decussatelyarranged laterally exceptthe carp, this and the seed impressed over uppermost few, each branch subtended the hilum; seed ellipsoid, rounded at by a bract'ranging from very prominent, apex, 15-16 mm. long, 9-10 mm. in acute,and 6-7 cm.long at the lowermost diameter,vasculature much anastamosed. to ca. 1 mm. long and liplike at the up- Seedling with 2 bladeless lear,es, the PRINCIPES

2. Rhopaloblaste augusta growing in the Botanic Gardens at Roseau, Domincia (Bail.ey 7)l'). Photo by L. H. Bailey. r9701 MOORE:GENUS RHOPALOBLASTE BI eophyll with ca. 7 pinnae on each side, sheath 62 70 cm. long, densely lepi- rachis with shining castaneousmembra' dote'tomentose;petiole 15-20 cm. long, nous scales. densely appressed lepidote-tornentose Distribution: Nicobar Islands. below, densely lepidote with basi{ixed Specimensexamined: NICOBAR IS- membranousscales above; rachis 2.9-3.8 LANDS. Kauonra: Feb. 1875, S. Kurz m. long, denselylepidote or dark-punc- s. z. (K, isotype; photo BH, neg. 4913- ticulate below, densely lepidote above 14) ; locality and date not stated, E. H. with basifixed,twisted, castaneous, mem- Man s. n. (Fl, Hb. Becc.,Rec. Oct. 1886; branous, errtire to lacerate scales or photo BH, neg. 3878-80). CULTIVA- densely puncticulate in age; pinnae TED. IlvloNBsrl: Java, Botanic Gar' about 90 on eachside, to 63 cm. long,2.5 dens,Buitenzorg IBogor] XII. E. 89, sub cm. wide, the midnerve prominent and no. 407 (FI, Hb. Becc.;photo BH, neg. elevatedon the upper surface, more or 3BB1-83), same,April-May, 1936, C. X. less densely membranous-lepidote,at Furtado, S. 1. A/. 30946 (BH). CusA.: leastnear the base,with scaleslike those Atkins Garden, Harvard University, of the upper surface of the rachis, the Soledad,Cienfuegos, 22Feb.1952, H. E. midnerveand other nerveson lower sur- Moore 6088 (BH). Donrlutca.: Botanic face brown-puncticulate, the midnerve Gardens,Roseau, Feb.-Apr. 1922, L. H. and sornetimes the secondary nerves Bail,ey711 (BH). Pa.weui.: Canal Zone also with basifixed or medifixed, dull- Experiment Gardens, Summit, Feb. 15, brown. twisted. mem.branousscales near 1936, W. R. Lindsay s. n. (BH). TnItrr- the base. l,l.o: Royal Botanic Gardens, Port-of- Inflorescenceto 95 cm. long, 164 cm. Spain, Mar. 1922, L. H. Bailey 687 wide acrosslower branches,Iower bract (BH) ; St. Clair Experiment Station, ca. 37.5 cm. long, lowest branchesto B0 April, 1923, W. E. Broadway 7030 cm. long, twice branched, ultimate ra- (BH). chillae to 55 cm. long, drying angled Rhopaloblaste augusta is a palm that but not rugose, 2.5-'4 mm. in diameter deservesto be more widely grown for its at base,rather denselybeset with small, foliage attractive and bright {ruits. The thick, often branched, pale or red-brown generalhabit of the species is well exem- to black trichomes,these often expanded plified by the individuals in Fig. 2, apically and irregular in shape; bracts though in its native islands trunks may subtending triads low, rounded, ca. f attain a much greaterheight. mm. high; bracteoles surrounding the Rhopaloblaste Brassii H. E. Moore, pistillate flowers subequal, imbricate, sp.nou. l-1.5 mm. high. Ab speciebus omnibus aliis Rhopalo- Staminateflowers (Brass 7135) 4-4.5 blastis dilfert fructibus 2 cm. longis, 1- mm. high, symmetric; sepalsca. 1.6 mm. I.3 cm. in diam., rachillis non rugosis high; petals with very minute depres- dense lepidotis, floribus masculis 4--4.5 sions giving the appearance of being mm. Iongis. puncticulate; stamens6; pistillode conic, Stem solitary, dark brown or black, about as long as filaments. concentrically ringed, sometimes with residual leaf-sheathsbelow the crown- Pistillate buds at staminate anthesis shaft, 8.5-10 cm. in diam. at base and (Brass7135) ca. 3.5 mm. high with val- 5-6 cm. in diam. below the purplish- vate apicesof petals evident; sepals2-3 sreen crownshaft. mm. high in fruit; petals 5-7 mm. high "rather Leaves 7-LI, flat spreading;" in fruit; staminodesca. 4, distinct. 82 PRINCIPES tVol.14

Fruit red, ovoid- to obovoid-ellipsoid, ca. 2 cm. long including short excentric beak, 13-14 mm. in diam.; seed13 mm. Iong, 10 mm. in diam., the hilum shal- lowly impressed. Seedlingwith pinnate eophylls. Vernacular name: Kuwehleh (Ome language, Wantipi, fide Darbyshire & Hoogland) Uses.: used for arrow points (bird arrows) but part not specified. Distribution: New Guinea Specimensexamined: NEW GUINEA. Wnsr InLtw; frequent in Agathis forest, 4 km. southwest of Bernhard Camp, Idenburg River, 900 m. alt., March, 1939. L. I. Bross13305 (A, holotype; L, isotype, photo BH neg 447I); common in rain-forest of moist alluvial flats, Bern- hard Camp,Idenburg River, April, 1939, L. l. Brass13809 (A) ; in Agathis Iorest, Dalman,45 km. inward from Nabire, 500 m. alt., Mar. 2, 1940,R. Kanehira & S. Hatusima 12131 (A). TsnnrronY Nnw 3. Rhopaloblaste Brassii (Brass 7145). Note GurNnl: Sepik District; Aitape Subdis- the slender stem, spreading pinnae, and twice- trict, near Wantipi village (on Bliri branched, sharply divaricate lower branches of the inflorescence. Photo by L. Brass. River) in tall forest on foothills, alt. ca. J. 800 ft., 3 Aug. L961, P. I. Darbysh'ire & R. D. Hoogland, 8373 (BH). PApu.q'. Leilermanniana with which Burret had Palmer River, 2 mi. below junction with earlier identified the last two numbers. Black River, sporadic in ridge {orest Lacking the type oI R. Led'ermanninna, substage,alt. ca. 100 m., June 1936,L. I. which is presumed to have been de- Brass7135 (A) ;71354 (A). stroyed, or material from the type-local- None of the collectionscited is com- ity, I am unwilling to equate the mate- plete and most are not truly comparable, rials cited here with that speciessince the being in various stages of flower and fruit is twice as long. There is, more- fruit, yet correspondenceis suf{icient so over, no way of assuring from the de- that I consider them to represent the scription oI R. Ledermanniana alone same species. The type has mature fruit, that it is truly a Rlropaloblaste and not a Brass 13809 has immature fuuit, Kane- species of Hetero'spathe, while there is hira & Hatusima 12131 has pistillate the suggestionthat it may be Hetero- buds, Darbyshire & Hoogland' 8373 has spathe in the nature of the pistillode of a portion of inflorescence in very young staminateflowers, the size of fruit, and bud, Brass 7135 and 71354 have stami- the stature of trees. I have therefore nate flowers, pistillate buds, and unripe described the above as new, drawing fruit which measuresless than that in the upon all materialsbut specifyingas holotype but more than that describedfor R. type fruiting material from West Irian. r9701 MOORE:GENUS RHOPALOBLASTE 83

Suchcontrasts as can be made with other Pistillate flowers green, ca' 4 mm. speciesappear in the key to species. high,6-7 mm. wide; sepals4 mm. high in fruit; petalsca. 10 mm. high in fruit; Rhopaloblaste ceramica (F. A. W. staminodesunited in an irregularly 6- Miquel) Burret, Repertorium Speci' lobed membranousring. erum Novarrm 24: 288. 1928. Fruit scarlet,3.0-3.5 cm. long, 16-18 ellipsoid'ovoid; seed ca. Bentinckia ceramicaF. A. W' Miquel, mm. in diam., long, 1.3 cm. in diameter, acut- De palmis Archipelagi Indici B. 2.I cm.. over the hilum. 1868. ish, impressed Seedlingnot described. Rhopaloblaste hexand'ra Scheffer, Distribution: Bachan (Batjan) , Ce- Annales du Jardin Botanique de ram in the Molucca Islands. Buitenzorg1: 156.1875. Specimensexamined. CULTIVATED. Solitary, to 15 m. high or more, trunk SrNsA.pono.Botanic Gardens, 11 Dec. (BH) brown or becominggrayish. 1963,H. E. Moore,1r.9039,9077 ; (BH) Leavesto 3 m. long or more; sheath 23 Aug. 1925,M. Nur s. n. ; 4 Oct. (BH) denselylepidote-tomentose; petiole short; 1929, M. Nwr s. n. ; INool,rBsrA.: rachis with 80-90 pinnae on each side, Java; Giardino Botanico di Buitenzorg, (FI, Becc.; denselylepidote-tomentose below, becom- Maggio 1878,O. Beccari Hb. ing brown-puncticulatein age, clothed photoBH, neg.3854-61). with rusty or dark-brown, twisted and Rhopaloblaste cera,tnica, often culti- often lacerate scalesabout the basesof vated under the name R. h.exandra,has the pinnae and along the central ridge been studied by me only from cultivated above;pinnae pendulous, to 1.1m. long, specimens. The type oI R. h'exan"drawas 2.5 cm. wide, glabrous above, densely supposedly from the island of Bachan and minutely brown-puncticulatebelow (Batjan) in the Moluccaswhile that of R. but membranousscales on midnerve ap- ceramica was from Ceram. Burret con- parently lacking. sideredthe two speciesidentical and took Inflorescencesseveral; peduncleca. B up the earlier epithet. I have accepted cm. long; rachis ca. 55 cm. long, densely his work and the similar conclusionsof tawny-tomentoseto glabrescent when Beccari (ex Martelli, Nuouo Giornale young, with ca. 16 branches,the lowest Botanicoltaliano, series2, 42:32. 1935) ca. 45 cm. long, twice-branched into in the absence of authentic material. stout, glabrous rachillae to ca. 45 cm. Completecollections from the wild state Iong,5-7 mm. in diameterat base; bracts are much to be desired. triads low, not prominent, subtendingthe Rhopaloblaste dyscrita H. E. Moore, margin of the triad not ele- the upper notn.nou. vatednor liplike; bracteolessurrounding the pistillate flowers markedly unequal, Rhopaloblastemicrantha Burret, Not- the larger to 2.5 mm. high, bracteolesof izblatt Berlin 15: 10. 1940; not ft. (Beccari) the staminatepairs small' micrantha Bentham & J. Staminate flowers symmetric, green, D. Hooker ex B. D. Jackson,Index to ca. 7.0 mm. long at anthesis; sepals Kewensis2: 7I3.1895. 3 mm. high and wide; petalsca. 6'5 mm. Stemsto 20 m. high, 30 cm. D. B. H. long, adnateto floral receptaclefor about Leaves with rachis softly white-floc- 2 mm.; stamens6 or 7, yellowish; Pis- cose-tomentosebelow, soon glabrescent tillode orange,more than half as long as above; pinnae numerous, spreading, stamens. rigid, linear, to 95 cm. long, 3.3 cm. wide, B4 PRINCIPES lVol. 14 slightly curved below above the base,the obvious that the pendulouspinnae asso- apex acuminate,bifid, obscurelybilobed, ciated with the type-speciesand R. au- the upper lobe produced,nerves minutely gustd ate perhaps exceptional and that dark-puncticulate below and the mid- spreadingpinnae may be more the rule. nerve with large membranous scales. It has also becomeobvious that the stam- Inflorescencelarge, broadly and divar- inodes in Rhopaloblaste are variously icately twice- (or ?thrice-) branched; united in irregularly lobed rings or in rachis 45 cm. long or more, broadly pairs, or distinct. I have not been able angled-rounded,4 cm. wide at base or to locate authentic material of the speci- more; primary branches subtendedba- mens iited by Burret when he described sally by a short, broadly triangular- R. micrantha (Clem,ens 7987, type; rounded bract, fruiting rachillae numer- Clemens8297), but from the description ous, robust, to 85 cm. long, 5 mm. in (translatedand adaptedabove) it seems diam. at the middle, bearing loosely spi- likely that the taxon represents a true ralled triads in the lower two-thirds, Rhopaloblaste. Since I cannot equate the paired staminate flowers in the upper description with that of any other species, third; bractssubtending the triads lightly since no material is available,and since produced and broadly rounded; bracte- Burret's epithet is a later homonym, I olesbroadly rounded. propose the new name Rhopaloblnste Staminate {lowers (fallen) about 4 dyscrita from the Greek dyskritos (hard mm. high, broadly and shortly ovoid, to determine, doubtful) . The closest more or less oblique, apex broadly relationship would appear to be with R. rounded; sepalsbroadly ovate,rounded, ceramica from which R. d,yscrita seems imbricate, dorsally carinate; petals ro- to dif{er chiefly in spreading pinnae and bust, smooth,ovate, obtuse; stamensand smaller staminate flowers. New and pistillode unknown. complete collections from the type local- Pistillate flowers (based on fruiting ity are much desired to provide a basis perianth) with broadly imbricate and for undersLandingR. dyscrita. rounded sepals3 mm. high; petals ca. I Rhopaloblaste elegans H. E. Moore, cm. high, broadly rounded with short, Principes I0:94. 1966. broadly triangular apex; staminodial ring at length divided into a few denti- Solitary, tall palms tapered from a form staminodes, somewhatenlarged base with a mass o{ Fruit, including perianth, 3.5 cm. long, shortostout adventitousroots, ca. 12 m. oblong in outline, broad, the part slightly high or more, 15 cm. in diam. near base, above the middle 1.8 cm. in diam., with 7.5 cm. in diam. below crown, the bole a thick rostrum and oblique apex above, gray-brown to rather light gray upward slightly acute and narrowed to the calyx with chocolate-brou'nscales on new inter- below; seed ovate-oblongin outline, te- nodes. rete, acute,3 cm. long, I.4 cm. in diam. Leaves rather numerous in a hand- Distribution: New Guinea. Territory some spreading crownl sheathsca. 8.5 of New Guinea: Morobe District. dm. long, light green with a denseindu- Specimensexamined: none. ment of brown scales and appearing In an earlier paper (Principes 10: 98. brown, pinkish inside, tightly appressed 1966), I suggestedthat spreading pinnae and sticky inside; petiole short, 5-7.5 and a staminodial ring were not associ- cm. long, green; rachis straight, covered ated with Rhopaloblaste as I then under- with denseindument of shining, brown, stood the qenus. It has since become fimbriate, peltatescales interspersed with 19701 I{OORE: GENUSRHOPALOBLASTE B5 deciduouswhite tomentum below, densely long, 3 mm. wide, slightly keeledbasally; covered with shining fimbriate scales petals 5-6 mm. long, 3 mm. wide; sta- abovewhen young, ca. 3.4 m. long; pin- mens 6, the flat white filaments ca. 2.5 nae ca. 76 on each side, bent downward mm. long, anthers yellow, 3 mm. long; at an acute angle up to 45o with the pistillode dull yellowish,about as high as rachis, light green, stiff, papery, with filaments, narrowed from an ovate base prominent pale nerves and appearing to a trigonous 3-lobed apex: pistillate sub-plicateabove, green below, the mid- flowers glabrous, surrounded by 2 low nerve elevated and clothed with dark- imbricate bracteoles ca. 1 mm. high, brown or grayish, dull, twisted,basifixed rounded in bud, 4 mm. high; sepals 3 scales above as are also the margins, mm. high, 5 mm. wide; petals 4 mm. below clothed with short and moderate, high, 3 mm. wide; staminodes4, more shining, brown, twisted,basifixed scales, or less united basally; pistil ovoid with the secondarynerves ca. 3 on each side, 3 short erect stigmas; pistillate perianth prominent below and scaly or brown- 6-7 mm. high in fruit. puncticulate, lower pinnae ca. 30 cm. Fruit maturing crimson with a copious long, 1.1 cm. wide, pinnae at mid-leaf glaucescence,green, yellow, to orange 64*68 cm.long,2.7-3.4 cm. wide, upper when immature,globose-obovate, 2.6 cm. pinnae 4I-26 cm. long, 19-B cm. wide, high,2.3 cm. in diam. when fresh, drying the apex acuminatelyand obliquely prae- to 2.5 cm. high, 1.9-2.1 cm. in diam.; morse. seed globose or depressed-globose,I.7 Inflorescencesl-3 below the leaves, cm. high, I.8 cm. in diam., hilum deeply enclosedin bud in subteretegreen bracts impressed, vascular bundles numerous with brown lepidote-tomentoseindument, and reticulatedorsally. the lower bract ca. 36 cm. long, laterally Vernacular names: d,ai'e (Kwara'ae keeled, the keels extending into a flat languageKwai dialect) ; angiriri (Akui rostrum ca. 2 cm. long, the upper bract dialect). enclosedwithin the lower, slightly keeled Specimensexamined: BRITISH SOL- and rostrate, more densely lepidote- OMON ISLANDS PROTECTORATE. tomentose; expandedinflorescence with GuAtltc.tN"{r: north coast. vicinitv of a short peduncle6-8 cm. long, orange- Honiara; gully forest I mile from coast green at base,then green; rachis ca. 30 and 1 mile east of White River head- cm. long; branches 15-16, the first two waters, alt. ca. 500 ft., 27 March 1964, divaricate at about a 90o angle with the H. E. Moore,Ir., G. F. C. Dennis & T. C. rachis and with a main axis ca. 23 cm. Whitmore 9310 (BSIP 4085) (BH, holo- long, again branchedinto simple or fur- type; BSIP, isotype). cate, pendulous, glabrous, non-rugose Rhopaloblatsteele gans probably occurs rachillae to ca. 40 cm. long, tipped with on ChoiseulIsland also. It is distinctive a brief sterile spinose apex, middle in its fruit and once-branchedbasal in- branches once-branched, the apical un- florescencebranches. branched and to ca.25 cm. long; triads borne on lower half of rachillae, paired Rhopaloblaste Ledermanniana Bec- or solitary staminateflowers above,sub- cari, Botanische Jahrbiicher fiir tendedby a prominent rounded bract. Systematik 58: 45. 1923. Staminate flowers glabrous, subsym- Moderate tree 3-8 m. high and thick metric, 6-7 mm. long, greenish,leaving as an arm or a leg. an orange scar on the green axis when LeaI 2.5 m. long, with very many equi- fresh, acutish at apex; sepals 2 mm. distant pinnae, the intermediate ca. 45 PRINCIPES tVol.14

4. Rhopaloblaste elegans at the headwaters of the Matinikau River near Honiara on Guadalcanal. Photo by H. E. Moore. 19701 ,\'IOORE;GENtis RHOPALOtsl,AS'IE 87

5. Mr. G. F. C. Dennis holds the expanded in{lorescence oI Rltopaloblaste elegans in one hand. inflorescence still enclosed by the bracts in tire other. Photo by T. C. Whitmore. cm. Iong, 15 18 mm. wide, ensiform, In{lorescences ample, 34 times straight, the apex acuminate and not or branched, lightly reddish furfuraceousl scarcell, falcate, with minute scales on rachillae 25 30 cm. long, 2 2.5 mm. midnerve belorv, secondary nerves 3-5 thick. on each side below, prominent, margins Staminate flowers yellow, regular, and secondary nerves puncticulate. ovate in outline; stamens 6, filaments BB PRINCIPES tVol.t4 conspicuouslyinflexed at apex; pistillode cm. long, densely lepidote with inter- conspicuous,trigonous, apically 3-lobed. locking, peltate,brown-centered, pale and Fruit ovate, symmetric, acute, l0 mm. often lacerate-fimbriate-margined scales long, B mm. in diam.; seedovate, acute, or becomingmerely brown-puncticulate; endosperm ruminate; fruiting perianth petiole elongate, 60-70 cm. long, with sheathing the fruit one-third, cupular, 4 lacerate, castaneous,membranous scales; mm. long, 5 mm. wide, subtendedat base rachis ca. 1.I m. long, rounded and cas- by reniform conspicuousbracts approxi- taneous-lepidotebelow, angledabove and mate to the calyx. densely lepidote with basifixed, twisted, Distribution: New Guinea. Territory membranous scales or brolvn-puncticu' of New Guinea, along the April River. late; pinnae 40-50 on each side, spread- Specimensexamined:none. ing, slender,to ca. 30 cm. long, 13-14 Beccari cited two Ledermann collec- mm. wide, the midnerve and usually a tions from the April River (Aprilflusse) prominent secondary nerve on each side -9718. 8648-and, in text mentioned on lower surfacewith basifixed or medi- another, Led,ermann 870,8, Irom which fixed, twisted, brown, membranousscalbs measurementsof pinnae have been taken. at least near tlre base,all nerves minutely The above is a free translation of the brown-puncticulate,median pinnae dark original description published in Latin green,to 3I cm. long, 12 mm. wide. and German. Neither authentic material Inflorescencesdecurved, to ca. 20 cm. nor photographsof specimenshave been long, densely covered with simple to seen,hence it is not possible to determine branched or even stellate, brown to pale whether the species is truly a Rhopalo' hairs or glabrescent,simply branched, or blaste or perhaps a species of Hetero' the lower branches forked; lower bract spa,the.The type presumably having been to 14 cm. long, denselybrown-lepidote- destroyedat Berlin, it is to be hoped that tomentose, upper bract densely whitish- ultimately new collections will be made lepidote-tomentose;peduncle 1.5-3 cm. from the type region to resolve the prob' long; rachis 2-6 cm. long; rachillae lem of identity. green-brown, transversely rugose, to 34 cm. long, subtendedby low, acutebracts; Rhopaloblaste singaporensis (Bec- triads subtendedby roundedbracts to ca. cari) J. D. Hooker in Bentham & 1.5 mm. high; bracteoles surrounding J. D. Hooker, GeneraPlantarum 3: pistillate flowers subequal,to about 1.4 892. 1883. mm. high. Staminateflowers yellowish, 3-4(-5) Ptychosperma singaporerrce Beccari, mm. long at anthesis; sepals1.5-2 mm. Malesia L: 6L IB77 (osingaporen- sz,r'). high, margins ciliolate; petalsca. 3 mm. high, adnate to a receptacleca. I mm. Pty ch,oraphi,ssingaporensis ( Beccari) high; stamens 6-9 but frequently B; Beccari, Annales du Jardin Botani- pistillode conic, about as high as the sta- que de Buitenzorg2: 90. lBB5. mensin bud. Ptychoraphis longillora H. N. Ridley, Pistillate flowers at anthesis about 5 Journal of the Royal Asiatic Society, mm. high; sepals2 mm. high; petals4.5 Straits Branch 41: 38. 1904. mm. high; staminodes3-6, membranous, Stems cespitose, to about 4 m. high, broad, obtuse,discrete or variously par- gray-brown, prominently ringed. tially connate in an irregularly lobed Leavesto 1.7 m. long; sheathaboat26 ring. 19701 MOORE:GENUS RHOPALOBLASTE

6. Rhopaloblaste singaporensis{orms clumps in the Singapore Botanic Garden. Photo by G. Addison.

Fruit orange-yellowto red, 13-14 mm. Specimensexamined: SINGAPORE: high including prominent beak 2 mm. a Woodlands, Marzo 1866, O. Beccari high, 9-10 mm. in diam.; endocarp Hb. No. 11180(FI, holotype;photos BH, about 11 mm. long, 8 mm. in diam., neg. 3870_72) ; nella parte selvatica del prominently impressedover the hilum, orto botanico, Gennaio 1878, O, Beccari operculum round, hilum impressed in Hb. No. 11181(.FI, photo BH, neg.3873- the seedthe length of one side. 77; wooded slopes of nature reserve' Seedlingwith pinnateeophyll. Bukit Timah, 14 Dec. 1963,H. E. Moore, Distribution: Malay l?eninsula and Ir. & T. D. Pen"nington9047 (BH) ; Singapore. Krangi, 5 Aug. IBB9, H. N. Ridley 2136 90 PRINCIPES [Vol. 14 "Ridley's (FI) ; Chan Chu Rang, L892,H. N. Rid- observations and descrip- ley 2124 (FI) ; Loos, l.BgL,H. N. Ridlcy tions on Malayan plants are sometimes 3137 (Fl\ MALAYSIA. M.lr-A.vA: Jo- uncritical. The diagnostic difference he hore; north of Labis, 5 miles, M. R. Hen- stated from Ptychoraphis singaporcn- d,erson38203 (BH) . sjs-longer male petals, which he later 'slender RhopoJobla'stesingd,porenszs is here amended to branched in- taken to include Ptychoraphis Xongillora florescences with distant flowers and which seemsnot to differ in significant lanceolate petals' (Flora ol the Malay detail. The little-branchedinflorescence Penins-y,la5: 19, I925)-are not fully with transversely rugose, mostly hairy substantiated by the full descriptions. I rachillae,the cespitosehabit, and orange' very much do,ubt i{ there are two species yellow fruit set the speciesapart. of this genus in Malay, and it seemsthat Dr. T. C. Whitmore has independently Ptychoraphis longiflora, a very weak concluded that Ptychorapltis longillora speciesif one at all, is now probably could not stand as a distinct species. extinct." With his permission, I include here two The description above was drawn paragraphswhich were to have appeared largely {rom my own notes taken {rom a in a forthcoming article of his own. plant at Bukit Timah where the leaf "Ptychoraphis longillora was based sheathswere tubular and the apparently on Ridley ll2l collected in Johore on mature fruit was orange'yellow. Dr. the top of Gunong Banang at Batu Pa- Whitmore, however, has noted in other 'oln hat. I could not find the collection at localities that R. singaporenszs,the Kew or Singapore. There is now a road crownshaft is obscuredby old persistent to the top of G. Banang and a telecom- leaf sheathswhich slowly rot away rather munications tower on the top. The orig- than abscissingneatly at the bottom and inal forest has been destroyedall along falling in one piece. The inflorescences the summit ridge and the slopes have develop in the axils of these moribund been culled for timber. Now there is leaves, sometimes burst through their secondary forest and degraded, climber- basis and so{netimes persist after their tangled, primary forest patches. I very sloughing. The fruit eventually ripen much doubt if Ptychoraphis, a delicate red. There are several records from the palm of primary forest undergrowth Dindings in lower Perak including my shade, has survived all this; I myself own FRl 099L I have also extendeclits could not find it nor any other of the range into E. Pahang, Aur Forest Re- palms of primary forest undergrowth. serve (1R1 3685\ ."

ExcrulBl ANDUNCERTATN SpEcrES AND SoME Tn.s.Nsrsnsro HErERospArHE

The following do not belong to or are Ptychoraphis uncertainly placed in Ptych.orapftisand P. cagayensis : Heterospathe cagay- Rlrcpal,oblaste.They are referred chiefly ensis to Heterospathewith complete synonymy P. Elmeri: Heterospathe Elmeri provided under the listing {or that genus P. intermed,i.a= Heterospathe sp. below. Two epithets have not been trans- = ferred owing to the need for a detailed P. microcarp@ Heteroepathe sp. study of Heterospathe in the Philippine P. ph.ilippinensis = Ifeterospathe phil- Islands. ippinensis 19701 MOORE: GENUSRHOPALOBLASTE 91

P. SiebertianaHort. Sander, Gardeners' nervatureand acuteapices of the pinnae, Chronicle,series 3, 43:257 and SuP- together with the morphology of the plementary lllustration, APt. 25, staminate flowers in combination are 1908 : ? congruent neither with Ptychosperma nor with Rhopaloblaste. They are con- The description accompanying this gruent with Heterospatheas arc fruits a juvenile plant and is so gen- name is of from a separate packet in the Beccari eventhe genuscan be prop- eral that not Herbarium (.No. 11163) which were The name should be erly ascertained. associated with the type by Beccari future consideraiion since reiectedfrom though not described. These fruits are to be no further description thlre appears 13 mm. long, B mm. in diameter,with a requisite for generic assign- with detail perianth 3 mm. high. They are within ment. the range of expected size for mature (Burret) Rhopaloblaste fruit of Heterospathe pilosa Burret {rom the Cyclops Mountains of R. arlakiana = Heterospathe arfaki- West Irian. Should thesetaxa ultimately ana prove identical, the narne Heterospathe R. Elmeri: Heterospathe Elmeri arlakiana will have priority. R. intermedia = Heterospathe sp. Heterospathe cagayensis Beccari, R. Macgregorii : Heterospathe Mac- Philippine Journal of Science,Bot- gregorii any4: 6Il. 1909. R. micrantha = Heterospathe micran- Ptychoraph.is cagayensis (Beccari) tha but see also Rlnpaloblaste d,ys- Beccari, Philippine Journal o{ Sci- crita ence14: 328.19L9. nticrocarpa: Heterospathe sp. R, Heterospathe Elmeri Beccari in El- R. princepsHort. Bull, Gardeners'Chron- mer, Leaflets of Philippine Botany icle, series2, 13: 759. 1880 : ? 2:646. 1909. The description, if such it can be Ptychoraphis Elrneri (Beccari) Bec- called, cannot be applied even at the ge- cari, in Philippine Journal of neric level. The name should be rejected Science14:328, I9I9. from future consideration. Rhopalobla,steElmeri (Beccari) Bec- cari in Martelli, Atti della Societi Heterospathe Toscana di Scienze Naturali resi- Ileterospathe arfakiana (Beccari) H. dente in Pisa, Memorie 44: l3B E. Moore.tr. nou. freprint 271. L934. Ptychosperma arlakianutn Beccari, Malesial: 57, 101. 1877 ('art'aki- Heterospathe Macgregorii (Beccari) ana''5. H. E. Moore,tr. noD. Rhopaloblaste ? arlakiana (Beccari) RhopaloblasteMacgregorii Beccari in Beccariex Martelli, Nuovo Giornale Martelli, Atti della Societi Toscana Botanico Italiano, series2, 42: 76. di Scienze Naturali residente in 8L 1935. Pisa,Memorie44: I34 [reprint 23]. 1934. Mature fruit of certain origin is not part of thetype (BeccariHb. No. 11187) The type of this species,collected by but the nature of the inflorescence,the Sir W. MacGregor on the Fly River, PRINCIPES lVol. 14

Papua, is fragmentary with some sections provisional key to speciesruns to the of leaf, two branchesof an inflorescence vicinity of Heterospa,the Clemensiae in very young bud, and loose fruits. It because of the ellipsoid fruit. There do conforms well with a more recent collec- appear to be adequate specific differ- tion from limestone banks of the Kikori encesbetween the two taxa however. River, Gulf Division, Papua (K.I. White N. G. F. 10714, BH), which has a long- Heterospathe philippinensis (Bec- pedunculate inflorescence in its entirety cari) Beccari,Philippine Journal of and three-nervedpinnae characteristic of Science,Botany 4:610. 1909. Heterospathe. The fruit has an essen- Ptychoraph,is philippinensis Beccari, tially apical stigmatic residueas in ftho- Annales du Jardin Botanique de paloblaste but unlike that genusthere are Buitenzorg2: 90. 1885; Malesia 3: diagonal fiber-sclereids in the outer 109. 1886. mesocarp and anastamosing red tannin bodies between the longitudinal fibers Heterospathe sp. and the endocarp. Transferred to Heter- ospathe,the speciesapproaches H. pilosa Pt;ychoraphis intermedin Beccari, (Burret) Burret from West Irian but Philippine Journal of Science 14: differs in having much larger though 328. Mar. l9I9; Leaflets of Philip- ellipsoid fruit. pine Botany B: 3011.Aug. 1919. Rho'paloblaste intermed,ia (Beccari) Heterospathe micrantha (Beccari) Beccariex Martelli, Nuovo Giornale H. E. Moore, tr. nou. BotanicoItaliano, series2, 42:75, Ptychosperrna micranthum Beccari, 81.1935. Malesia I: 52. lB77 ('micrantha'). The relationship of this and the next Rlr,opalobl,astemicrantha (Beccari) to other species oI Heterospathe in ilre Bentham & J. D. Hooker ex B. D. Philippines is not clear hencethe epithets Jackson, Index Kewensis 2: 713. are not transferred. 1895; Nuovo Giornale Botanico Italiano,series 2, 42: 78,81. 1935. Heterospathe sp. Ptychoraphis microcarpa^ Beccari, Examination of the type material of Philippine of Science 14: this speciesmakes clear that it belongs in Journal Heterospathe rather than Ptychosperma 327.r9r9. or Rhopaloblaste. It falls among those Rh,opaloblastettl.icrocdrpa (Beccari) speciespreviously referred to Ptycha,nd,ra Beccariex Martelli. Nuovo Giornale but recently incorporated in Heterospathe Botanico ltaliano, series2, 42: 75, (Principes 13: 99-105, 1969) and in a BI. 1935.