124 PRINCIPES lVoL. 36

Principes,36(3), I992, pp. 124-127

Voanioala,the Forest

Herbarium,Royat BotaniciH.TilT,t#, nd,surrey, Twe sAE, u.K.

By now, many members of the Society much like that of the coconut but the male may be aware of the existencein Mada- flowers have 12 stamens.Pollen is unlike "the gascar of a palm nicknamed Forest that of other members of the Butiinae Coconut." It was Mardy Darian who first (Harley 1989). The fruit is 7-8 x 4-5 brought the possibleexistence of this palm cm and appearssuperficially like a fruit of to my notice, but it was not until October an . Inside the fruit there is a very I986 that good scientificspecimens were thick endocarp that has irregular bumps collected in the northeast of the island. on the insidethat penetratethe seed,rather The Forest Coconut has several unusual like a walnut. Seedlings gro\rn at Kew features that make it necessaryto include causedgreat excitementas, from their root- it in a new . The name Voanioala tips it was discovered by *y colleague gerardii has been validly publishedfor the Margaret Johnson,that the Forest Coco- ForestCoconut (Dransfield I9B9a). At first nut has a very high number of chromo- sight the genericname may be horrifyingly somes-at least 596 (Johnson1989). In unpronounceable; however, it now triPs fact, the mass of chromosomeswithin the euphoniouslyoff my tongue! Pronounce it cell nucleus looks, under the microscope, "voh-ann-ee-oh-al-a,'obut run the first two almost like scrambledegg rather than the 'ovuan," syllablestogether to give, almost, usual small ribbons. What the significance 'oee" 'ooh" and the and the should be run is of this astonishingly high chromosome together too. This will then approximate number, the highest recorded for any to the Malagasypronounciation! It is based monocotyledonous , no one really on the Malagasy name for the palm, and knows. "Forest means'literally, Coconut.'oIn using A popular account of the collecting of a Malagasy name as a generic name, I the first scientific specimensof Voanioala "Plant followeda long establishedprecedent-the (Dransfield19896)is givenin hunt- narnes Votuitra arrd,Raphia are based on ing for Kew,'o a book that describesthe Malagasy plant names while Antongilia, plant hunting activitiesof the Royal Botanic Masoala and Marojejy@ are based on Gardens Kew. Since that first expedition Malagasy place names. The palm has I have returned with David Cooke of the proved to be of exceptional interest on Palm House at Kew to the original locality many {ronts. It is indeed a relative of the to look at the other palms that we had coconut and represents the first member been unable to study for lack of time on of the subtribeto which the coconutbelongs the first trip. It seem$ certain that this (Butiinae) to tre recorded for Madagascar, palm is of very limited distribution, occur- though not the first cocoidpalm (the others ring as scattered trees in the rain forest. are the African Oil Palm, Elaeis guineen- Villagers elsewhere in Madagascar have sls, and the endemic Beccariophoenix obliged by reporting the presenceof other oovoanioala" madagd;cariensis). It is a big tree palm trees of but thesehave always with rnassive bases and big pinnate turned out to be individuals of tall tree . The inflorescence appears very palms such as Neodypsis lastelliana. ln t992ll DRANSFIELD:VOANIOALA 125 l9B9 in the National Park of Marojejy, I rachillae, each subtendedby a small triangular rachis was told of the existence of five different bractl rachillae numerous. most with a basal bare oovoanioala"; portion, the rachillae bearing 0-7 triads near the typesof sadly not one of these baseand paired or solitary staminate flowers distally, was the true Voanioala, further empha- the flower groups spirally arranged, or becoming sizingthe restricted distribution of the true somewhat distichous by close"packing. Staminate Forest Coconut. flowers asymmetrical, broadly or narrowly triangular News travels fast. Within a year of the in outline, sepals3, distinct, slightly to strongly imbri- cateat the base,triangular. acute to acuminate.mem- collecting of the first scientific specimen, branous,glabrous; petals 3, distinct, unequal, valvate, the villagers in one part of northeast Mad- glabrous,thinly coriaceousexcept at the thick angular agascar where seyeral palm enthusiasts tips, broadly and irregularly triangular-ovate, with have visited, were well aware of the Forest acute or acuminate tips, abaxially smooth, adaxially marked with the impressionsof the stamensand papil- Coconut and were busy collecting seed to losenear the thick tips; stamens12(-13), filaments sell to anyone who wanted to buy. Several subulate, very short to moderate in length, anthers private collectors may now already have basifixed, basally sagittate, apiculate at ihe tips, seedlingsof this remarkable palm. latrorse; pollen circular or elliptic in polar view, mono- A detailed scientific description of the sulcate, intectate with microbaculae, granules and pilae (seeHarley 1989). Pistillate flowers only known gerrusVoanioala is provided here, set out as buds, much larger than the staminate, irregularly in the format of the descriptionsin Genera triangular; sepals3, distinct, unequal, strongly imbri Palmarurn (Uhl and Dransfield i9B7) so cate, broadly ovate, with triangular, keeled tips, oori that you may make a copy of the descrip- aceous,glabrous, the margins minutely toothed; pet- longer the sepals,basally irregularly (Fig. include it as als 3, distinct, than tion and plate 1) and imbricate, conspicuouslyvalvate at the triangular tips, genus 177A, after Parajubaea. abaxially with scaly indumentum towards the apex, adaxially strongly papillose towards the tip; stami- nodial ring high with 9 irregular, triangular teeth, 177A. VoANtoALA J. Dransfreld,Kew 0.1-0.5 mm; gynoeciumsyncarpous, tricarpellate, and scaly, Type: V. gerardii triovulate, stigmas 3, angled, papillose Bulletin 44(2):192. ovules with axile placentation. Immature fnrit green J. Dransfield. covered with dense chestnut brown scaly indumen- tum. Mature fruit one-seeded,somewhat irregularly monoecious Robust solitary, unarmed, pleonanthic ellipsoid, tipped with a short beak and stigmatic erect, with a large root boss, tree palm. Stem basally remains; epicarp red'brown, densely covered with very "stepped" distally the stem bare, conspicuously brown scaly indumentum; mesocarp with an outer and ringed with oblique leaf scars. Leaves pinnate, fibrous zone just below the epicarp, and an inner reduplicate, cleanly abscissing;sheath tubular at first, fleshy zone; endocarp ellipsoid, apically pointed, basally fibrous, apparently soon disintegratingto leave a mas- truncate, very heavily thickened, pale trrown when sive elongaterectangular leaf base,forming an appar- fresh, becominggrey with age, very deeply and irreg- ent petiole with sparsely fibrous margins, abaxially ularly longitudinally grooved, with 3 very deep basal densely covered with caducous brown indumentum; impressionseach with a central germination pore, in leaf basesuddenly contracting into rachis, true petiole section the body of the endocarp traversed by lon- absent, the rachis more or less rectangular in cross gitudinal irregular vertical canals and fibers, inner section in the mid leaf region, abaxially densely cov- surface of the endocarp with numerous irregular ered with'caducous brown indumentum as the leaf rounded excrescencesintruding into the cavity. Seed base; leaflets numerous, rather stiff, scarcely pen- irregularly ellipsoid, filling the endocarp cavity, lat- dulous,very coriaceous,concolorous, mevenly bilobed erally attached with a narrow irregular hilum, endo- interfoliar, appar- at the tips. Inflorescencessolitary, sperm homogeneousbut irregularly intruded by the to one order, erect in ently protandrous, branching endocarp protruberances, very hard, white, with a bud, later horizontal; prophyll tubular, 2-keeled, narrow, irregular central lacuna; embryo basal, top- fibrous, remaining hidden among the leaf bases; shaped, positioned opposite an endocarp pore. Ger- green strictly tubular in peduncular bract bright and mination remote-tubular; eophyll entire, lanceolate. bud, later splitting longitudinally, flattening and 2n:596* (V. gerardii, Johnsonl9B9). becoming somewhat cowl-like, abaxially deeply and closely longitudinally grooved, bearing scattered brown Anatomy.-Not studied. scales on the ridges between the grooves, adaxially -A smooth, glabrous, pale cream-colored; peduncle cir- Distribution. single speciesin cular in crosssection; rachis bearing spirally arranged northeasternMadagascar. r26 PRINCIPES [Vor. 36

r- r! ft&$ I i I t

L N IIt\

\_/ \ ,D

t. Voanioafa, ?, portion of rachilla with staminate flowers X 2/z; b, rachil.la with triads x lti C, pair of staminate flowers removed to show bracteoles x $t4' d, pair of staminate flowers X llZ; 9, staminate flower x 2; f, staminate sepals x SVz; 9, staminate petals x 2; h, pistillode x 5r/z; I staminate flower in vertical section x 2; i, stamens x 3Vs; k, triads of flowers removed to show bracteoles x 2/ti l, pistillate flower x 2; m, pistillate sepals x 2; l'1,pistillate petals x 2; O, pistillate flower in vertical section x 2; P, gynoecium with staminodial ring attached x 2; Q, cross-sectionovarian part of gynoecium x 6; lr staminodialring x {; 5, 161 x /z; l, fnit in vertical section X Ts; l!, base of endocarp showing pores x 7s| Y, endocarp in two views X Te. Voanioala gerardii, Dransfield 6389. r9921 DRANSFIELD:VOANIOALA r27

Ecology. -V. gerardii grows in Lrrnnerunu Cttoo valley bottomsand on gentleslopes swampy DRANSFIELD,J. 1989a. Voanioala (Arecoideae: at an elevation of about 400 m in primary Cocoeae:Butiinae), a new palm genusfrom Mad- rain forest. agascar. Kew Bulletin 44(2): 191-198. Common Names and Uses.-Yoan- 19896. Searchingfor a forest coconut in "forest (ed.). Plant hunt- ioala (Malagasy) (literally coco- Madagascar.1n: F. N. Hepper ing for Kew. Royal Botanic Gardens, Kew, Her nut"). The apical bud is said to be edible. Majesty'sStaionery Office, London, pp. 5t-60. In view of the great rarity of the palm, it HARLEv,M. H. 1989. Pollen morphology of Voan- is perhapsnot surprisingthat no other uses ioala gerardii (Palmae: Arecoideae: Cocoeae: have been recorded. B utiinae) lr om Madagascar.Kew Bulletin 44(2): I Notes.-Voanioala possessesseveral 99-205. JoHNSoN,M. A. T. I989. An unusuallyhigh chro- unusual features, such as the multistami- mosome number in Voanioala gerardii (PaI- nate flower, the pollen and the endocarp mae: Arecoideae: Cocoeae: Butiinae) ftom with deep longitudinal grooves and inter- Madagascar. Kew Bulletin 44(2): 207 -210. nally with irregular swellings.The unique Usr, N. W. exo J. DRANSFIELD.1987. Genera Palmarum: a classificationof oalms basedon the high chromosomenumber is remarkable, work of H. E. Moore.Jr. L. H. BaileyHortorium the only other high number recorded for and the International Palm Society. Allen Press, the palms being in Jubaeopsis caffra, also Lawrence, Kansas. belonging to the Butiinae. Taxonomic Account.-Dransfield (l989a).

Acknowledgment I should like to thank Marion Sheehan for preparing the analytical drawings.

BOOKSTORE ESTRY (H. B. Pedersen and H. Balslev, 1990, I05 pp.) .....,.."...... 15.00 ExorrcA (4) (A. Graf, I982, pictorialen- A Guroe ro rHE MoNocorYLEDoNS cyclopedia,2 vols.,including 250 plant oF PAPUANew Gurruea,Panr 3, families,16,600 illust., 405 in color, Plr-ule (R. J. Johnsand A. J. M. Hay, 2590 pp.) -..--.---.,-...--.,--.---.-..-I87.00 Eds.,1984. l2a pp.) 8.00 FloRe NeorRoPrcA lNTRoDUcrtoN BRAztLtANPALMS, Notes on their uses AND THE IRIARTEINAE (A. Henderson, and Vernacular Names (C. Pinheiro and 1990, 100 pp.)--..-..--.--...-..--.,-- 23.00 M. Balick. 1987. 63 pp.) ...... -...... 9.25 FLoRA oF Tnoprcl|- EAsr AFRtcA, PALMS (D. Hodel, Cxaureoonea 1992, Peuare (J. Dransfield, 1986, 52 350 pp., I27 pp. of superb color) .,-..--.,-..59.95 ^. \ ------23.00 CocoNUT PALM FRoND WEAvrNe PP.t Fldne oes MascnRercnes (La Re- (Wm. H. Goodloe, 1972, 132 pp.) ..-...... 8.95 union, Maurice Rodrigues, 1984, 3t CocoNur RESEARcH lNsrrrurE, nn.) 5.00 MANADO (P. A. Dards, H. Sudasrip, Fi6nroa Patus, Handbook of (8. and S. M. Darwis, 1985, 165 pp., 79 McGeachy, 1955, 62 pp.) -..-..--..--..--..--..--.--..2.95 pp. color) 35.00 FLoRTDATREES AND PALMs (S. A. Rose, Cutttvlreo PALMS oF VENEZUELA A. A. Will, Jr., T. B. Mack, 1984, 30 (A. Braun, I97O,94 pp. and 95 photo- palm ,120 pp.) -.. 6.00 graPns., t -95 GeNem PALMARUM(N. W. Uhl and J. DESERT Pat-u Olsts (J. W. Cornett, Dransfield,I987, 6 I 0 pp.) --.--..--...--..--..--.-..,79.00 1989, 47 pp., 41 pp. color) ..-..-...-..-..-...-.....8.95 Hrnvesr or rrtE PALM (J. J. F ox, I97 7, DrsEAsEs AND DrsoRDERs oF ORNA- 244 pp.) 24.00 MENTAL PALMS (A. R. Chase and T. lNoex ro PRrNcrpEs(Vols. 1-20, 1956- K. Broschat, 1991, 56 pp., color on 1976, H. E. Moore, Jr., 68 pp.) --..--.--.---.-4.00 each page) 29.OO EcuaooRrlnPelus ron Acnoron- (Continued on p. 132)