19861 RAUWERDINK:

Principes,30(4), 1986, pp. 165-180

An Essay on Metroxylon, the Palm

JeNB. ReuwnRomx Department of , Agricultural Uniaersity, Wageningen, the Netherlands P.O. Box 8010, 6700 ED Wageningen

Metroxylon is a of arborescent under cultivation. The aim of my survey palms of Papuasia and several island and the present paper has been to report groups of Micronesia and Melanesia. There on the variability of M. sagu in PNG, in are five occurring in five separate the context of the diversity found in the areas. The most widespread taxon, M. genus as a whole. This paper may con- scLgu, covers , , Min- tribute towards an eventual monograph of danao, and . The other four Metroxylon. taxa are endemic to the aforementioned island groups. Historyof the Genus The palms accumulate in the pith of their trunks and are a traditional source The first and most competentpublica- of carbohydrate. The best known r-epre- tion on sagopalms is by Rumphius(1741). sentative of the genus in this respect is In the Herbarium Amboinensehe gives M. sagu, known as the sago palm. This a meticulousdescription of the sagopalm species occupies the largest area. esti- as it occurs in Ambon. and he Dresents mated to cover 4 million ha in natural the taxonomic views of the inhabiiants on stands and about .2 million ha under cul- this palm. Four Ambonesespecies are tivation. With the exception of M. salo- described under the seneric name of monense.the other tp".i"t of Melroxylon Sagris.This namewas adopted by Caert- are not exploited for their starch content. ner (1788), but unfortunately alsoincluded At present, interest on Metroxylon as a Rctphia P. de Beauvois.Giseke (1792) starch-producing crop is focused on M. does not include Raphia and gives the sogu which grows in swampy areas usu- earliest binomial applying solely to the ally unsuitable for plantations of other descriptionof Rumphius'sS. genuina. The crops. So far, two international symposia name Sagus is, however, antedated by have been held to evaluate the exoloita- Metroxylon Rottboell(1783), basedpartly rion of sago starch (Sago I 977. Sago on Rumphius'sdescription, but in larger l9B0). In addition, an FAO expert con- part on specimensprovided for Rottboell sultation took place in Jakarta in 1984. by Dr. Koenig. The next author to adopt Studies were carried out in Indonesia Metroxylon was Von Martius (1845). (Team Proyek I9B2) and New Nearly eighty years after Rottboell, Guinea (DME 1980, 1982). Wendland (1862) proposedthe generic The present author has made a taxo- name Coelococcusin which he described nomic survey for the Papua New Guinean a sagopalm from the Islands.Coelo- government in the Sepik River flood plain coccushas henceforwardbeen adopted by (Fig. l). An estimated one million ha of various other authors (Dingler lBB7, natural sago palm stands occur in PNG, Warburg 1896, Heim 1904) to classify and an additional 20 thousand ha are several species of sago palm from r66 PRINCIPES [Vor. 30

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Micronesian and Melanesian islands; DiagnosticCharacters Metroxylon is used exclusively for those sago palms occurring throughout New A brief treatment of the diagnostic Guinea, Indonesia, and Malaysia. Beccari characters wrthin Metroxylon is appto- (1918) included Coelococcusin Metrox- priate. Especiallywhere the ylon as a section,an opinion that has since is concerned,the application of correct teen followed. Somecommotion may have terminology to refer to particular units is arisen in the early sixties when Moore vital. In this paragraph,diagnostic char- l96D disclosedthat Rottboell'sMetrox- acters are treated in an order reflecting ylon is actually antedatedby SagrzsSteck the comparativeimportance of each char- (1757). However, Steck's publication is acter within the taxonomy of the genus: not effectively published, and Metroxylon the most significant character is treated tt conserved (t968) against this first. *".,T3. Inflorescence: The inflorescenceof r eB6l RAUWERDINK:METROXYLON t67

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2. of . 3. Spines on sheathsand petioles of a mature specimen. 4. Fruits of M. amicarum. 5. M. sagu, somaticchromosomes in equatorialplane (photo by J. C. Arends).

Metroxylon is extensively discussed in gle basal ovule in each locule, usually only Tomlinson (I971). For reasons of brevity a single matures in each . This of this paper, reference to this publication seed contains a considerable amount of is made for interested readers. As far as hard, bony, homogenous endosperm which diagnostic characters are concerned, the has a very characteristic horseshoe shape shape of specific units of the inflorescence in vertical section, with a large chalazal has oroved useful. Most characteristic is cavity. The embryo is situated in a smaller, *hether branches of a certain order are second peripheral cavity. Differences pendulous or not. This single character between the taxa are found in the number enables unambiguous distinction of three of longitudinal series of scales on the fruit. out of the five species recognized in The divisionol lhe genusinto two sections Metroxylon. is based upon the number of scales. At the Fruits (Figs. 2,4): The fruits of specific level, the shape and the size of Metroxylon are covered with longitudinal the mature fruit is also diagnostic. series of imbricate rhomboidal scales (Fig. Spines (Fig. 3): Spinescence has been 4). The apex of the fruit is always emphasized in Metroxylon as an impor- mucronate, the mucro representing the tant diagnostic character throughout. remains of the style. Although the pistil Authors, such as Rumphius (1741) and of Metroxyktn is three-locular with a sin- Beccari (1918), as well as sago-subsisting r68 PRINCIPES lVoL. 30 people, all greatly value spinescencein length of spines may be governed by a iheir respectivetaxonomies. Among other comparatively simple genetic mechanism. thines this led to the division of the section Thus, different formae in the section Meioxylon into 17 taxa bY Beccari. Metroxylon (e.g. spiny vs. smooth) are However, in the section Coelococcushe considered to be only slightly different distinguished 7 taxa only. Probably this genotypesof a single species,M. sagu, was becauseBeccari lacked the extensive instead of two or more different speciesor collectionsand experiencein the field with subspecies.This assumptionwill be dealt these taxa that he had in Metroxylon' with in the Discussion. Like Beccari, the present author is famil- Srimatic Chromosome Numbers: iar with the section Metroxylon, but has Somatic chromosomenumbers of M' sagu scant knowledge of Coelococcus aparl were determined from tiPS of and descrip- from herbarium specimens I) Seedlingsgrown at the Departmentof the data, all taxa in tions. According to Plant Taxonomy at Wageningen. The are spinescent, the section Coelococcus seedswere collected in the Sepik area but details such as length and abundance (PNG)by Mr. Van Kraalingenand all Such charac- of spinesare not available. specimenswere spiny. One seedlinghas betweentaxa ters enableme to distinguish beenpreserved at WAG (Rauwerdink in the section Metroxylon, and they may lsB). the section Coe- be of the same value in 2) grown at the Department o{ in transverse lococcus.Spines are found TropicalCrop Scienceat Wageningen, the abaxialside crescent-shapedseries on from obtained from the Singa- surface of the of a leaf, over the entire pore Botanical Gardens. Spiny as well of about 3 leaf sheath, and along a strip as unarmed plants were available. cm wide on the and rachis. Spines are pectinate, flattened, of a brownish Permanentsquash preparations were made color, and confluent by their bases,i.e. accordins to the methods describedin De arrangedas in a comb(Figs. 12,13). Their Wilde and Arends (1979). The slides are length rangesfrom a few cm tp Io 22 crn. retained in the slide collection of the Furthermore, they are unequal' the mid- Department of Plant Taxonomy. No living dle one of a series usually being the lon- maierial from the other taxa in Metrox- gest. Reduction in length of spinesoccurs' ylon was available for an investigation. in which casethey are referred to as spi- The Singapore plants had already been nules. Spinulesare short, up to l0 mm investigatedby Verhaar (1977) ar 2n: long, and usually of a triangular form. The 26.The presentauthor also obtainedthe ultimate reductions of spinesare knoblike somaticchromosome number of 2n: 26 structures which occur exclusively on the in M. sagu (Fig. 5), both in the PNG baseof leaf sheaths.The variability in spine specimensand in the Singapore material. morphology in the section Metroxylon is The basicchromosome number of M. sagu : threefold. The first is related to the age can therefore be put at x 13, which of the palm. Crown of a mature actualJymarks M. sogu as an exception specimen usually have spines that are in Calamoidpalms. The basic chromosome : much shorter than those on leaves from numberin thesepalms is x 14, accord- offshootsof the samepalm' Secondly,the ins- to Moore and Uhl (1982)' exact location of the spines is important. Analysis of Pollen: Recentanalyses on Especially in a mature palm, the spines the pollenmorphology of Metroxylon were on the leaf sheath are longer than the undertaken by Sowumni (1972) and spines on the petiole and rachis. In the Thanikaimoni(1970). This paragraph third place, I am of the opinion that the presents a few observationsby the latter r9861 RAUWERDINK:METROXYLON r69

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DT a, saga, overview, bar : 1.0 6-7. 6. Scanningelectron microscope(sEM) photo of a pollen grain; lltrgxyl.on 0. oI M' micrometers; b, detail of exine ."ol'ptn.", Uai: O.t *i".o*"t"rr. 7. SEM photo of a pollen grain N. ,uarburgii; a and b, as above tot M. sagu. il author on the sculpture of the exine in wide. On the basis of the sculpture of the exine of the grains, the genus can be le Metroxylon. divided into two groups. One group has a )e Pollen grains were obtained from her- reticulate ornamentationon the equatorial re barium specimens on loan from various sides.The lumina of this reticulum dimin- le institutions. The specimensanalyzed are colpi, resulting in rg listed below. Voucher slidesof pollen have ish in size towards the and the respec- a perforatetectum (Fig. 6a,b). This group !t- been filed at Wageningen consists of M.. salomonense and M. n. tive institutions. uitiense.The other group displaysa per- t: M. sagu: Balt'huizen u/d Brink 74BB almost the total sur- (B). forate tectum over face of the grain (Fig. 7a,b). This group he (K). M. salomonense:Moore 9317 consistso{ M. warburgii, M. amicarum, z6 (K). M. uitiense:Smith 9339 and M. sagu. The exine-sculptural types .C s.n. (B) and M. amicarum: leg. Kramer I found were also recognized by Thani al. (K\. Furtado 25868 kaimoni and Sowumni; neither author, (P). {u M. warburgii: Moore 9319 however, had the opportunity to investi- ch have been analyzedwith the gate all taxa in the genus. Thanikaimoni QN Pollen grains (SEM) and used the terms coarsely reticulate and Ire scanningelectron microscope (LM). finely reticulate for the exine sculpture of rd- with the light microscope Samples were preparedaccording to the acetolysis M. salomonenseand M. sagu resPec- Sowumni uses the terms reticulate on procedure that is commonly used at the tively. and vermiculite. Praglowski and Punt, in fre Department. The pollen grains of all taxa their 1973 paper, proposethe definitions nd in Metroxyl,onare dicolpate,i.e. possess- I have adopted ph ing two germinationapertures. They are for exine sculpture which here. At present, it is difficult to relate the ler oval, about 30 pm long and about 15 Pm 30 t70 PRINCIPES lVoL. sheath two types of pollen grains in Metroxylon spreadingand eventuallyhorizontal; semicir- to the taxonomy of the genus' The divi- broad, claspingthe trunk, having seriesof -F sion of Metroxylon into two sectionsbased cular transverseridges bearing knobs, or on fruit characters is not paralleled by a conspicuousspines, prickles or petioles up to several division based on Pollen tYPes. totaliy smooth: meters in tillers but considerably shorter in mature leaves, strongly channeled, TaxonomicTreatment abaxially convex, sometimes armed with -F MetroxylonRottboell in NyeS. Kon. D. transverseseries of conspicuousspines Vid. Sels. Skr. 2: 527. L7B3 norn. or piickles, adaxially concave; rachis cons.:Koenig in Ann. of Bot' ed. K. & straight, if armed, similar to the petiole S. l: I93. 1804; Endlicherin Gen.Pl. along its convex abaxial side, the adaxial Sec. Ord. Nat.: 250. IB37; Kunth in sidelifaced with the salient angle smooth; + Enum.Pl.3:213-215. 184I; Meisner pinnae straight to drooping, equidistant in Pl. Vasc.Gen. l: 35aQ65). I8a3; or in clusters,green and shiny, faintly paler Martius in Hist. Nat. Palm. 3: 2L3. underneath, linear, unicostate, acute with 1845 (ex parte Pigafetta Bhrne); a spinulous costa and margins. Inflores- Miquel in Fl. Ind. Bat. 3: I39. lB55 ceniet each branching to five orders, the (ex parte Pigafetta Mart.); Drude in E. bracts sometimes covered with spinules, & P. Pfl. Fam. 2(3):47 ' IBBT; Beccari sometimes in transverse series. Flowers in Ann. Royal Bot. Gard. Calc. 12, 2: reddish-brown, arranged in dyads of one 156-I57. lgtB; Ridleyin Fl. Mal. Pen' male and one hermaphrodite flower each, 5: 3l. 1925; Blatter in PalmpBrit. Ind. similarin appearance,the dyadsvery reg- in & Ceylon:256-259.1926; Lawrence ularly distributed along third-order axis in Gent.Herb. 8: 438. 1956. TYPe:M. spirals, depending on developmental -f sagu Rottboell. stage, immersed in the reddish-brown (greyish)hairs of the flower bracts. Male 21. Dis,q.Inaug. Sagus Steck in De Sagu flower borne on one side of the hermaph- 1757. Nominum Medica de Sagu' rodite; precocious and dropping before Moore in ?axon I l: -l- invalidum teste anthesis of the hermaphrodite; calyx and Taxon 17: 85' 164-166. 1962 deeply three-lobedin the upper third, the r968. Iobesbroadly rounded at the apex, coria- Ord. Nat. Pl.: 93' Sae.rsGiseke in Prael. ceous, ovate, obtuse, tapering below to a (excl. et S. IlgZ S. palmaPinus narrow base;the corolla I /z to more than Rumphia 2: I44-L54' f.laris); Blume in 2 times longer than the calyx, divided in 127. 1843 (ex t. 86, 126 and Parte its upper two-thirdsinto three + wood). in Palms Brit' Ind' Pigafeua); Griffith oblong-ovate,glabrous, boat-shaped seg- TYPe:S. genuina 24: IB-20. 1850. mentso the lower part entire, campanu- 1741. in Herb.Amb. I: 75. late: stamens 6, the united bases of the in Bonplan- CoelococcusH. A. Wendland filaments forming a cup adnate to the TYPe:C. uitiense dia t0: I99. L862. undivided part of the corolla, broadly lin- H. A. Wendland' ear in their free part, flattened: anthers Arborescent, solitary or clustering, elongate-elliptical, obtuse, cells parallel, attached armed or unarmed, pleonanthic or hapax- latrorse, basesdisjunct, dorsally of the con- anthic, polygamouspalms of lowlands and to the filaments at the middle represented ,*u*pt, aiso known from hillsides.Trunk nective: ovary rudimentary. + bod- columnaroto 20 m, ringed with leaf scars by three, small, coalescent,oblong and partly covered with clasping, persis- ies, merging into a prominently three-sided tent Ieaf sheaths.Leaves pinnate, erect, style. Hermaphrodite flower with calyx, L9861 RAUWERDINK:METROXYLON I7I corolla and androecium similar to the male Metroxylon syluestreMartius in l.c.: 215. flower; ovary globose and shiny with yel- lB45; Miquelin l.c.: I46. 1855. lowish rhomboidal scales. trilocular. ovules Metroxylon rumphii Mart. var. syluestre 3, reportedly basal, erect, anatropous; style Beccariin Ann. Royal Bot. Gard. Calc. narrow, prominently three-sided, grooved. 12,2:174,p. 107.l9IB. Type:Sagus Fruitscovered with scales,containing one syluestrisin Herb. Amb. l: 75. 174I . ,.eed. Metroxylon rumphii Mart. var. cera- mense subvar. ceramenseBeccari in l.c.:175, p. 107.1918. Key to the Speciesof Metroxylon Metroxvlon rumohii Mart. var. cera- *enie rubuar. platyphyllum Beccarr 1 Fruit covered with 18 longitudinal series of scales. Clustering palm, unarmed or spiny to in l.c.: 175. l9lB. Type: Beccari 12 various degrees (section Metroxylon) New (isotypeat BO). Guinea, Indonesia, Mindanao, Malaysia ...... ,.-.. Metroxyktn rumphii Mart. var. cera- mensesubvar. rubrum Beccariin l.c.: r F;;;";;;;;-ittt in iu i",'l*l,fJ,,i1,"' (isotype series of scales. Spinescent, solitary palms lack- 175. 1918. Type: BeccarlB ing basal offshoots (seclion Coelococcus) -.--..".-..2 at BO). 2. seemingly lateral, first-order Metroxvktn rumphii Mart. var. cera- branches arising from the axils of unreduced menie subuar.album Beccariin l.c.: leaves. Guam, and crown 175. l9lB. Type:Beccarl s.n. (isotype planted in the ...... 3. M. amicarum 2. Inflorescences seemingly terminal, first-order at BO). branches arising from the axils of reduced, Sagus genuina (excI. Sagou duri-rot' bractlike, crown leaves ,...... -...--...3 tozg) Rumphius in l.c.: 75. I74I; 3. Second-order branches of the inflorescence Blumein l.c.: I50. 1843. pendulous, very short (20 cm). Fiji Islands Sagus rumpAil Willdenow in Sp. Pl. 6: 3 a;;;;.;;;;;;;;;il ;;;;;;:;i;,1;!- l"'l"*n 404. 1805. 4. Third-order branches of the inflorescence pen- Sagusspinosus Roxburgh in Fl. Ind.: 623 dulous, very long (20 cm), adaxial pedicellar excl.syn. | 832. parts of all branches pubescent. Solomon MetroxvlonrumDhii Martiusrnl.c.: 214. Islands, Bougainville, St. Cruz Island...,...,-..,,..,,. " lB45; Miquelin l.c.: 140. lB55; Bec- 4.;il ;;;;;h; ;i ;; i;n;.".;;,l:,.Y; ":i.^ff^::;' cari in N. Giorn.Bot. Ital. 3: 30. 1879 pedicellar parts of all branches not pubescent. et in Malesial: 91. IBBT; Hooker in New Hebrides,Western ...,...... ,...,-..,...,,...- Fl. Br. Ind.6: 48I. 1894.Type: Sagas .,.....5.M. uarburgii genuina in Herb. Amb. I: 75, t. 17. 1741. Metroxylon rumphii Mart. var. cera- i. Metroxylon sagu Rottboellin Nye. mensesu.bvar. nigrum Beccariin Ann. S. Kon. D. Vid. Sels. Skr. 2: 527. Royal Bot. Gard. Calc. 12, 2: 175. l7B3; Miquelin 1.c.:I47. lB55; Bec- 1918. Type: Beccari s.n. (isotypeat cari in N. Giorn.Bot. Ital. 3: 29. IB79; BO). Hookerin Fl. Brit. Ind. 6: 48I. 1894; Metroxylon rurnphii Mart. var. buruense Beccariin Denk. K. Akad. Wiss. M. Beccariin l.c.: 177- l78. 1918.Type: Nat. Kl., Wien,84: f. 6. 1913.Type: Labillardieres.n. Koenig specimensand drawings and Metroxylon squarrosum Becc. var. Rottboelll.c. 1783. souarrosnmBeccari in l.c.: I79-180. Sagus syluesrrisRumphius in Lc.: 75. l9tB. l74I; Blumein l.c.: 153. I843. Metroxylon seuarrosum Becc. var. kil- Sagus genuino. bis syluestris Ciseke in woi Beccariin l.c.: lB0. 1918. Type: Lc.:94. 1792. Beccari 1 (isotypeat BO). lVoL. 30 t72 PRINCIPES laPia, sagu, Lumbia, Metroxylon squanosum,Becc' var' kil' kiraj, ambulung, lasi Beccariin l.c': 180. 19lB' Type: 'pohon sagu. Examined: These sPeci Beccari 2 (isotYPeat BO)' Speciiens hurn"been identified as belonging to Metroxylon squatosum Becc' var' ldl- rn"n, M. sagu. but a further iden- karuaBeccatiin l.c': I80' 19lB' Type: the species as one of the four formae in M' Beccari 3 (isotyPeat BO). Lificaiion not been possible(veg' : veg- Metroxylon squarrosum Becc' var' lcila' sagu has no flowers or fruits available; tan Eeccariin l.c': lB1. I9lB' TYPe: etJtive, i.e. : flowers; fr. : dispiaying Beccaris.n. (probably4, isotypeat BO)' fl. displaying : the specimenis past fl:Ig: Metroxylon sqiarrosum Becc' var' kil' fruitsl pfl. not forrned any fruits)' NEW kour-Beccariin I'c.l lBl. l9l8' Type: ine bui has Sepik, 1953, FloYd 5ao9 $' Beccari s.n.(probably 5, isotypeat BO)' CiitNg,t. L); VogeikoP, 1948, Kostertnans G Metroxylon sqiarrosum Becc' var' kil- BO, (fr., Bb). INDONESIA, CELEBES, tafuiBecciri in l.c.: IBI. I9lB' Type: et H KoorrJers19722 bis (veg', BO, L); B-eccari6 (isotYPeat BO)' 1895, Korthalss.n. (veg.,L); ' Metroxylon squ&rrosum Becc' var' kil- , Schoute s.n. (fl', L), 1882, Treub kikii Beccaiiin l.c.: l8l' I9IB' Type: 1903, (fr., B), lBB5, Treub s'n' (fr'' B); Beccaris.n.(probably 7, isotypeat BO)' s.n. LBB3, Warburg s'r'' (fr', B); Metroxylon rumphii Mart. var' flyriuer- BATJAN, d'Atleizette77l5 (fr', L, enseBeccari in l.c.: l78-179' I9lB' CULTA, 1909. Bakhuizenu/d Brink 1544 Type: D'AlberJis s.n. SBG).1922. (fr., 1958, Heine s.n' (fr', BO), Meiioxyton ox'ybracteaturn Warburg BO), 35664 bis (fr', BO), ined. nomen nudum; Schumann & Lau- 1899, Koorders s'n. (fr., B). terbachin Fl. d. Sudsee:202' I90I' Schlechter stem Clustering Palm, 8-20 m high, Key to the Formaeof M. sagu l5-60 cm diameternear baselleaves up ages of the to ? m in mature Palms, uP to 12 m in I. Leaf sheaths and petioles at all -l- ""'-"""lorma sagu offshoots,unarmed or armed oalm unarmed, totally smooth vigorous and petioles not quite smooth or on transverseridges I. Leaf sheaths wilh spinesor spinules armed -..,-.-....-.,..-....-. "' """"'-"""""""""' 2 o, ,"ri"r. Infloiescenceapparently termi 2. The bases of leaf sheaths at all ages of the nal with first-o second-, and third-order oalm with knob'like structures (spine reduc- fotma tuberatun brancheserect to spreading(Fig' 8)' Bracts iior") .,,,-.....-.,.,....-.,,,. petioles bearing spines '- 3 and third order with spi' 2. The.leaf sheathsand of first, second, and petiolesbeaing spinesshorter flower, 3. Leaf sheaths nules or smooth; in hermaphrodite than 4 cm forma nicracanthum style attains apex of the anthers at anthe- 3. Leaf sheathsand petioles bearing spines of 4- sis. In male flower ovary rudimentary and 20 cm long fotma longisPinunt' style not exceeding filaments' Fruit glo- bose, apple-shapedto shaped like an sagu formasd$u (Fig' 't la. Metroxylon inverted pine "o.t", "ty variable in shape e). and size within an inflorescence,4-4'5 laeuisBlume in l.c.: I46' lB43' cm broad, lS (19) longitudinal seriesof Sasus laeue Marritts in l'c': 214' straw colored scales;seed frequently abor- Mitroxylon TyPus: Koenig sPecimensad tive" fruit then filled with a fleshy hyper- 1845. et Rottboell ut omittam quo trophic mass. Rottboell bistribution: New Guinea and widely ad Rumphius. Rumphiusin I'c': 76' 1741; planted throughout Indonesia, Malaysia, Sasuslaeiis in Calc. J. Nat' H. 5' 1845 et and the PhiliPPines. driffith Ind. 24' IBSO' Vernaculir Nam'et sah-salt, rumbia, in Palms Br. RAUWERDINK:METROXYLON

"terminal" B lI. B. Metroxylon sagu, habit showing inflorescence.9. M. sagu, fotma sagu. IO. M. sagu forma tuberatum. I I. Same, petiole at higher magnification showing knoblike structures (reduced spines). PRINCIPES

";;;; l2_li.12.Metroxylonsaguformamicracanthum,ia,.sheathanilbaseofpetioleshowingcurvedrowsoflong rows of spines' itt^' tongispina' leaf sheath showing spines; b, close up "f tpi;'';;:'M' taeuisGiseke inr'c': sagusgenuina s. li:lff;lt:?t $";;:t:1'li:T"ifi:Tl 94' 1792' are smoothtoo' Fl'r Ind' 3: l'u"t' or the inflorescence So,gu,iner*is Roxburgh5' inur apex Th" pi""u" may be spinulousat the 623. rB32' l: and the costa' S"S^'t""rit Jackin Comp')'rrv' Bot'uvL' Mag' Vernacular Names: ambutrurn' 266. L836. in kaparang' auirkoma (PNG)' Metroxylon herntaphrod'ilumHasskarl Tp"'til""nt Examined:NES/ GUINEA' Tiid. Nat. Gesch.9: I75' lB42' (fr'' A' 215' Mihie Bay' 1953' Brass22064 Metioxylon inermeMartiusin l'c': (ft'' ijt S"ptfa lgSl"Rauwerdink 112 1845. Typus, Sogu' i;"is in Herb' WnCl, Rauwerd'ink 119 (pfl'' WAC)' Amb. I: 76.t.17. L74;' MetroxylonsogusRottb.var.molatBec-Rauwerd'ink123(veg.,wAG),Rauwer-ne (veg., INDONESIA' cariinr.c.:167,p. ioz. tqta. Typus: il* -wAG). SUifnATRA' I"933' Rohmat si Torus Beccaril0 (isotypeBO)' siis (pfl'' A' L)' PHILIPPINES'MIN- Metroxylon ,ogu, ttott var' peeke' (ft'' leNe'b' !9I2' Fenix 1112 P)' lianumBeccari in l'c': 167-l'68' p' 1gt5' Mt'os M-edj1'a24526 (pfl'' A)' 106. 1918. Typus:Peekell evtueLll5"" (not uereysu,. SABAH,1964, Moore 9218 seen). .'""" gU)' CULTA' 1929' Bahhuizenuf MetroxylonsogzrsRottb' vat'gog2l1rys1: ift- i il'ink 7488(fl'' B' BO' L'P)' 1952' Beccari""ryp,",in l.c.: 168, p' i06' l9l8' e6i(not seen). r"*,,,ao,r, X;;fr{.?Ka;:-,art#,n )},7},".!.tili:, K)' The leaf sheath,petiole, and rachis of Le86l RAUWERDINK:METROXYLON 175

lb. Metroxylon sagu formatubera- WAG), Rauwerdinlt 118 (pfl., WAG), tum Rauwerdinkform. nov. (Figs. Rauwerdink 127 (fl., WAG), Rauwer- 10,1 l) vaginae frondinum tuberatae. dink 134 (veg.,WAG). Typus: New Guinea. E-Sepik prov., (pfl., I9BI, Rauwerdink 715 holo- ld. Metroxylon sagu forma longi- type WAG; isotypi BO, BH, CANB, spinum (Ciseke)Rauwerdink stat. K, L, LAE, SIN). nov. (Fig. l3). Typus:Rottboell pro The base of the leaf sheaths covered parte quo ad Rumphius. ,,'ith knoblike structures, irregular to the Sa.guslongisplna Rumphius in l.c.: 75. touch, at all ages of the palm. Petiole, l74l; Blumein l.c.: 154. 1843. rachis. and all bracts of the inflorescence Sagusgenuina longispina Gisekein l.c.: perfectly smooth. The pinnae can be spi- 94. 1792. nulous at the apex and costa. Sagus longissimus Hamilton in Mem. L'ernacular l,lames: koma, oliato,goe Wern. Soc. 5: 320. 1824. (PNG). Metroxylon longispinum Martius in l.c.: SpecimensExaminecl: NEW GUINEA. 2I5. 18450;Miquel in l.c.:146. 1855. Sepik, 198I, Rauwerdink 115 (pfl., Metroxylon rumphii Mart. var. longi' WAG), Rauwerdinh 135 (veg.,WAC). spinum Beccariin l.c.: 173, p. i07. 1918. Typus: Sagas longispina in Herb.Amb. I:75. 1741. tc. Metroxylon sagu forma micra- canthum (Blume)Rauwerdink stat. The leaf sheath,petiole and rachiswith nov. (Fig. l2a,b). Typus: Blume in spinesthat are 4-20 cm long. Bracts of Rumphia 2: I53 teste Rumphius p. inflorescencewith spinules,scattered or in 75, sub.no. l, SaguDuri-Rottang. transverseseries. Vernacular Names: uakar. ketro. Sagusduri rottang Rumphiusin l.c.: 75. tring. t74r. anum"ninginam6. nago. passin. kangrum. nombarang.moia p. Sagus micra.cantha, Blume in l.c.: 153. SoecimensExamined: NEW GUINEA. I 843. Malalaua.1966. Grauen& Schodde9'tr'tr Metroxylon micracanthum Martius in l.c.: (fl., A, L), Grauen& Schodde945 (fl., 146. 1845; Miquel in l.c.: 146. 1855. A, L); 1938, Meyer Drees 378 (fl., fr., llletroxvlon rumnhii Mart. var. micra- BO); Sepik,1981, Rauwerdink 100 (fr., canthum Becc. subvar. micracanthum WAG), Rauwerdink 101 (fl., WAG), Beccariin l.c.: 176. 1918. Typus: Saga Rauwerdink 102 (fr., WAG), Rauuser- Duri-Rottang in Herb. Amb. l: 75. dink 104 106 (veg., WAG), Ra,uuer- 174I. dink 107 (pfl., WAG), Ra.uwerdinhl0B Metroxvlon rumnhii Mart. var. micra- (veg., WAG), Rauwerdink 110 (veg., ncanthum Becc. subvar. makanaro WAG), Rauwerdink 111 (pfl., U/AG), Beccariinl.c.: 177.l9lB. Tvous: Bec- Rauwerdink 113 (fr., WAG), Rnuwer- ca.ri 14 (isotype BO). dink 117 (veg.,WAG), Raunerdink 120 The leaf sheath. oetiole and rachis from (veg., WAC), Rauwerd,ink 121 (fl., an early age of the palm with spines not WAG), Rauwerdink 124 (veg., WAC), longer than 4 cm. Bracts of the inflores- Rauuerdink 125 (veg.,WAG), Rauwer- cence smooth or with scattered spinules. clink 126 (veg.,WAG), Rauwerdinh129 Vernacular Names: makapn, uttipi, (veg., WAG), Rauwerdink 130 (fr., Itangrum, mandm (PNG). WAG), Rauwerdink 132 (fr., WAG), Specimens Examined: NEW CUINEA. Rauwerdink 133 (fl., V/AG), Morobe, Sepik, 1981 , Rauwerdink 114 (fr., 1964, WontersleyltrGF 19234 (pfl., fr., 30 PRINCIPES lVoL.

14_15.Fruitshapesandsizes,eachfromasinglecollection.Figure14fromRauwerdinltll2'Figwe|5 from Rauwerdink 100' (H' A' Wendl') ISLANDS1964, Moore Metroxylon crmicarum A, L); SOLOMON Beccari in l'c': %ls asP 4ogo(fr,., sIN, K). INDo- Becc. var. commune lglB (note:This nameis a homo- NESIA.Bengkalis, I9I9, Beguin 19.19 188. synonym.even thoughBeccari (fl."fr., gO);CeLgeES, 1895, Koorders tvpical 'lB3s7 L"d"rmann 134091' bis (veg.,BO): AMBON" l9I3' typifledi *iri in Bot' Robinson28 (ifl., L, K, P);JAVA, 1843' SagusamicarumH. A. Wendland Rei- Zollinger 14d4 (fl., fr., B, P). PHILJ-I' Zeit. 36(8): l15. 1878' TYPus: W)' PINEs.r94r, Ebalo999 (fl.,BH); MIN- chenbachs.n. (not seen,possibly at DANAO,1909, Elmer 11160 (Pfl',fr', ProposedneotyPe: Furtado 25868 A, C, L, K, P), L927,Ramos & Edano (holotypeK, isotYPeA)' 49399(fl., fr., BO).CULTA ' 1924,Bak- CoeLococcuscarolinensis Dingler in Bot' (veg',-BO)' huizen u/d. Brink 3497 Centr.Bl. 32:349, t. 2' IBBT; Schu- Bloembergens.n. (Dl) (fl', B0]r Bl2'!- mann & Lauterbachin Fl' D' Sch' Sud- bersen.'i-t. (VL 28) (veg',BO), lS3S' see:606.1l901. Typus: Schneider s'n' BuTrrt3J3 (fr.-BO)' 1952' Moore 6O95 (not seen). 1895,RidIeY s'n' (fl', K)' (veg.,BH), Coelococcuscarolinense Beccari in Wiss' M' Nat' Kl' (H' A' Denkschr.K. Akad. 2. MetroxYlon-B"c"a.i amicarum (Rechingered.), Wien, 84: 60, fig' 5a in Ann' Royal Bot' Wendl.) & d. lOl3. TYPus:Beccarl s'n' (not 12,2: IB7-I92, ILI- Gard.Calc. P' seen). l9l8; Mooreet Fosbergin Gent' I13. (H' A' Wendl') Herb. 8, Fasc.6, att' 20: 438-444' Metroxylon annicarum majus Beccariin Ann' Royal 1956;Barrau in Econ'Bot' l3: 15I- B"cc. ,r.r. CaIc.L2,2: tB8' 19iB' 159.1959. Bot. Gard' RAUWERDINK:METROXYLON 177

Typus: Gibbons 1189 (not seen, pos- Bonplandia l0: 199-200, 1862. sibly at B). Typus: Seeman 658 (holotypeK, iso- type P). Solitary, pleonanthicpalm, 6-8(20) m high, stem to 30 cm in diameter near base. Solitary, hapaxanthicpalm' 5-i0(15) Leaves ca. 6 m long, with spines. Inflo- m high, stem to 50 cm in diameter near rescences arising from axils of crown base. Leaves 4-5 m long with slender leaves,i.e. thesenot bractlike,reduced in spines,brown to chestnut-brown.Inflores- size as in other Metroxyloz species.Var- cence apparently terminal, first-order ious inflorescences differing in develop- branches erecto-patent,2-2.5 m long, mental stage. Infructescence pendulous second-order branches pendulous, those due to weight of numerous big fruits. In seen by me 20 cm long. Style in male hermaphrodite flower, the style attains flower not exceeding the filaments, style apex of anthers at anthesis, but in1 male in hermaphrodite flower as long as sta- flowero gynoecium much reduced Ito3to mens. Fruit turbinate, yellowish-brown, connate, oblong bodies, reaching only to about 7 cm high and 5 cm wide, with the baseofanthers. Fruit globoseto apple- 27(28) longitudinal series of scales. shaped,broader than high, 6-9 by 5-B Distribution: Fiji-Islands' cm, or higherthan broad,7-I3 by B-12 Vernacular Name: songa. cm, covered wtth 24-28longitudinal series Sp ecimensExamined : FIJI-ISLANDS. of chestnut-brown(reddish) scales. 186O,Seeman 658 (fl., K, P); Viti Levu, Distribul ion: Guam, Caroline-lslands. 1953. Smith 9339 (fl., GH, K, L),1969, Vernacular lYames:r6pung (Truk), o7 Tomlinson & Kuruuoli 16551 (fl., fr., (Ponape). BH). SpecimensExamined: GUAM. 1914, lt{elson s.n. (fl., A, P). CAROLINE- 4. Metroxylonsalomonense (Warb.) ISLANDS. Truk, Fosberg 24522 (fl., fr., Beccari in Denkschr. K. Akad. Wiss. BH), Kriimer s.n. (fl., B), 1947, Wong M. Nat. Kl. (Rechingered.),.Wien, 84: 244 (fl., A); Ponape,I9L4, Kersting s.n. 60-62, fig. 5b & e & f & i,lt 7. (fl., B). CULTA. 1903, Loher s.n. (fr., t9l3 et in Lc.: I92-L93, p' l14. K), 1907, Lyon s.n.(fr., K), 1929, Anon. l91B; Barrau in Journ. Agr. Tr. B. (fl., fr., SIN), 1932, Furtado 25868 (fl., Appl., 3: 32-36. 1956 et in Econ. fr., A, K), 1954, Kiah ONO 114 (fl', Bot.l3: l5I-159.1959. .Warburg BH). Coelococcussalomonensis in Ber.D. Bot.Ges. 14: I4I, t. 10. 1896; (H. A. Wendl.) 3. Schumann & Lauterbach in Fl. D. Sch. Bot. Gard. Calc. Beccariin Ann. Royal Sudsee:606. I90I. Typus: Warburg ll0. l9l8; Bar- 12,2: 185-186,p. s.n. (fr.), (not seen, possiblyB). Pro- Appl., 3: 32- rau in Journ. Agr. B. posedneotype:. Kajewski 2300 (B). et in Econ.Bot. 13: l5I- 36. 1956 Metroxylon bougainuillense Beccari in r59.1959. Denkschr.K. Akad. Wiss. M. Nat. Kl. .Wendland Sagusaitiensis H. A. in Seem. (Rechingered.), Wien, 84: 60*62, fi9. Fl. Vit.: 279, t. 80. 1865-68 et in 5c & g & h & j. l9l3 et in Ann. Royal App. to Seem.Vit.: 444(teste Beccari); Bot. Gard. Calc. 12, 2: I93-I94, p. Benth.& H. in Gen.Pl. 3: 935. 1883; l14. 19l8; Barrau in Journ. Agr. Tr. Drake de Cast. in lll. Fl. ns. Pac. B: B. Appl., 3: 32-36. 1956 et in Econ' 323. 1886-92; Warburg in Ber. D. Bot. l3: l5I-I59. I959. Typus: Bot. Ges.14: I4I. 1896. Rechinger 4878 (fr.), (not seen, pos- Coelococcusuitiensis H. A. Wendland in siblv W). 30 PRINCIPES lVoL. I7B ridges or series' Inflorescence haPaxanthic 9-20 m verse Solitary, Palm, terminal with erecto-patent 55 cm in diameter near aonarently high, stem up to l-l'5 m long' sec- with spinesin fi'r.t-otd"i branches, Uu".".t"uro"t 9-II m long, erecto-pat' appar- ond- and third-orderbranches transverse series. Inflorescence "nU udu*iul sidesof all pedicelsglabrous' ".ritv ,"t-i"al with first-order branches as Flowerslarge, corolla(10 mm) twice "."Jto-pate"t, 3-4 m lorrg, second-order lonq as calyx and apex minutely punctate; ;;;;"t"t sPreading but third-order at in t"r*uphtodite flower style attains p".td.llo"t, ca' 20 cm long' b.un"h". lengthof filaments;style.in greyish-silvery pubes- i"urt tutfit't" Fio*", bracts with go*". fused part of fila- flowers ,nui" as lo-ng as cence. Male and hermaphrodite seed- th: reflexedsub- ;;;t. Fruit Pear-shaPed, .T with filamentswith a sharply (4i?-l cm high and not ,ro"o"r.ftoua.r'parr, 2 ulate top, male flower with stYle l{.ilo-s cm bioad. with 24 longitudinal """""ding f.rted part of filaments' whereas seriesof chestnut-brownscales' l" tt".*"lpn-dite flo*er, style not reach- Distribution.'New Hebrides' W-Samoa' irrs tr"vondpoint o[ attachmentof anthers' Vernacular Nam'ex tenebee'niu oLo' f."uit gtolot". depressedat apex3ld-bu-t"-' )9 broadand 5'5-6'5 tuma. .tru*?oto."a,7 cm NEW HEBRI- series S,pecimensExamined: "- ttlglt, with 24-27 longitudinal (fl', P); DE5. 1964, Moore 9319 of-- scales. rrilrnrure,lg7l, HalleRSNH 6344 iistribution.' ' Sol- K, r); PENTqgglr Cruz-Islands' il G- fr., A, omon Islands,Santa ig+g,Ma,Daniels 3011 (fr'' Names: bia, naml' hae- isreNbs, Vernacular eH).W-SAX'{OA' UPOLU, 1968' An-on-' sa,n. (fr'' Tiist (fl.,'nr*totK, P), Bristol217I GH' SnecimensExamined: PAPUA NEW (ft', GH)' 1971' ISLANDS, K-f), 2180 CUT.TqC,q.BOUCAINVILLE (fl.,fr., BH), 1978,Mo-ore 2300 (fl', fr', tti"oii'loisa 9985 tOiO,,tcolr*ski P' 99' (fl.,f'.;BH). cULTA' 1891, Rid- Schod'd'e4013 et 4013a (fr"' Eul,'tgo+, l"y slil ifl.,fr., K, sIN)' L94L,Anon' 'i"rrr'r.i.tl.,'t , L). soLoMoN-ISLANDS'I936' (fl',fr'' K), 1964Moore 9317 s.n. (fr., B). A-S!PAOSI (fl', fr', K, L, SIN)'Mg!ry sioz (rr.,BH).sANTA cRUZ-ISLANDS' Discussion L' WhitmoreBSIP 1818(fl" K' is 19t4,' The first question to be considered sIN). (the sago why M. rumphii \4art' spiny .yttotty*ous-with sagu (Heim)Bec- f.f*) tft.tfa b" !I' 5. Metroxylonwarburgii My main argu- Bot' Gard' Calc' ithe spinelesssago palm)' "uri in inn. RoYal more than one l9iB; Bar- ment against recognizing 12,2: lB2'IBa,P' 109' basedup.on 3t .o""i"rIn the real sagopalm is rauin Journ'Agt' tt' B' APPI'' ??- tn ltt- plr.ot"l observationand references 36. 1956et in Econ'Bot' 13: 151- erature: a spmy sago palm can produce ls9.1959. from-,a both spiny and'spinelessseedlings Coelococcuswarburgii Heimin S"L !:: sineleinflorescence' For exampleJumelle been Bot. Fr. 5o: 572, fig' l-5' l9o4' iiSiSl menlionsthis' and I have rn Tvnus: Heim s'n' (fr')" (not seen'pos- able to verify that this situation occurs ,i6iv HAL). Proposedneotype: Ridley pttlC (lt"ifiikir area, road to- Maprik' questron .lIi I tholotvP"StN'isotYPe K)' West-SepikProv')' The second I have not upheld 6-7 m ,. U" uo'.*"ted is why Solitary' haPaxanthic Palm' the real sagopalm nearbase' Beccari'staxonomy on hieh,stem to 3d cm h diameter a srn- which, even though not applying to LJ"lr", ca' 3 m long, with spinesin trans- r986l RAUWERDINK:METROXYLON ele species situation, covers a remarkable range of variability. It might not prove too hard to maintain at least some of his vari- L L L K eties. As can be seen in his key to the taxa (Beccari in Lc.: 160 162. l9l8), Beccari divides species in the real sago L L K palm into varieties and subvarieties on the bases of fruit characters such as size and shape. I do not agree with these distinc- tions because I have observed a consid- L K erable variability in fruit shape on the inflorescence of a single palm. Figures 14 to I 5 represent samples of varying shapes K I S ,.elected from material collected in the Sepik-area. Each photograph represents a single number from my collection. I have 16. Diagram presenting the possible genotypes of compared my materials with type material the formae of M. sagu where two genes control spine Iength. L, longispinuml K, micracanthuml I, tu- of Beccari at the Herbarium Bogoriense beratum; and S, sagz. and have found that a single accession collected by me may include several of Beccari's taxa. For example, M. rumphii that a minimum of two genes or alleles Mart. var. ceramense subvar. nisrum and governs spine length in its broadest sense. subrar. platyphy!lum material both dis- Such a situation can be expressed in Fig- play fruits that are similar to Ra,unerdink ure 16, which presents the genotypes of 112 (Fig. l4). Therefore, I have to con- the occurring forms in this concept. The ciude that fruit shape and size cannot be thesis is rather academic, however. and useful in typifying taxa in the real sago can only be tested in the field, but this is palm. These characteres are most likely unfortunately seriously hampered by the very dependent on age and location of the lack of knowledge on the pollination fruit in the infructescence. mechanism in Metroxylon, and the fact For each part of the palm and for each that fruits are eaten by a number of ani- developmental stage, spine characters are mals. Not much is known for certain, but independent of age, and hence suitable fbr it is suspected that hornbills and fruit bats use in classification. It is convenient that eat the fruits at various stages of devel- this classification can take place nearly at opment (local inhabitants Sepik-area pers. ground level in the field. Furthermore a comm.). Wild boars are known to eat fruits classification on spine characters offers a on the ground (Rumphius l74l). - very clear distinction between the various thermore, particularly the formae saga., forms of M. sasu. The taxonomic rank of tuberatum and micracanthum, long the taxa in M. sagu should be low. The domesticated by man (e.g. the Sepik in fact that a spiny palm may produce spiny PNG), are almost always cut down for and spineless seedlings indicates that it is starch extraction long before fruits even heterozygous for this character and may start to develop. In the rare cases that produce embryos of at least two geno- fruits are oroduied in these formae I have types. The classification forma should been unaLle to find seeds with a well- apply to these taxa rather than variety or developed embryo and endosperm, seeds subvariety. The offspring of the aforemen- containing instead a hypertrophic cellular tioned young palms may not be of the mass. In the taxon longispinum fruits do genotype of the mother-palm. I assume frequently develop an embryo and endo- lVoL. 30 180 PRINCIPES A Taxonomic Revision' Misc' taxon is much less often cul- eonia Warb' sperm; this Faoers 19: 377 -421. Landbouwhogeschool' Liber Gratula' tivated. Wue"ninge.,-hotor"- The Netherlands' t..iJs in H. C. D. De Wit' Acknowledgments DE WILDE, J. J. F. E. axl AnrNls, J' C' 1979' Besonia loranthoides Hook' F' (Sect' Tetro' I would like to acknowledgethe assis- ptlta t. DC.). Acta Bot. Neerl' 2B(4/5):357- tance and kind attention of everyone con- 374. Newcombe, Holmes and Paivoke in with the Department of Plant Tax- DME 1980. nected Palm energy-alcoholfuel from the sagoand nipa StudY on sago Guinea' Report 6/80' onomy. Palms, .at 'Energyoalms of Fupta N"w Wageningen was' howeYer, initiated by Planning Unit, Dept' of Minerals and P.oi.dr.it. M. Flach of the Departmentof Energy, Konedobu, PNG' and Holmes in Implemen- Tropical CroP Science. DME 1982. Newcombe alternative fuels in developing coun- are due to Prof'dr'ir' L' tation of Specialthanks alcoholfuels in Papua New Guinea' Report .rran Bos tries: J. G. der Maesenand Dr.ir' J' J' 7 /82, Energy Planning Unit, Dept' of Minerals for their supervisionover my work, to Dr' and EnergY, Konedobu, PNG' Coloniales,Plantes a J. Dransfield(Kew) for his' much appre- Juutrr-n, tt. ISZS. Cultures et Cereales' Paris' ciated, interest in my work, to Ir' J' C' Fecule eNo N. W. Uur. 1982' Major on the cytological part Moonr, H. E., Jn. Arends for his help trends oI evolution in palms' Bot' Rev' 48: 1 of this paper and the photograptr h"-pt:- 69. vided of ihe chromosomes,to Dr' R' G' PRecrowsrt, J. er'{o W. PuNr' 1973' An eluci of the van den Berg for his help on the palyno- dation of the Microreticulate Structure Grana I3: 45-50' logicalpart of this paper and the photo- Exine. I97?' Papersof the lst' Int' SagoSymp'' of the pollen grains, SAco'76. er;phs he provided Kemajan Kanji Sdn. Bhd', Kuala Lumpur' a nat' io MissH. C. D. Zewaldfor her finedraw- SAGo'80. 1980. The equatorialswamp as ings and the preparation of all illustra- ural resource. Proceedingsof the 2nd' Int' Sago Hague' to Mi. J. W. Mugge who printed Svmp.. Martinus Nijhoff, the tiolns,and Pollenmorphology of the- photographs' Sowuvni, M. A' I972. all macromorphological and its bearing on taxonomy' Rev' of to the Palmae Furthermoie I am indebted Pal. Bot. and PalY' 13: l-80' Directors and Curators of the herbaria of Teau Pnovrr. 1982. P3 S.B.P'E' (SAGU)' Hasil : Bagain A, B, BO, BH, GH, K, L, LAE, P ANd survai potensi sagu di kepuluan II dan bagain III' Jakarta' SIN for kindly putting their material at I, bagain TueNireil,loNr, G. l9?0. Pollen morphology' clas- my disposal. sification and phylogeny of Palmae' Adansonia' ser. 2, I0(3): 347-365' P' B. I97I' Flowering in Metroxy- Ltrenerune Ctrun TouLtNson, loz. PrinciPes 15: 49-62' 1977. Papersofthe lst Int' SagoSymp': DE WILDE, J. J. F. E' (taxonomy) ANDJ' C' ARENDS VrRtr,len. (cytology). 1980. Begonia section Squamibe' 155-l)o.