l99sl ESSIG:PALMS OF BISMARCKARCHIPELAGO

Principes,39(3), I995, pp. 123-129 A Checklistand Analysisof the Palmsof the Bismarck Archipelago

FneoenlcrB. Essrc Institutefor Systematic Botany, Department ofBiology, (Jniuersity ofsouth Florida, Tampa, FL 33620

This analysisof the palm flora of the Bismarck them. However,surprisingly few New Guineapalms Archipelago, although certainly incomplete, is have crossedthe small gap. The intended both as a practical guide to the palms, begin officially with Bougainville, some 120 miles and as a small contribution to the phytogeography east of , with only a few very small of the . The Bismarck Archipelago is a sub- islandslying betweenthe two archipelagos.A num- stantial chain of islands off the north-east coast ber of palms have apparently migrated from the of , and is part of the independent Solomonsto the Bismarck Archipelago. nation of New Guinea. The chain itself is Most palm in the western Pacific region "U," bent into I Iying on its side just south of are fleshy and rather heavy, and are presumably the equator. The two large islandsof dispersedonly short distancesby animals (Nypo and New Ireland form the bulk ofthe archipelago, and Cocosare obvious exceptions).Some genera but both sidesof the U-shapedchain extend west- with large fruits, such as Orania, are absentfrom ward in strings of small islands. The chain ter- the Bismarck Archipelago,although abundantand minates on the north side with (the diverse on New Guinea and some of the islands ), and on the south side with a to the southeast.Even genera with smaller fruits seiies of small volcanic islandslying closely along (e.9., Calamus, Gulubia, Gronophyllum, the north coast of New Guinea (Fig. l). Hydr i as t ele, Caly p t r oc alyx, H et er os p at h e, and. The largest of the islands, New Britain, is rug- Cyrtostachys) are each representedby only one ged and mountainous, consisting of over 36,000 or two speciesin the islands. On the other hand, squarekilometers, with peaks tp to 2,250 meters somegenera with large fruits(Ptychococcus, Acti- in elevation. The geology is complex, large areas norhytis) are found in the Bismarck chain and in of raisedlimestone alternating with extensiveareas the Solomons. of volcanic rock and a number of active volcanos. Genera with endemic speciesin the Bismarck New Ireland is much smaller and for the most Archipelago are , Gronophyllum, part quite narrow, but the southeastern end is Heter os p athe, Hydr iastele, Caly p tr ocalyx, Cyr - broad and mountainous, with peaks over I,800 tostachys, Physolcentia and, Clinostigrna. The meters high. The higher elevationson both islands first six genera are widely distributed in New sustain montane forest and some of the unique Guinea. The last two are distributed through the elementsof the palm flora. Solomon Islands and down to but not in New Moore ( I 969b) consideredthe palm flora of the Caledonia. The remaining genera found in the Solomon Islands to be primarily an extension of region are represented by common, widespread that of New Guinea, with some contribution (Cli- species. nostigma, Physolcentia) from the ancient Pacific Genera for:nd on New Guinea but lacking in flora that survives primarily in .As the Bismarck Archipelago arei Bora.ssus,Bras- one would expect, the palms of the Bismarck siophoenix, Corypha, Daemonorops, KorthaL Archipelago show an intermediate character sia, Linospadix, Pinanga, Sommieria, andPiga- between those of New Guinea and those of the fetta. Most of these genera are fairly restricted Solomon Islands. New Britain is separated from in their distribution, even in New Guinea, and not New Guinea by only 83 kilometers, the volcanic suitably positioned for migration eastward. Kor- serving as a stepping stone between thalsia, however, is widespreadand common in t24 PRINCIPES [Vor.. 39

Manus I. EF New Hanovert]

Long I. ""rr.,t"Ut\ (Solomon Is.\.-)

"Q)

(\1

Map of the Bismarck Archipelago.

New Guinea, and its fruits are small. It \ryouldnot annotations(as well as some by Moore), and my be surprising eventually to find it somewherein mention of them in my Palm Flora of New Guinea the Bismarck Archipelago. (Essig1977), regrettablymay have lead to erro- Rhopaloblaste and,Liuistona are found in both neousreports of the occurrenceof Drymophloeus N_ewGuinea and the Solor4onIslands, but seem in the Bismarck Archipelago (as in Moore L969b, to be lacking from the Bismarck Archipelago. Hay 1984). Drymophloeus is found only in west- Moore's(1969b) indicationthat Liuiston@occurs ernmost West Irian, skips over the remainder of here appearsto have been in error, although one New Guinea,and reappearsin the SolomonIslands, would certainly expect it. Several speciesof this assuming one accepts the inclusion of Burret's occur in New Guinea, including L. wood- gentlrisR ehdero p hoenix in D ryrnop hloeus (Moore of fordii Ridley, which also occurs in the Solomon 1969a).The tremendousgeographic separation Islands(Tulagi, Esa'ala).To date, however, I have these two parts of the genus, and the fact that seen no collectionsfrom this region. the characters by which Drymophloeus is distin- One genusthat has diversified among the islands guished from the other genera of the Ptychos- is Ptychospenna) a large genus centered in New permatinae are primitive for the alliance as a Guinea, and representedby two endemic species whole, suggest that the issue should be further in the Bismarck Archipelago,and at leastone more examined. in the Solomon Islands. None of these seems to One final observation-These islandswere set- have any closerelatives in New Guinea, however, tled by seafaringpeoples many centuriesago, and with P. hentyi being an unusual endemic in New there has undoubtedly been much interaction Britain, andP. gracile appearingto be most closely among the peoplesof the islandsand New Guinea related to Solomon Island and Australian species. since then. We must take into consideration' Ptychosperma subgenusActinophloeus, on the therefore, the possibility of human introduction other hand, is lacking altogether from the islands, and cultivation of some of the palms. Ptychococ- although it is widespreadin New Guinea. cus and Caryota, for example, have very hard 'When "wood" I first encountered specimensof Pty- used by the local people throughout the chosperma hentyi, I assignedthem to Drymo- region for bows,spearheads and a variety of other phloeus based on the elongate peduncles of the purposes(pers. obs.).It is quite possiblethat early ,the broadly cuneate, apically con- settlers or traders brought seedsof these species vex leaflets,and the apparently globoseseeds. My with them. In both genera mentioned, the single r9951 ESSIG:PALMS OF BISMARCKARCHIPELAGO

species present in the Bismarck Archipelago 9a. Palmswithstiltroots...... I0 appears to be indistinguishablefrom speciesdis- 9b. Palms without stilt root. . . . : : : . : : : : . : : : : : : . : : Il tributed broadly through New Guinea and into the I0a. Fruits dark purple; seedsharply angled; montane for- Islands. est,NewBritain ...... Physokentiaaaia Solomon Similar interpretations can be l0b. Fruits redi seedsterete in cross-section;montane for. argued for Cocos nucifera, Metroxyl.on sagu, est, New Ireland . .Clinostigma collegarum Areca catechu, andAreca macrocalyx, as lla, Palms massive, clumping, producing a massive inflo. all are ' of economic importance. rescence at the apex of the trunk; bases with sinuous spiny ridges . . .. Metroxylon sagu I lb. Palms of moderatedimensions, or with solitary tnmks, Synopsisof the Palmsof the producing inflorescencesin sequencefrom the axils .; BismarckArchipelago of the ;leaf basessmooth ...... 12 I2a. Pinnae conspicuouslytoothed or ragged at the tips l3 The only previous work on the life of the I2b. Pinnae acute or briefly notched at the tips ...... 13 BismarckArchipelago was that of Peekel(1984), l3a. Pimae induplicate; chntering palrns with coarse fibrous leafbases;crownshaft lacking . -, Arenga microcarpa a German Catholic priest who resided on New l3b. Pinnae reduplicate; solitary or clustering palms with Ireland for many years. A truly enlightenedindi- non-fibrousleafbases; crownshaft present ...... 18 vidual, Peekel studied not only the botany of New I 4a. Staminate soft, with long, pointed, loosely closed Ireland, but also the language and cultuie of the ,mature when first exposed; terete in cross- people there. We are indebted to Peekel for the section...... rJ I 4b. Staminateflowers hard, bullet-shaped,maturing slowly indigenousnames from New Ireland and much of after exposure of the inflorescence; Slobed in the ethnobotanicalnotes included in the followins cross-section,at least when immature ...... 16 list. I5a. Small,clustering palms ...... Hydriastele kasesa ISb. Tall, solitarypalms; restricted to ManusI...... Kry ro Gronophyllum manusii rHE SpECTES I6a. Pinnae broad. wedge-shaped,apically "oru"*l .eed subteretein cross.section..... Ptychosperma hentyi la. Leaves palmate; small, solitary palms of the forest I6b, Pinnae lanceolate, apically oblique to concave; seed undergrowth ...... Licuala lauterbachii clearly SJobed in cross-section lb. Leaves pinnate or bipinnate; habits various ...... 2 17a. Fruits less than 2,5 cm long, with thin, fibrous endo- Leavesbipinnate; large solitary palms producing inflo- carp . .. .. Ptychospermagracile rescencesfrom the top downward I7b. Fruits more than 2.5 cm long, with thick, elaborate, . .. Caryota rurnphiana bony endocarp Ptychococcus lrraemerianus 2b. Leaves once pinnate . 3 I8a. Inflorescenceappearing as severalelongate mbrmched 5a. Fruit in a denie, globosehead; trunkless palms forming ":l:ll: "";';;:'ff,;,""., spikes emanating from a single leaf axil . ::i:l':::'::i:l ::::' r8b.r;i;;;.;;;"" u""""r,"i, ;:"; * ;;;;:"!:t':::t:: 3b. Fruit loosely arranged or in dense, elongate heads; T; l9a, Fruit massive, more than 15 cm long. Widely culti palms with distinct trunks and growing in more ter- vated, and growing wild along coasts Cocos nucifera restrial habitats 4 19b. Fruit lessthan l0 cm long ...... 20 4a. Stems weak, climbing by means of spiny leaves, or 20a. Fruit black; flowers and fruit crowded, slightly sunken by spiny whiplike extensionsfrom the leaves or leaf in the thick rachillae Cyrtostachys peekelii axils...... 5 20b. Fruit and flowersnot as above ...... 21 4b. Stems erect, not climbing, and mostly not spiny . . 6 2Ia. Fruit grayish with conspicuous lighter ribs; pinnae 5a. Leaveswith irregularly placed,broadly lanceolateleaf. pendulous ..... Gulubiacostata lets, and extended into an elongate, spiny cirrus . . 2lb. Fruitorangetored...... 22 ... Calamus hollrungii 22a. Ftduitsmall, spherical ..... Heterospatheparuiflora 5b. Leaveswith regularly placed,narrowly lanceolateleaf. lets, with whiplike flagella attached to the leaf sheaths "o * ...Calamus ralumensis 1':l::i:::: :li::::: i Iiii::,;ir,,;;ipp,o,a 6a. Pinnae irresularly multiribbed 7 6b. Pinnae r"g,ilu.ly'a.ranged, each with a single prom- inentrib ...... I Actinorhytis calapparia H. Wendl. & Drude- 7a. Palms stilt-rooted; pistillate flowers and fruit distrib- New lreland. uted along most of the length of the rachillae; fruit red...... Arecaguppyana This specieswas reported from the Namatanai 7b. Palms not stilt-rootedl pistillate flowers and fruit on region of New Ireland by Peekel, but not seen the lower parts of the .u"hiil"" only; fruit yellow-green elsewhere,despite being found in both New Guinea ...... I and the Solomon Islands. Moore (1969b) reports 8a. Fruit in a densespike; small palms ofthe forest under- story ...... Areca macrocalyx that this speciesis widely cultivated as far west 8b. Fruit in a panicle; robust palms cultivated near villages as Malaya, so cultivation may be a factor in the anddisturbedsites ...... Areca catechu palm's distribution. Indigenous names (New Ire- t26 PRINCIPES [Vor.. 39

Iand)-Kuanu a: 1)ek ao elt e i P ala:H ak aheke, k ak' Calamus ralumensis Warb.-New Britain. Lamekot" kafa. ahiaka; This speciesis quite abundant in the lowland forest of New Britain. It is probably on New Ire- Areca catechu L.-Universally cultivated in land as well, but has not been collectedfrom there. the region. It resemblesthe widespreadC. uestitu.sBurr. of which Moore alsoidentified from the This is the widely used betel nut, and was most New Guinea, Islands. It is likely that the two are one certainly spread to the Bismarck Archipelago as Solomon the same species. Indigenous names (New a cultivated plant. Indigenous names (New Ire- and Ireland)-Kuama: kada; Pala: siribe, buso;"'. land)-Kuanua: buai; Pala: buei; Lamekot: aua iogel. note these names all appear to be slight variations Lamekot; on the widespreadPidgin English name (buai) fot Calyptrocallx sp.-New Britain. betel nut, suggestingrelatively recent introduction of the palm to New lreland. This one rather robust speciesresembles one collected in the Central Province of Papua New Areca macrocalyx Zipp. (as A. jobiensis n Guinea,but neither has been identified. The genus Peekel)-New Britain, New Ireland. is particularly in need of revision.

This species has spike-like infructescences Caryota rumphiana Mart.-New Britain, New derived from inflorescencesin which the pistillate Ireland, Manus, Long I. (Fig. 2). flowers are confined to the very base of each and common, rachilla. The speciesserves as a substitutefor the This speciesis both widespread Indonesia (Ternate) to the Phil- commercialbetel nut. Indigenousnames (New lre- occurring from northeastern , the land)-Kuanua: Kunxuli P ala: kurnulo; Lamekot: ippines, New Guinea, and the Solomon Islands makega, kega. Bismarck Archipelago, (Gudalcanal).Varieties have been described for geographicalvariants in some parts of the species guppyana Becc. (as A. noao-hibernica Areca range,includin g v aiety p ap uana Becc. from West Britain, New lreland. in Peekel)-New Irian. There is too little information available to This is a slender, stilt-rooted speciescommon determine the usefulnessof such varietal names' in lowJying coastalareas, with noncongestedfruits As its wood is valued for construction purposes similar to A. catechr.r..It is found only in the (pers. obs.),its spreadmay have been aided by Bismarck Archipelago and the Solomon Islands, human migration or trade. Indigenousnames (New nothing like it being found in New Guinea. It has Ireland)-Ku antra:gelep, kuraga; P ala:.galah; not been reported by Peekel as used for betel nut. Lamekot:gelat; (Manus)-Kali: Sahun. Indigenous names (New lreland)-Pala" misle, butno; Ugana: uapbua, bua. Clinostigma collegarum J. Dransf.-New lre- land. Calamus hollrungii Becc.-New Britain, New Clinostigma is widespreadin the Pacific, from Ireland, Manus. the Bonin and in the north to and Samoain the east. This specieswas found in The two rattans known from this region are New Ireland in 1975, growing on ridges in mon- quite different from one another, but each apPears tane forest at elevationsof about 1,350 m. It to belongto very widespreadspecies. C. hollrungii resemblesGulubia costata in general aspect, but Warb. has leaves with widely spaced,irregularly is distinguishedmost readily by its fruit with Iateral arranged, broadly lanceolate leaflets, and a ter- stigmatic residue. The speciesis also notable {or minal flagellum, while C. ralurnensi.shas closely the mass of distinctive stilt roots at the base of spaced,narrowly lanceolateleaflets and a cirrhus the trunk. arisingfrom the leaf sheathrather than a flagellum leaf tip. C. hollrungii is common from the Cocos nucafera L.-New Britain, New lreland, New Guinea, the Bismarck Archipel- throughout Manus, Long I. ago and the Solomon Islands. Indigenous names (New Ireland)-Kuanua: ntagu; Palaz kalaua, The widespread coconut palm is cultivated kalawa: Lamekot: ni. throughout the Bismarck Archipelago. Indigenous reesl ESSIG:PALMS OF BISMARCKARCHIPELAGO

2. Villagers from Matafuma Village on Long Island drive spikes of Caryota rumphiana "wood" into a log, during construction of a mskeshift outrigger canoe. 3. The bright red fruits oI Heterospathe paniflora are widely spaced on the branches of the inflorescence. This palm is known only from New Britain. 4. A villager from Sampantobil in the Nakanai Mountains of New Britain holds a leaf and of Ptychosperma hentyi, a rlnigue and beautiful species found only on this island. 39 I28 PRINCIPES lVoL. names (New Ireland)-Kuanta: la'mamas; Pala: Metroxyfon sagu Rottb. (M. laeue Marr., M. lamas; Lamekot: ni. rumphii (S/illd.) Mart.)-New Britain, New Ireland, Manus. Cyrtostachys peekeliahd Becc.-New lre- land. This is the common sago palm found from Malaya and the Philippines to New Guinea and Cyrtostachys extends from Thailand to the beyond. It is found in swampy areas throughout SolomonIslands, with the greatestnumber of spe- the larger islands of the Bismarck Archipelago, cies in New Guinea. Only the one speciesis found and on Guadalcanalas well, while M. salomonense in the Bismarck Archipelago, and so far only on is found also on Guadalcanaland elsewherein thd, New Ireland. One specieshas been found east- Solomons. Another species, C. bougainaillensis wardo on Bougainville (C. Kisu Beccari). Peekel has been describedfrom Bougainville. As it is an reports that the hard outer woodis usedfor planks. important economic plant throughout its range, Indigenousnames (New lreland)-Kuanual. a-iai Metroxylon sagu may also have been spread by Pala: a-ihul; Lamekot: lifur;Ugana: aariuan, iuin. humans.According to Moore (1969b) the genus Gronophyllum manusii Essig-Manus. is Melanesianin origin, rather than Asian or New Guinean.Indigenous names (New Ireland)-Pala: Found on Manus Island, but similar appearing 6ia (spiny form called bia-gargarat); Lamekot: palms have been noted from a distance on Long sasak. Island. Nypa frutic?hs Wurmb.-New Britain, New Gulubia costata (Becc.)Becc.-New Britain. Ireland. This widespreadspecies of New Guinea has so Nypa is found in brackish estuaries from Sri far been found in only a few spots in the Hoskins 'West Lanka and eastern India to the Solomon Islands, area, New Britain. Ryukyu Islands,and Australia. Peekelreports that Heterospathe parviflora Essig-New Britain Nypa fronds provide excellent material for atap "wild (Fie.3). matting, and is commonly called sagopalmo' " ' or sak-sak.' Indigenousnames-Kuanua: rnan- . Heterospathe is a widespreadgenus with many imuai Pala: bia-to, bia-ta; Lamekot: ulagai. Iocally endemic speciesin both New Guinea and the Solomon Islands. This is the only speciesknown Physokentia av:a H. E. Moore-New Britain. from the Bismarck Archipelago. The genus has aaia is related to speciesin the not been found in New Ireland and was unknown Physokentia and is found at relatively to Peekel. Solomonsand beyond, high elevations. Hydriastele kasesa (Lauterb.) Burr.-New Ireland, New Britain. Ptychococcuskraemerianus (Becc.) Burr.- New Britain, New lreland. This speciesresembles H. microspadix (Beec.) Burr. from northeastern New Guinea, but dimen- This specieswas describedfrom New Ireland, sions overall are smaller. Widespread in lowland but the type consists of only a few large fruits forests. Indigenous names (New Ireland)-Pala: with deeply grooved and heavily armored endo- kasesa;Lamekot: kasi;Ugana: oakase,kase. carps, and seedswith homogeneousendosperm. Recent collections from New Ireland and New & K. Schum. Licuala lauterbachii Damm. Britain, with matching fruits, are indistinguishable Britain, (Licuala peekelii Lauterb.)-New from P. elatusBecc.,a speciesabundant in north- BagabagI., New Ireland. eastern New Guinea. Furthermore, the species This is the only speciesof this widespreadand from the Solomon Islands also closely resemble diverse genusto be found in the Bismarck Archi- this one. Ptychococcusis valued by the indigenous pelago. It is common in eastern New Guinea and people for its hard wood, which is used for bows is found also throughout the Solomon Islands. In and spear heads. It may have also been spread New Britain it is the most common palm in the among the islands by humans, and in fact, dis- r:nderstory of lowland forests' Indigenous names persalof its large fruits by any other meansseems (New lreland)-Kuanua: luga, uban; Pala: unlikely. Thus it appears that there is but one ;n lowland salaho; Lamekot:,f.. widespread species of Ptychococcus r99sl ESSIG:PALMS OF BISMARCKARCHIPELAGO t29 eastern New Guinea, the Bismarck Archipelago, This specieswas described from New Ireland and the Solomon Islands. The name P. kraerner- from incomplete material. The leaves are clearly ianus has nomenclatural priority. of Ptychosperrna, possiblyP. gracile. The inflo- rescencefragment is unusual, however, as it has Ptychosperma gracile Labill.-New Britain, what appears to be an elongate main axis and New Ireland, Long I. mrmerous short rachillae. It resembles more a This specieshas elongateleaflets and ruminate, Heterospathe, leading me to suspectthat the type weakly groovedseeds, and is similar to P. elegans collectionwas mixed. Flowersand fruit were lack- of Australia, and P. salontonenseof the Solomon ing, and the specimen at Berlin was probably Islands. The group of solitary medium-tall palms destroyed, so it is not likely that this specimen with red fruit and ruminate endosperm has no will be further identified. obvious close relatives in New Guinea, and thus could be considered part of the ooextra-New Guinea" element of the palm flora. Acknowledgments Ptychosperma hentyi Essig-New Britain (Fig. I thank the staff of the Division of Botanv. 4). Office of Forests, Lae, , for their assistanceon my several trips to the Bis- This species is widespread and abundant on marck Archipelago, and the National Science New Britain. but has not been found elsewhere. Foundation and the National Geographic Society It has unusualbroadly cuneateleaflets with convex for financial support. tips, that gradually decrease in size toward the end of the strongly arched leaves. Its seedsare nearly terete at maturity (although clearly S-lobed Lnenerunr CIreo when immature) with ruminate endosperm. Esslc, F. B. 1977. The palm flora of New Guinea. Botany Bulletin. Papua New Guinea Office of Forests. ADDENDUM Hev, Attsretn. 1984. A guide to the of Papua New Guinea. IIL Iz (R. Johns and A. J. M. Hay, Elaeis guineensis Jacq., the African Oil eds). Palmae. Papua New Guinea Office of Forests. Palm, has recently become widely cultivated on Moonr, H. E., Jn. 1969a. New palms from the Pacific,II. New Britain. There is no indication, however that Principes13: 67-76. it has escapedand become part of the flora. MooRE, H. E., JR. I969b. A preliminary analysis of the palm flora of the Solomon Islands. Phil. Trans. Roy. Soc. Ptychosperma novo-hibernica Becc. in 8255: 589-593. (English Lauterb., Beitrdge zur Flora von Papuasien.IV. PEEKEL,P. G. translation by E. E. Henty). 1984. Flora of the Bismarck Archipelago for Naturalists. Office Engl. Bot. Jahrb.52:29. I9I4. (Holotype:Kroe- of Forests, Division of Botany, Lae, Papua New Guinea. nler s.n.) I909, New Ireland (B)). Biographical notes by H. Sleumer.

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