r9821 ESSIG:GULUBIA

Principes, 26(4), 1982, pp. I59-I?3

A Synopsisof the GenusGulubia

FRnoeRrcrB. Essrc Department ofBiology, flniuersity ofSouth Flnrida, Tampa, FL 3362O

Gulubia is a genus of nine known tandrous genera, petals of the pistillate species in the Areca alliance of palms. flowers have long, tapered tips ihat are The genus is distributed from the Moluc- closed over the stigma at the time the cas in Indonesia to the Palau Islands, New inflorescence opens. Staminate anthesis is Guinea, the Solomon Islands. the New first (though not immediate), with pistillate Hebrides, Fiji, and Australia. All are soli- anthesis following sometime later (details tary, moderate, elegant palms with prom- of timing have not been observed). inent crownshafts and clean tmnks. Two The present account summarizes what species have leaves with essentially straight is known about Gulubia, but is clearlv rachises and pendulous pinnae (Figs. 1- preliminary in nature. Specimens avail- 2), while the remaining species have able for most species are meager and there strongly arched leaves with erect pinnae is much that is not known. A large part (Figs- 3-8). These palms are eminently of the range of the genus has not been suitable for cultivation in the tropics, but adequately explored for palms, so new have not yet been widely planted. They species may come to light in the future, seem to have little economic use, but in just ag two new speciesare described here. at least are used for floor- It is hoped that this paper will stimulate boards and sometimes for siding on build- the further exploration that is needed. ings. The genus is closely related to Grono- Fruit phyllum, Hyd.riastele, and Nengella, with Structurein Gulubia which it forms a natural subunit of the The structure of the pericarp has been Areca alfiance (Essig and Young 1979). found to be of considerabletax;nomic sis- From Hydriastele, Gulubia is separated nificancein the arecoidpalrns (Essig 1928, only by its emergent, solitary rathlr than Essigand Young l97t), so a spe;ial sec- caespitose habit. Inflorescence and flowers tion on the fruit of Gulubia is included are essentially indistinguishable. However here. The known speciesof Gulubia share it seems useful to provide an account of several basic features: a prominent pali- these distinctive emergent palms, even sade layer derived from the locular epi- though their generic status may hav'e to dermis, a series of fibrovascular bundies be reviewed in the future. From Grono- of varioussizes and shapes,a prominent, phyllum and, Nengella, Gulubia and dense zone of tanniniferous parenchyma, Hydriastele are disiinguished by a pro- and a subepidermal zone of compr;ssed togynous rather than protandrous mode of parenchyma. There are no sclereids or flowering and related morphology. In the crystalsin the outer pericarp, featuresthat p.rotogynous genera, pistillate petals are are commonin other arecoidDalms. short and the stigma is exposed it the time Representativesof all nine speciesof the inflorescence - opens. Pistillate anthesis Gulubia were examined using standard is immediate and staminate anthesis fol- histologicaltechniques (see Es-siglg7B). lows in 24 hours (Essig l9Z3). In the pro- Eight speciesare illustrated. Preparations PRINCIPES lVoL. 26

L. Gulubia moluecana,cultivated at Bogor (from Beccari,Am. Jard. Bot. Buitenzorg2, plate 7. 1885)' 2. Gulubia costdta' in lowland forest near the '

from dried material of Gulubia pct'lauensis tion of bundles (Fig. 9), which may be taken as. a more generalized condition were not adequate for drawing, but some 'n information was obtained, which will be probably ancestral to that G. costata. summarized below. 2. Gulubia longispatha can be char- Variation in fruit structure.is striking acterized by the presence of a distinct at every level, including the subspecific, series of fibrous bundles located outside of so detailed conclusions about the fruit the tanniniferous zone and apparently not structure in particular species cannol be connected with the fibrovascular bundles made based on limited samples. This is to the inside of the tanniniferous zone (Figs' iflustraled in the specimens of Culubia t 2-r4). longispa.tha, taken from three widely 3. Gulubia ualida appears to be most separated populations (Figs. l2-14). The similar to G. longispatha, bur lacks the following tentative remarks do seem to be outer series of fibrous bundles' and has a warranted however: broader zone of non-tanniniferous paren- I. Gulubia costdta clearly stands out chyma between the fibrovascular bundles because of the presence of very large and the tanniniferous zone (Fig. ll). fibrovascular bundles, alternating with 4. Gulubia hombronii and G. cYlin- much smaller ones, that give the fruit a drocarpa appear to be very similar in their ribbed appearance (Fig. l0). Gulubia fruit structure (Figs. 15 and l8). nith the moluccana clearly lacks this differentia- latter species apparently having smaller, l 9821 ESSIG:GULUBIA

3. Gulubia long^ispatha, growing +' culubia on the slopes of Mt. Suckling. rongispatha'type prant oi i. u^*li,t "irBella vista,c""oJn"u*"l photoby L. H. Brass, s. Gutuu toto nffi;t":i",T"T:'#,fiiT:XT;" FriedaRiver . 6. Culubia ualida, from the Torricelli Mourtalr, 1typ"il;j. PRINCIPES lVoL. 26

7. Gulubiahombronii, Santa Ysabel Island. Photo 8. Gulubia microcarpa,Viti Levu, Fiji Islands. by H. E. Moore,Jr. (courtesyL. H. BaileyHorto- Photoby H. E. Moore,Jr. (courtesyL. H. Bailey rium). ' Hortorium). more numerous and crowded fibrovascu- lar bundles. This difference may not be pericarp similar Io G. microcarpa, btrl reliable however as it is based on only one with a much thicker locular epidermis, well specimen from each species. developed as a palisade layer. There is no 5. Gulubia macrospadix is distinctive, sisn of fibrous bundles external to the tan- nol only in having ruminate endosperm. niniferous zone (not illustrated). but also in having a series of fibrous bun- In summary, it appears that each dles in mid-periclrp, which apiears to be species of Gulubia possessesdistinctive distinct from the fibrovascular system' but features in the pericarp, and isolated fruits not"as widely separated as in G. longis- might be identifiable. Subspecific variation patha. The fibrous bundles interrupt the appears to be significant, in at least one tanniniferous zone rather than being specieshowever, and this may confuse the external to it (Fig. 16). Also, the locular picture. Ultimately, pericarp structure may epidermis is wavy, following the uneven prove to be a sensitive marker for affini- contours of the seed. ties at the subspecific and population level. 6. Gulubia microcarpa has the thin- nest pericarp of all species, with markedly TaxonomicTreatment flattened fibrovascular bundles, and a locular epidermis only slightly modified in Gulubia Beccari in Ann. Jard. Bot. Bui- the direction of a palisade layer (Fig. l7). tenzorg2: l2B, l3l. 1885; Beccariin 7 . Gulubia palauensis, viewed in dried Martelli, Nuov. Giorn. Bot. Ital. ll, 42: material from Tuyama s.n., has a thin 84. 1935; Beccariand Pichi-Sermolli t9821 ESSIG:GULUBIA

o ta I O a a O

12

10

r3

Epidermls

Tshniniferous cetle

Fibroua bundlea Fibrovascular bundl6s -1J Locular, €pidermis

Figures 9-14. Diagrams of typical segmentsof the pericarp in cross-section. 9. Gulubia moluccana, from Beguin 2098. I0. Gulubia costata, from Essig s.n. (Brahman River area, Madany Province). lI. Gulubia aalida, ftom EssigLAE 55099. 12. Gulubia longispatha, from Essig LAE 55231(Mt' Suckling, Milne Bay Province). 13. Gulubia longispatha from Brass 5457 (Central Province). 14. Gulubia longispatho, from DssJgLAE ($test Province)' PRINCIPES lVoL. 26

l5

Figures l5-I8. Diagrams of typical segmentsof the pericarp in cross-section. t5. Gulubia hombronii, from Moore & Witmore 9296. 16. Gulubia macrospadix, from Moore & Whitmore 9305. 17. Gulubia microcarpa, from Moore & Phillips 1O543. 18. Gulubia cylindrocarpa, from Raynal 16256-

in Webbia ll: 40. 1955; Moore & scars; crownshaft well-developed; leaves Fosbergin GentesHerb. B: 455. 1956; reduplicately pinnate; sheath tubular, Moorein CentesHerb. 9: 263. 1963: elongate, cylindrical; petiole short, almost Moorein PrincipesI0: BB. 1966. Type flat or concave adaxially, convex abaxi- species:Gulubia moluccana (Beccari) ally; rachis straight or moderately to Beccari (seeBeccari & Pichi-Sermolli, strongly arcuate, pinnae regularly op. cit.). arranged, pendulous or horizontal to Kentia Blume in Bull. Sci. Phys. Nat. ascending, sometimes upper Ieaflets Neerlande l: 64. lB3B (i4 part, see recurved and drooping, linear-lanceolate, Moore in Gentes Herbarum 9: 264. unicostate, acute or notched at the apex, 1963); Beccari,Malesia 1: 36. 1877. sometimes with prominent ramenta on the Type: Kentia moluccana Beccari lower surface; leaf axis densely brownJep- Gulubiopsis Beccari in Bot. Jahrb. Syst. idote-tomentose to minutely brown-dotted. 59: lI. 1924; Beccari & Pichi-Ser- Inflorescence infrafoliar, solitary at each molli in Webbia 1l: 40. 1955. Type: node, but often several in different stages C. palauensisBeccari of flowering and fruiting at one time. pan- Paragulubia Burret in Notizbl. Bot. Gart. iculate, branching to l-3 orders, pedun- Mus. Berlin-Dahleml3: 84. 1936; cle short, often becoming bulbous in fruit, Beccari & PichiSermolli in Webbia I l: prophyll compressed, broadly oblanceo- 46. 1955. Type: P. macrospadixBtr- late, completely closed around the bud, ret. rounded at the tip, second peduncular Tall, solitary, unarmed, monoecious bract smaller, incomplete, triangular to palms; trunks smooth, with annular leaf Ianceolate, or rudimentary in a horizontal r9821 ESSIG:GULUBIA 165 scar, rameal bracts rudimentary, axes gla- 3. Pinnae more than 50 on each side of the brous to slightly glaucous, whitish, rach- rachis. 4- Fruit ca. I0 mm long, with very thin illae elongate, straight or somewhat flex- pericarp; incompletely known species uous. Flowers white to rose-pink in from Fiji. G. microcarpa. decussatetriads for most of the length of 4. Fruit usually larger with thicker peri- the rachillae. Staminate flowers one on carp; species from New Guinea, New flower, Hebrides and Solomon Islands. either side of the central pistillate 5. Stamens 6. ..-...-...-...... G. cylindrocarpa. asymmetrical, inserted at an angle to the 5. Stamens9-24. rachilla, with the outward-facing petal . 6. Pinnae stiff, not drooping at tip; substantially larger than the other two, fruit dark red. .,,-.,...... -Gl ualida. sepals 3, short, gibbous, pointed, briefly 6. Pinnae lax, drooping at tips; fruit bright red...... G. Iongispatha. united at the base, petals 3, 4-5 times 3. Pimae 30-40 on each side of the rachis, longer than the sepals, valvate, broadly 7. Flowers cream-white; staminate flowers lanceolate, tapering to a fine point; not 3.5-4 mm long with 6 stamens; fruit tightly closedin bud, stamens6-24, erect, 9 X 4 m. Palau Islmds. .- G- palauense. staminateflow- to somewhat 7. Flowerspink to rose-red; basifixed, somewhat shorter ers 8-lI mm long. longer than the petals, pistillode lacking 8. Fruit dull red; endosperm homoge- or rudimentary and representedby 3 small neous: stamens l0-12. .-...... ,..-...,...,..,,.- papillae. Pistillate flowers globose, much C. hombronii. smaller than the staminate flowers, sepals 8. Fruit bright crimson; endosperm ruminatel stamens6-9. broadly rounded, imbricate, gibbous, pet- G. macrospadix als similar but briefly mucronate and not gibbous, sometimes ciliate, parting only t. Gulubia moluccana (Beccari)Bec- slightly at anthesis, staminodes usually cari in Ann. Jard. Bot. Buitenzorg2: 3(l-6), dentiform, oppositethe inner petal 131.1885. only, pistil conic-globose,stigmas 3, short, Kentia m.oluccanaBeccari, Malesia I: 35. sessile,exposed and receptiye when the IB77; H. A. Wendland in Kerchove, inflorescencebracts open. Fruit ellipsoid Les Palmiers: 248. L878. Type: Bet- stig- to globose, symmetrical with apical cari s.n. lB74 (Il sheetsfiled under matic residue, bright to dull red or blue- accessionnumbers 11150, 11151) gray with pale stripes, pericarp thin with (holotypeFI, photosat BH). a prominent tanniniferous layer in mid- pericarp, and straight, little-branched Solitary palm 20-30 m high (to 55 m fibrovascular bundles running longitudi- high f.de de Vogel), stem to 30 cm in nally, these forming prominent rihs in one diameter.Leaf sheathto 95 cm long, blade species;seed ellipsoid-ovoid, embryo basal, 250-300 cm long, upper sheath,petiole endosperm homogeneous or (in one and rachis finely brown-lepidote, pinnae species)ruminate. pendulous, ca. 45 on each side (from photo, Fig. 2), to 105 cm longo 3.3 cm Key to the Speciesof Gulubia wide, lower surface with prominent, basi- fixed, pale brown ramenta along midrib at I. ?innae pendulouson slightly bowed rachis. base. Inflorescencebranching to 2 orders, X 4 mm (dry), not conspicuously 2. Fruit 9 cm long, ca. 2 mm thick, ribbed; leaf blade of about 45 pinnae on rachillaeto 39 each side. Moluccas. .-..-...--..-...--..G. moluccana. bearing up to 240 triads; staminate flow- 2. Fruit 7-8X4 m (dry), conspicuously ers cream-colored,ca. 6 mm long, sta- ribbed; leaf blade of usually 65-77 pinnae mens 6; pistillate flowers reddish at base, on each side. Widespread in New Guinea, cream-coloredat tip; fruit 7-9 X ca. 4 Aru Islands, northeastern Australia.,.,,,..-.."--- .-..-...G.costata mm, Iacking prominent ribs, color l. Pinnae ascending on strongly arcuate rachis. unknown. r66 PRINCIPES [VoL.26

Distribution: Moluccas-Halmahera, mann 8228 (holotype B, believed Ternate, Bacan, on hillsidesup to 1,200 destroved). m altitude. Gulubia costatd var. nl,inor Beccari in Local name: ifu (Ternate language). Ann. Jard. Bot. Buitenzorg 2: 135. SPECIMENSEXAMINED: Ir,rnoNnsI.{. 1885. Type: Beccari s.z. 1875 (filed "sul Ternate: abundant Picco di Ternate under accessionnumber lll53 in FI) ad Aequi Conora," alt. 600-I,200 m, (holotypeFI). November 1874, Beccarl s.z. (accession Gulubia costata var. pisifunnis Beccari numbers11150, 11151)(FI, type); Hal- op. eit., p. 136. Type: Beccari s.n. mahera:Loci Tabaroi,alt. ca. 800 m, 29 lB78 (filed under accessionnumbers JuIy 1922, Beguin 2098 (K); Gunung 11149 in FI) (holotypeFI). Jailolo,rather denseprimary forest, 20 m Gulubia costata var. gracilior Burret in high with little undergrowth, steep hillside Notizbl. Bot.. Gart. Berlin-Dahleml3: with deep, loose,porous, black, volcanic Bl. 1936. Type: Bross 5BBZ(holotype soil; solitary, emergentpalm from 650- B, believeddestroyed; istotypes NY, A). 900 m alt., 12 October I974, de Vogel Tall, moderatepalm to 20 m or more 3386 (K); Bacan Island: Gunung Sibela, in height; stem 20-30 cm in diameter; near Waiaua, alt. 1,050 m, rather dense leaf sheath75-150 cm long, green,tinged primary forest 35 m high, with little with lilac, glabrous to finely light-brown- undergrowth, steephillside, rather dry, on tomentosenear the tip, petiole35-60 cm shallow, clayey soil with stones, bedrock long, blade 220-420 cm long, petiole and gray schists,26 October I974, de Vogel finely white-woolly and brown-lepi- s681(K). rachis dote above and below, pinnae pendulous, The several recent specimensof de 43-77 on each side,97-127 cm long, Vogel have improvedour understanding 5.5 cm wide with the apex deeply bifid of this species,which in general is very (or hoodedin life), upper surface glabrous, similar to Culubia costata. The principal lower surface minutely brown-dotted and differenceis in the frurt, Gulubia moluc- with large, whitish, basifixedramenta along canalackirl'g the very prominent seriesof the lower third of the midrib. Inflores- fibrovascularbundles that give the fruit of cence branchingto 2-3 orders, 70-125 G. costata its distinctive ribbed appear- cm long, the upper peduncularand rameal ance. The inflorescenceof G. moluccana bracts rudimentary, triangularo to 6 mm appearssmaller, from de Vogel's speci- long or represented only by horizontal mensowith rachillae only half the length scars; rachillae straight or sometimes of thosein G. costata. It is lik6ly that G. somewhat flexuous, to 62 cm long, 2-3 molucca,narepresents a relictual, more mm thick, bearingup to 280 triads. Sta- generalizedspecies directly ancestralto the minate flowers cream-colored, soapy- widespreadG. costata. scented,6-7 rnlr:'long, stamens6, shorter than the petals, pistillate flowers 1.5 mm 2. Gulubia costata (Beccari)Beccari in high, flushed with pink; fruit 7-9 X 4.5- Ann. Bot. Buitenzore 2: I34. Jard. 5 mm, blue-gray with whitish stripes over IBB5. prominent fibrous ribs; endospermhomo- Kentia costatcl Beccari, Malesia l: 36. geneous. I877. Type: Beccari s.n. lB73 (filed Distribution: Ubiquitousin lowland New under accessionnumber 11152 in FI) Guinea, also found in the Aru Islands, Bis- (holotypeFI, photosat BH). marck Archipelago, and northeastern Gulubia ffinis Beccari in Bot. Jahrb. Queensland. Syst. 58: 444. 1923. Type: Leder- SPECIMENS EXAMINED: INloNnsre. r9821 ESSIG:GULUBIA 167

Aru Islands:Vokan, April 1873, Beccari BH); Port Moresby Subprovince,along the s.n. (11152 in FI) (type number), s.n. SubitanaRoad, near the Musgrave River, (11152A-n (FI, photosat BH); Ansus, disturbed rain forest in hilly country, alt. April 1875, Beccaii s.n. (11153) (type 1,800 ft, 23 February1972, EssigLAE of G. costata var. minor (FI); Java (cul- 55178 (LAE, BH); Western Province: tivated): Buitenzorg (Bogor) Botanical near Wuroio along the , Carden,May 1878, Beccaris.n. (11149) common, a conspicuousfeature of the riv- (type of G. costatavar. pisifurmis),s.n. erbank forest, January-March 1934, (11149) (FI); Irian Jaya:Aria, near Uta, Brass 5881 (type of G. costatavar. gra- alt. 4 m, 28 June 194I, Aet 386 (K); cilior,holotype B, believeddestroyed; iso- Pepue New GuINne. W'est Sepik Prov- types NY, A); commonamong river banks, ince: Aitape Subprovince,near Sumo Vil- Palmer River, 2 miles below junction with lage, Rhinbrum River, in tall forests on Black River, July 1936, Brass 7245 (A, river flats, alt. ca. 50 ft, 5 July 1961, BH); occasionalin rain forest near Lake Darbyshire & Hoogland B0BZ (CANB, Daviumbu,August 1936, Brass 7951 (A, BH); near Walwali Village, along Pieni BH); West New Britain Province: Hoskins River, in forest, alt. 100 ft,20 June Subprovince, cutover rain forest at the 1961, Darbyshire & Hoogland 7971 foot of Mt. Otto, 26 April 1972, Essig (CANB, BH); East Sepik Province: allu- LAE 55213 (LAE, BH). vial forest along a creek near Mt. Hun- Gulubia cost&ta is one of the most stein, 200 m, July-August 1912, Leder- common and abundant palms in lowland mann 8228 (type of G. afi.nis, B, believed New Cuinea.The speciesoccurs on mesic, destroyed;along Frieda River, a few miles well-drainedsoils in hilly terrain as well as downstreamfrom Carpentaria Exploration in swampy or seasonally flooded situa- Pty. airstrip, swampy forest, 27 April tions, Gulubia ffinis was describedfrom 1978, Essig & Young LAE 74053 (LAE, a specimen collected in the Sepik River 'Wewak BH, USF); Subprovince,2 miles Basin in an alluvial forest near Mt. Hun- west of But Village, alt. 20 ft, disturbed stein. It differed from Gulubia costatci lowland forest, 12 January 1972, Essig supposedlyin that the petals of the sta- LAE 55129 (LAE, BH); Angoram Sub- minate flowers were drawn out into lone province, near Kabriman Village, along bristle-liketips. Fruil were lacking from the Blackwater River. in a sago swamp. the specimen, but in other important alt. 30 m, 29 October 1972, Leach NGF respects,Beccari's description conforms 34312 (LAE, BH); Morobe Province: to Gulubia costata. The habitat, overall Kaiapit Subprovince,base of KassarilPass, dimensions,nature of the foliage, number gallery forest along stream in broad, open of stamens, size of the flowers, plus the ravine, alt. 600 ft, 3 June I97I, Stone fact that Gulubia costata is abundantin 10249 LAE 53549 (LAE, BH); Lae Sub- the upper Sepik Basin, all suggestthat the province, wet lowland forest along Mark- unusual shape of petals in G. affinis rep- ham River on Lae-Bulolo Road, 7 March resents a minor variation in the broadly 1964, Moore 9273 (LAE, BH); Milne Bay distributed species. The several varieties Province: Raba Raba Subprovince, be- that were describedalso appearto be based tween Kwagira and Moi Biri, coastal rain on minor variationsof no laxonomicsie- forest,alt. 50 ft, 7 July 1972, EssigLAE nificance.GuLubio costoto var. minir, 55521(LAE, BH); CentralProvince: road collected from the Aru Islands, not far from Mori River to Yanu Village, ca. 15 from the type locality of the species,has km N.E. of Cape Rodney, rain forest on somewhat smaller overall dimensions, as gently.undulatingterrain, alt. ca. 30 m, well as smallerfruit. G. costata var. grcl- 5 September1969, Pullen 8218 (CANB, cilior was basedon a very similar speci- I68 PRINCIPES [Vor. 26 men from western Papua. In the collec- staminodes;fruit bright red, subglobose, tion notes for the latter specimen, Brass 7-13 X 4-9 mm when dry; endosperm indicated that the individual was extremely homogeneous. tall. Leaves, inflorescences and fruit fre- Distribution: Widespread in mountain- quently become smaller as palms get taller. ous regionsof ,between G. costata var. pisifornlis was based on (?I97) 600 and 1,450 m elevation,often a specimen cultivated at Bogor, which had in small, isolated populations on steep fruit somewhat more globose than the typ- ridges. ical form. There is no evidence that this SPECIMENS EXAMINED: PEPUE corresponds to a naturally occurring vari- New GurNae. West Sepik Province: on ation worthy of taxonomic status. steep slopesof mountains south of Frieda River, Carpentaria Exploration Company 3. Gulubia longispatha Beccariin Bot. helicopterpad # K-27, alt. I,000 m, I Jahrb.Syst. 52: 25. 1914. Type: May 1978, Essig& YoungLAE 74081, Schultze323 (holotypeB, believed 74083 (BH, LAE, USF); EastSepik Prov- destroyed). ince: Ettapenberg, alt. 850 m, Leder- ffLann 9133 (8, holotype of Gulubia Gulubia crenata Beccari in Bot. Jahrb. obscura, photos only seen at BH); Hun- Syst. 58: 445. 1923. Type: Leder- stein Mtns. alt. 1,050 m, Ledermann mann 8449 (holotype B, believed 8449 (8, holotype of Gulubia crenata, destroyed). photosonly seenat BH); Sepik River, alt. Gulubia obscuraBeccari, op. cit., p. 447. 197 m (location dubious), November Type: Lederrnann 9133 (holotype B, 1910, Schuhze 323 (8, holotype of believeddestroyed). Gulubia photosonly seenat BH); Gulubia brassii Burret in Notizbl. Bot. ffinis, tr{orobe Province: Mountains above Mo Gart. Berlin-DahlemI2: 336. 1935. River, 5 hours walk from Ana Village, 29 Type: Brass5457 (holotypeB, believed January 1972, Essig LAE 55166 (BH, destroyed;isotypes A, NY). LAE); Milne Bay Province; on Castan- Tall slender palms to 24 m or more in opsis dominated ridge, junction of Ugat height;stem l0-25 cm in diameter;leaves and Mayu Rivers, near Mayu I camp (Mt. I6-19 in a crown, strongly arcuate, SucklingExpedition), alt. 700 m, 15 July sheat\ 95 120 cm long, petiole L2-20 I972, EssigLAE 55231(LAE,BH), same cm long, bladeca. 240-250 cm long, pet- locality, 17 July 1972, Streimann NGF iole and rachis thickly brownJepidote 28921 (LAE, BH); Central Province: Bella aboveand below;pinnae erect: but droop- Vista, common, sporadicin forests, sur- ing at the tips, 50 69 on each side of the viving on cleared land, alt. 1,450 m, rachis,65-95 cm long, 2.I-4 cm wide, November 1933, Brass 5457 (A', NY, deeply bifid at the apex, with the upper isotypesof Gulubia brassii). INoonnsIe. margin much prolonged, upper surface Irian Jaya: plentiful in mossyforest, 4 km minutely and sparsely dotted, the lower southwest of Bernhard Camp, Idenburg surfacethe sameand with severalto many River, alt. 900 m, March 1939, Brass large, pale ramenta on the midrib (lacking t3099 (A). in Mt. Suckling population). Inflorescence Having carefully examined the cited 60-90 cm long, branching to 2 orders, specimensoin comparison with the type axes and flowers white, glabrous;rachillae descriptionsfor Gulubia longispatha, G. ca. 50-60 cm long, l-4 mm wide,bear- crenat&.G. obscuraand G. brossii,I have ine 80-185 triads.Staminate flowers l0- reached the conclusion that they all rep- IB *- long, with 9-24 stamens.Pistil- resent elementsof a single, variable, and Iate flowers 2-3.5 mm hieh, with 2-3 widespread species. The examination of r9821 ESSIG:GULUBIA 169

the pericarp anatomy of several of the from the pericarp, it became evident that specimens, representing widespread pop- Gulubia brassii could no lonser be main- ulations, reinforced this conclusion. The tained. specimens examined shared a unique fea- ture in the genus, namely a distinct series 4. Gulubia valida Essigsp. nov. of fibrous bundles in the outer pericarp, G. longispathae affinis sed robustior,foliis similar to that found in some species of rigidioribus,pinnis erectis, apicibus the closely related genus Gronophyllum. non pendulis,staminibus 12, fructu atro- As far as can be determined, the fruit in sanguineo vice pallide-rubro this species are all bright red at maturity. differt. Typus: Papua New Guinea, Essig LAE Leaves are characteristically strongly 55099 (holotypusBH; isotypi A, BRI, arcuate, with pinnae ascending but soft CANB, K, LAE). and drooping at the tips. The habitat pref- erence of mountainous terrain for this Solitarypalm, with stems15 20 cm in species contrasts with the lowland habitat diameter; leaves about 22 rn a crown, of Gulubia costata. strongly arcuate, with pinnae ascending In this regard, however, the locality and rigid, not droopingat the tips, sheath given for the type collection of Gulubio ll0 cm long, petiole 40 cm long, blade longispatha is troublesome and possibly 210 cm long, petiole and rachis brown- an error. The locality is given vaguely as lepidoteabove and below, "the pinnaeabout 57 Sepik River, altitude I97 m, Novem- on each side,to B0 cm long,2.2 cm wide, ber 1910." At 200 m, one is still in low- glabrousabove, brown-dotted below, lack- land alluvial forest, the wrong habitat for ing ramenta.Inflorescence branching to 2 this species as currently understood. It is orders, with 6-7 secondaryaxes, these recorded, however (Flora Malesiana l: white, glabrous;upper peduncularbract 478. 1950), that during November (Nov. present,20 cm long, narrow, triangular; 2-I3) Schultze made a side trip to the rachillae48 cm long with ca. 134 triads. mountains south of the Sepik, ascending Staminate flowers white, 17 mm long and to Peripetus Peak (alt. I,492 m), which I 4.5 -6.5 mm wide,slamens 12. pisrillode have not located, but which apparently is lacking;pistiliate flowers white, 3 mm high. in the vicinity of the Leonard Schultze staminodes3, dentiform; fruit 1l X 7 mm, River, 1Q-15 miles east of the Frieda to purple; endospermhomoge- River. This, then would be very close to *::1.:"t where I collected the species and observed Distribution: Elevationsaround 1,000 it in great abundance. It is possible and m in the Torricelli Mountains of north likely that the type of the species was col- central New Guinea. lected on this side trip at an altitude con- SPECIMENS EXAMINED: Pepue siderably above 197 m. Nnw GuINne.West SepikProvince: Lumi The populations in the Sepik Basin are Subprovince,Torricelli Mtns., near the homogeneous with respect to stamen village of Fatima, besidethe road running number (9), but those in the southern part eastwardfrom Lumi, alt. ca. 3,000 ft, of the range are more variable. Gutibio 26 NovemberI97I, Essis LAE 55099 brassii has 20-24 stamens, which at first (holotypeBH; isotypesA. BRf. CANB, K, prompted me [o maintain it as a separate LAE). species.Other populationsbridge rhe gap, Gulubia ualida appears to be most however. The specimen from the Morobe closely related to Gulubia longispatha, Province has 12 stamens, and the speci- but there are clear differences in the mens from the Mt. Suckline area have 18. foliage and fruit. The leaf rachis is not as With the addition of the anatomical data strongly arcuate as in G. longispatha and 170 PRINCIPES lVoL. 26 the pinnae are stiff and erect, not droop- brown-dotted,pinnae ca. 30 on each side, ing at the tips as in that species.The epi- 82-100 cm long, 2.5-3.5 cm wide, tiP thet ualida refers to the strength and very briefly praemorse or notched, lower robustnessof the foliage. The fruit lack surface glaucous,with ramenta lacking or the series of fibrous bundles in the outer few and inconspicuousat the base of the pericarpcharacteristic of G. Longispatha, midrib. Inflorescence,40-90 cm long, but otherwisethe pericarpstructure is very branchingto 2 orderswith 7-ll primary similar (Fig. ll). The fruit alsoripen to a branches and Il-24 rachillae, upper dark red color, as opposedto the bright peduncular and rameal bracts lacking; red of the neighboringspecies. In addi- rachillae25-36 cm long, ca. 2 mm wide, tion, G. ualida has staminate flowers with glabrous,bearing ca. 150 triads. Flowers 12 stamenswhile the nearest populations red to rose in color. Staminate flowers 8 (all those in the Sepik Basin) of G. longis' mm long, 3 mm wide, with 6-9 stamens. patha have staminate flowers with 9 sta- Pistillate flowers globose-pyramidal,3.5 mens. mm high, with 3 dentiform staminodes. The new speciesmight be confusedwith Fruit 12-16 X 8 mm, bright crimson,seed a speciesof Gronophyllum (G. cf. may' with ruminate endosperm. rli) that occurs in the Torricelli Mtns. at Distribution: Bougainville and Santa somewhat higher elevations (Darbyshire Ysabel in the Solomon Islands. 464 at CANB, LAE), and which appar- Local names: niniu (Kwara'ae lan- ently has a similar overall appearance.The guage), kuritu (Botgainville). genericdistinction (of longer, valvate pet- SPECIMENS EXAMINED: PEPUE als in the pistillate flowers) is clear, how- Nrw GuINne. Bougainville Province: ever. Also, the Gronophyllum has fiewer Kugumara, Buin, 28 Muy 1930, pinnaeper leaf (38 per sideas opposedto I(ajewski 1787 (holotype B destroyed, 57 per sidein the Gulubia), and the pin- isotypeA); SorouoN IsreNos. SantaYsa- nae of the Gronophyllur?rpossess numer- bel: Bogotu Peninsula, slopesof ridge on ous conspicuousramenta on the lower sur- mainland opposite Horara Village near face. Flower color is not known in the Tatamba,alt. 0-500 ft,22 March 1964, Gronophyllurn, and the fruit color was Moore & Whitmore 9305 (BH, BSIP); recordedby Darbyshireas pale brown. Maringe Lagoon, near Tiratona Village, on a broad ridge, alt. 1,600 ft, 23 Octo- ber 1963, Whitmore BSIP 2325 (BSIP, 5. Gulubia macrospadix (Burret) H. K). E. Moorein Principes10: 88. 1966. This treatment is based essentially on that of Moore (1966), as I have not seen Burret in Paragulubia macrospadir any specimensother than those used in Berlin-Dahlem 13: Notizbl. Bot. Gart. his analysis.Gulubia macrospadix is dis' Kajeu.tski1787 (ho\o- 84. 1936. Type: tinguished from all other species in the isotypeA). type B, destroyed; genus by the ruminate endosperm of its Gulubia niniu H. E. Moore ex T. C. 'Whitmore, seed.This promptedBurret to erect a new Guide to the Forests of the genus for the species, a moYe which he Islands,1966, name only. Solomon believed was bolstered by the praemorse A tall, solitary palm to 20 m in height; character of the pinnae. Moore rejected stem ca. ll-12 cm in diameter;leaves the new genus,pointing out that ruminate ca. 2O-25 in a crown, arcuate,with pin- and homogeneousendosperm coexist in nae droopingat the tips, sheath ca. 60- many genera, and that other species of 90 cm long, petiole35-50 cm long, blade Gulubia alsohave slightlypraemorse tips. ca. 150.-195 cm long, petioleand rachis The fruit is also distinctive by virtue of a 1eB2) ESSIG:GULUBIA r7l

ring of fibrous bundles that interrupts the November 1965, Corner 2892 (K, BH); tanniniferous zone in mid-pericarp. Choiseul (easternmost):Ultrabasic hill on coast opposite Bembalama Island, forest with much casuarina and thick leaf litter 6. Gulubia hombronii Becc.in Webbia layer, 3 March 1964, Whitmore BSIP 3: 161. 1910. Type: Hornbrons.n., 4009 (K); Big Nggela Island: l83B-I840 (holotypeP). west of Haghela School, ridge top, alt. 350 ft, A tall, solitary palm, to 15-20 m in well-drained primary forest, 28 June height;stem l2-15(-28) cm in diameter: 1969, Gafui & collectors BSIP 15257 leavesl2-20 in a crown,strongly arcuate (K); Guadalcanal:on ridges above Tam- with pinnaeerect, sheath50-75(-90) cm balusu on Suta-Kiki River. alt. 2.000- long, petiole IB-20(-30) cm long, blade 3,000 ft, 3 November1965, Corner 195 100-140(-lB0) cm long; petiole and (K, BH). rachis glabrous, slightly glaucous above; Gulubia hombronii appears to have pinnae 36-38(-46) on each side, 45- somewhat smaller dimensions than G. 67(-75) cm long, 2-3.7 crn wide, with macrospadix, and both, in turn, are sig- apex briefly notched, lower surface with nificantly more diminutive than any of the many small, pale, basifixed ramenta on speciesin New Cuinea.The arcuateform the lower l0-12 cm of the midrib. Inflo- of the fronds suggestsa relationship with rescence40-55 cm long, simply branched Gulubia longispatha, but the rosy color or the I-2 lower branchesforked, with of the flowers is distinct from the cream- 7-9 rachillae; peduncular and rameal white of that species,and the pericarp bracts lacking; axes white, glabrous to structure is different(Fig. l5). finely brown-dotted;rachillae 36-50 cm long, 2.5-4 cm in diameter, bearing up 7. G0lubia cylindrocarp? Beccari in to l3B triads. Staminateflowers l1 mm Webbia 3: I56. 19I0. Lectotype: long, 4 mm wide, deep rose in bud, Harland. s.n., 21 1905 (FI). becomingrose-pink shadingto ivory at the June tips when expanded,fragrant, calyx white, Solitary palm to 27 m rn height; stems stamens10-12. Pistillateflowers pinkish. to 15 cm in diameter;Ieaves ca. 18 in a Fruit 14-17 X 6-6.5 mm, ripening dull crown, strongly arcuate, with pinnae erect; red; seed with homogeneousendosperm. sheath to 95 cm long, petiole to 25 cm Distribution: Widespread in the Solo- long, rachis ca. 2-2.5 m long; upper mon Islands. sheath, petiole and rachis lepidote with Local names: bulntari (Kwara'Je lan- reddish and white scales;pinnae ca. 55 guage),bombua (Longu language). on each side, to 95 cm long, to 3:3 cm SPECIMENS EXAMINED: Sorol,roN wide, apex notched, lower surface with a "durante Isrenos. St, Georges Island: il few whitish ramenta near the base on the viaggio dell'Astrolabe e della Zelee" midrib. Inflorescencebranchine to 2 (1838-1840), Hombrons.n. (holotype P, orders,the lowerprimary branchesdivided not seen,photo at BH of fragment at FI); into ca. 4 rachillae,with ca. 25-32'rach- Santa Ysabel Island: Cape Prieto, plentiful illae altogether; rachillae to 43 cm long, on crests of steep, scantily vegetated 2-2.5 mm wide, bearingup to ca. 190 mountain spurs, alt. 200 m, 14 January traids. Staminate flowers (mature?) 2.5 1933, Brass 3749 (A); BogotuPeninsula, mm long (Kajewski 6-ll), stamens6. Pis- on ultrabasic ridge due west of Tatamba, tillate flowers2.5-4 mm high, 2.25 mm alt. 0-500 ft, 19 March 1964, Moore broad, staminodes2-6. Fruit apparently & Whitmore 9296 (BH); poor casuarina yellowishat maturity, cylindrical to some- forest on ultrabasic soil, Tatamba Bay, 6 what ovoid, slightly curved, l2-I3 X 5 172 PRINCIPES [Vor.. 26

6 mm, cupuleca. 3.5 mm high; seedwith 25 rachillae, upper peduncular bract tri- homogeneousendosperm. angular to strap-shaped,3.8 to 8.5 cm Distribution: New Hebrides. long, axesglabrous, rachillae 30-32(-40) Local name: motoual (fide Harland). cm long, 1.5-2 mm thick, bearingabout SPECIMENSEXAMINED: Nnw Heb- 90 traids. Flowers white. Staminate flow- BRIDES.Vanua Lava: nr. Mt. Garigona, ers ca. 4 mm long, I mm wide, stamens alt. 1,000-1,500 ft, 2l June 1905, Har' 6, somewhat exserted beyond petals in land s.n. (FI lectotype, photo and frag- bud. Pistillateflowers globose-conic, 2-2.5 ments BH; isolectotypeK); Erromongo: mm high; fruit ellipsoid,9-I0 X 4.5-5.5 I I miles west of Ipota, with Agathis, Cal- mm, color not noted; seed with homoge- ophyllum and. Hernandia, 28-3I May neous endosperm. 1968, Bernard,i 13369 (K, BH); along Distribution: Palau Islands on lime- forestry route, km 12, dense forest, 9 stone. AugustL971, Raynal RSNH 16256 (K, Vernacular names: bugelangererals, BH); Malekula, South West Bay, east of sz6ric (Palau). 'Wentoua, along the ridge crests,alt. 250- SPECIMENS EXAMINED: PETEU 300 m; SorouoN IsLeNos. Santa Cruz Istlt,tos. Urukthapel Island, east end, Group, Vanikoro Islands: common in rain common on limestone ridge, less so on forest, alt. 800 m, I I November 1928, slopes,alt. 200 ft, 2 April 1950, Fosberg Kajewski611(K). 324391 (US, BH, BISH); Koror, in cal- This speciesseems to be most closely careoushills, 26 August 1939, Tuyama related to Gulubia hom.bronii. Stamen s.z. (fragmentsfrom TNS at BH); seealso number is different, and there may be a specimenscited by Moore & Fosberg differencein ripe fruit color, but the cylin- (res6). drical nature of the fruit doesnot seem to . This treatment is distilled from Moore hold up well when specimensother than and Fosberg (1956), augmented with the type are examined. The anatomy of reexamination of the cited specimens. the pericarp is most similar to that of There apparently have been no more Gulubia hombronii. recent collections, and in fact, the species was said to be seriously threatened at the B. Gulubia palauensis (Becc.)Moore time of Fosberg's visit. & Fosbergin Gentes Herb. 8: 455, The genus Gulubiopsis was established fig. 135. 1956. on the basis of the presence in the peri Gulubiopsis palauensis Becc. in Bot. carp of rigid, fusiform fibers, shorter than Jahrb.Syst. 59: Il, fig. 133. 1924. the entire length of the fruit. The fact that Holotype: Led.errnann14149 (8, Pre- the staminate flowers expand early in bud sumed destroyed). has also been noted as a generic distinc- A tall, solitary palm to 18 m in height; tion. Moore and Fosberg did not find these stem 15 cm in diameter; Ieaves appar- characters to be distinctive enough to war- ently arcuate with erect pinnae (as judged rant generic status, as similar features from the attachment of the pinnae to the could be found in other species of Gulu- rachis),sheath ca. 57 cm long, petioleca. bia. The species appears to be most sim- lB.5 cm long, blade about 100 cm long, tlar to Gulubia I'ongispatha, but with sig- sheath, petiole and rachis thickly brown nificantly smaller overall dimensions, lepidote;pinnae 35 on each side, 54-59 including small, staminate flowers with only cm long, 2.2-2.4 cm wide, aPex deePlY 6 stamens, and possible differences in bifid, lower surface with large, pale brown pericarp anatomy. Cursory examination of ramenta along the lower 5-10 cm of the dried fruit from Tuyama s.n. did not reveal midrib. Inflorescence52-60(-90) cm the short, fusiform fibers mentioned by long, branching to 3 orders, with about Beccari, however. r9821 ESSIG:GULUBIA t tJ

9. Gulubiamicrocarpa Essig, sp. [ov. mature fruit is. The speciesis distinctive G. cylind.rocarpae simllis sed fructu in its small fruit with very thin pericarp minore, ca. l0 X 3 mm, pericarpio and poorly developed palisade layer. It tenuissimo.Typus: Flji, Moore & Phil- appears to be most closely related to lips 10543 (holotypusBH). Gulubia cylindrocarpa. The specificepi- Tall, slender palm to 15 m or more in thet was suggestedby ProfessorMoore, height; stem 28 cm in diameterl leaves to whosememory this paper is dedicated. 17 in a crown, arcuate, sheath 76 cm tE"aruo"o long, petiole 30 cm long, blade 220 crn Spucrss long; sheath (near top), petiole, and rachis minutely brown-dotted and thinly white- Gulubia liukiuensis Hatusima in Mem. Fac. Agric. Kagoshima Unr-n.I: 39.1964. Ryukyu Islands. : woolly, more thickly so above than below; Satakentia liukiuensis (Hatusima)H. E. Mmre pinnae erect, not drooping at the tips, 52 in Principes8: 5. 1969. on each side of the rachis, to ll0 cm Gulubia ramsayi Beccariin Webbia 3: I59. 1910. : long, 3.5 cm wide, bifid to briefly prae- Northern Australia. Gronophyllum ramsayi (Beccari) H. E. Moore in Gentes Herb. 9: 265. morse at the tips, Iower surface with up 1963. to 4 small, pale ramenta scattered along the lower l5 cm of the midrib. Inflores- Acknowledgements cence ca. 50 cm long, with about 28 branches, the lower few again branched Support of the National Science Foun- into several rachillae, incomplete pedun- dation (GrantsGB-20348X and DEB 77- cular bractslacking; rachillaeto ca. 42.5 173I9) is gratefully acknowledged. cm long, ca. 2.5 mm wide, bearing ca. Thanks are due also to Michael Galore 280 triads. Staminate flowers unknown. and the Division of Botany, Office of For- Pistillateflowers ca. 2 mm high and broad. ests ip Lae, Papua New Guineaofor logis- Fruit whitish when nearly mature, l0 X tical support of the field work, to Brad 3 mm, cylindrical and slightly curved; Young for assistancein the field, and to endospermhomogeneous. John Dransfieldfor suggestingexpansion Distribution: Fiji Islands, known only of this paper and for providing the nec- from the type locality. essary specimens.Thanks are also due to SPECIMENS EXAMINED: FITI the L. H. Bailey Hortorium for making IsleNns: Viti Levi, cutover forest on ridge facilities and specimensavailable to me. ca. 8.5 miles inland from Ngaloa,alt. ca. 260 m,22 March 1980, Moore & Phil- Lrrenerunp Crreo (holotype lips 10543 BH); inland from EssIc, Fnnoanrcr B. 1973. Pollination in some Galoa(sic.), on steepslope, alt. 600-800 New Guineapalms. Principesl7: 75-83. ft, l0 November 1977, Vodonaiualu L I9?8. A systematic histological study of 30688 (SUVA, photo only at BH). palm fruits. l.The Ptychosperma alliance. Sys- This species is not completely known tematic Botany 2: 15I-168. ANDBRADFoRD E. YouNG. 1979. A sys- yet. Staminateflowers have not been seen, tematic histologicalstudy of palm fruits. II. The and it is not clear what the color of the Areca alkance. Systematic Botany 4: 16-28.

ERRATA

Page 76, column l, Iine 20: for Manatanai read Namatanai. Page 84, column I, line 3: for Houeaforsterana read. H. forsteriana; column 2,line 4I; for Ptychosperma macarthurii read P. rnacarthuri