6B PRINCIPES lVoL. 33

Principes,33(2), 1989, pp. 68-73

Larpoxylont1 I macrospermum

Jottt'tL. Down e.rutNererm W. Usr 18 Amelia Street, Albion 4010, Queensland,Australia, and L. H. Bailey Hortorium, 467 Mann Library, Cornell Uniaersity, Ithaca, NY 14853

"Genera When Palmarum" went to tapering to an irregularly rounded, more press in 1986, Carpoxylon was known or less bifid tip, stiff, coriaceous, horizontal from fruit only. Since lack of information to erect, glabrous adaxially, with numerous made it impossible to include the genus in punctate scales abaxially, midveins most the hierarchy of the classification, it was prominent, marginal veins next largest, two "Incertae placed in an Sedis" group along other pairs of large veins conspicuous. with the Madagascar gerrus Masoala. The transverse veinlets not evident' Inflores- rediscovery of CarpoxyLoz, described in cences infrafoliar, branched to three orders the accompanying paper (Dowe 1989), basally, to one order distally, branches allows us to provide here a complete stiffiy spreading; peduncle short, stout, description of the genus and its one species elliptical in cross-section; prophyll com- and to consider further the relationships pletely encircling the peduncle at insertion, of this unusual palm. tubular, two-keeled, tapering distally, split- ting abaxially, tomentose; peduncular Carpoxylon H. A. Wendland& Drude, bracts 2, the first inserted shortly above Linnaea39: I77. 1875. TYPe:C. the prophyll, the second an equal distance maGrospermumH. A. Wendland& above the frrst, both tubular, complete, Drude.(Fig. l, CoverPhoto; see also tapering to rather short pointed tips,.gla- Fig. l, accompanyingpaper, p. 65). brous, caducous; scars of 2-3 incomplete bracts above the inner peduncular bract; Moderate, solitary, unarmed, Pleo- rachis about twice as long as the peduncle, nanthic, monoecious palm. Stem erect, rachis bracts low, ridgelike in slitlike cav- longitudinally fissured, swollen basally and ities, subtending ca. l0 primary branches; with a boss of adventitious roots, prom- primary branches stout, dorsiyentrally flat- inently ringed with slightly sunken leaf tened, with a short bare part and two large scars, internodes short. Leaves regularly lateral pulvini at the base, bearing very pinnate, spreading but arched towards the shallow bracts, each in a slitlike cavity, tips, neatly abscising; sheaths forming a subtending rachillae; rachillae angled, crownshaft, crownshaft glossy, glabrous to tapering, also with basal pulvini, rachilla lightly scaly, splitting opposite the petiole; bracts shallow, rounded, subtending triads petiole short, wider proximally, ridged of flowers for about one-third their length, adaxially, rounded abaxially; rachis flexi- paired staminate flowers with some inter- ble, broadly ridged adaxially at base, nar- mingled solitary staminate flowers above rowly ridged distally, rounded abaxially, the triads. and solitary staminate flowers extending beyond the apical leaflets in a distallv. in triads one staminate flower often flexible tip; leaflets subopposite, in one rank, distal and one lateral to the pistillate flower' apically and basally inserted at right angles flowers lateral to each other in staminate to the rachis, more obliciuely inserted at dyads, rachilla ending in a short bare por- midleaf, leaflets single fold, erect, linear, tion; first bracteole surrounding the pistil- r9891 DOWE AND UHL: CARPOXYTON ffi

ti,r1\ I n il{\1 n UJ(.(L//frii

Cafpoxylon macrospermum, a, portion of rachilla in bud x I %; b, scars of staminate dyad x 3Vz;c, staminate bud, x6/t; d, staminate bud in vertical section x6/t e, two stminate petals x6%; f, staminate petal in two views x 67ti g, portion of rachilla with staminate bluds x 2Tt; h, staminate bud sepalsremoved x 6Tz; i, slaminate pelals x 6%; j , scars from floral triad x 3Tt; k, triad x 3yst I, androecium x 6Ts.;m, stamen in three views x 6?4; n, pistillode x62A; o, young pistillate bud x67t; p, pistillate sepals x62A; q, pistillate bud in vertical section *6yu: ,, pistillate petals x6Ts; s,'gynoecium with staminodes x67t; t, end of stigma x I; u, end of stigma enlarged x3; v, fruit x/z; w, frtit in vertical section xTsi x, endocarp xTzi y, seed in three views x?6; z, seed in vertical section xVz. rU PRINCIPES lVoL. 33 late flower large, rounded, coriaceous, the scars whitish, prominent, internodes 7 cm second smaller and more shallow. Stami- long near base to 2 cm long distally. Leaves nate flowers very asymmetrical in bud, regularly pinnate, 3.5 4.0 m long; crown- rounded or pointed apically; sepals 3, dis- shaft green, I.5-2.0 m long, somewhat tinct, irregular, imbricate basally, keeled, larger in diam. towards base; petiole 25 prominently ridged when dry; petals 3, cm long or less, wider proximally; rachis distinct, valvate, tips thickened, ridged wide to 6.5 cm at base, 4.5 cm wide in when dry; stamens 6, filament- slender, middle, 4.0 cm wide distally, extending inflexed at tip, anthers more or less sag- beyond apical leaflets in a flexible tip about ittate basally, slightly bifid apically, dor- 12 cm long; leaflets about 70 on each side sifixed just below the middle, latrorse, ver- of rachis. proximal ones ll4 x 1.5 cm, satile, connective tanniniferousi pollen mid-leaflets I22 x 3.2 cm, distal ones 36 elliptic in polar view with finely reticulate x t.5 cm. Inflorescences infrafoliar; tectate exine; pistillode elongate, slightly peduncle stout, elliptical in cross-section, Ionger than anthers in bud, tip enlarged, about 14 cm long, 4 cm diam.; ProPhYll rounded. Pistillate flowers in young bud, 70 cm long, about 8 cm wide, peduncular irregular, rounded; sepals 3, distinct, very bracts two, the first inserted ca. 5 cm broadly imbricate, extremely thick basally; above the prophyll, the second 5 cm above petals 3, very broadly imbricate, thick the first, each 70 x 7 cm taPering to a basally, tips thick, valvate; staminodes woody tip ca. 5 cm long; scars of two to joined in a shallow ring with about 5 broad three incomplete bracts above the inner toothlike tips; gynoecium irregularly obo- peduncular bract; rachis ca. 36 cm long, void, unilocular, uniovulate, stigmas 3, rachis bracts subtending l0 primary fleshy, ovule erect at stage studied, ? anat- branches; primary branches stout, lower ropous. Fruit obovoid to ellipsoidal, red at ones to 2 cm wide with a basal bare portion maturity, stigmatic remains eccentrically 7-8 cm wide; rachillae stout, ca. 5 mm apical, epicarp smooth, wrinkled basally diam. and 30-40 cm long. tapering. also when mature, mesocarP thick, with close with basal pulvini, bearing spirally packed longitudinal fibers, endocarp rather arranged, rather distant triads, 1.5-1.0 thick, whitish, bony, longitudinally ridged, cm apart, for about one-third their length, large operculum over embryo. Seed obo- rachillae much reduced in diam. to 2 3 void, raphe elongate, branches longitudi- mm distally, first bracteole surrounding the nal, endosperm homogeneous. Germina- pistillate flower shallow, 2-4 mm high, tion adjacent ligular, eophyll bifid. rounded, coriaceous, evident in fruiting Distribution: One species rediscovered rachillae, second bracteole smaller and 30 November 1987 on Espiritu Santo, more shallow. Staminate flowers very ir- Vanuatu where growing in silty alluvium regularinbud,2.S 4.5 x 2.0 mmin young on the edge of a small stream. The pop- material examined, rounded or pointed api' ulation may have been planted (see accom- cally; sepals2.5 x 3.0 mm; petals 2.9 x panying article), thus the wild location is I.7 mm; stamens six, dorsifixed near the uncertain. The original description men- middle, filaments slender, 1.5 mm long' tions the mountains of the Vanuatu Islands. inflexed at tip; anthers 2.0 mm long; pis- tillode 2.0 mm long. Pistillate flowers stud- bud. 2 x 6 mm. irreg- Carpoxylonmacrospermum H. A. ied in very young ular; sepals various in size, abotr 4 x 2 Wendland & Drude, Linnaea 39: I77, mm; petals imbricate, also not completely Plate 1, Fig. 3. 1875. Type: Vanuatu, developed and varying in size, about 3 x fruit only,? in GOET,.not found. 2 mm1' staminodes 0.5 mm high; gynoe- Stem erect to IB m, ca. 35 cm diam. cium obovoid,2 mm high x 1.5 mm wide. DBH, baseenlarged, 50 cm in diam., leaf Fruit slightly obovoid to ellipsoidal, 6 x 19891 DOWEAND UHL: CARPOXYLON 7I

3.5 cm, stigmatic remains eccentrically 2- Seed irregularly ridged, furrowed arrd sculp- tured with adherent fibers ...-...,,.... ALsmithkt apical; epicarp thin, mesocarp 2 mm thick 2. Seed :t small, not ridged or sculptured ...... 3 thick, with large fibers, endocarp 3 4 mm 3. Staminate flowers borne in vertically oriented thicker basally, longitudinally ridged, bony pairs sunken in distinct depressions distally, below ridges, operculum circular, large. smaller than and lateral to pistillate flowers fruit large, with Seedlarge, ellipsoidal,4 x 2.5 cm, raphe proximally on the rachillae; apical stigmatic remains. Fiji .,.,,. Neoxeitchia extending laterally, little fibers abundant, 3. Starrrinate flowers borne in horizontally ori- anastomosing, endosperm homogeneous ented pairs distally, lateral to pistillate flowers with central cavity; embryo basal. Ger- proximally on the rachillae; fruit moderate, mination adjacent ligular, eophyll bifid. rarely large rvith apical, lateral, or hasal stig- matic remains ...--...... - 4 4. lnflorescence interfoliar; fruit covered rvith Discussion prominent corky warts; stigmatic remains basal fruit ...... 5 Curpoxylon, then known only from in ...... , 1. Inflorescence interfoliar, or infrafoliar, fruit fruit, was put in AreceaeIncertae Sedis in merely to granulose rvhen "Genera ,snooth or pebbled Palmarum" (Uhl and Dransfield dr-v; stigmatic remains various ...... -...... ,..6 1987).The newlycollected material allows 5. Peduncular bract inserted near the base ol a subtribalplacement. The large opercu- the peduncle; fruit more than 2.5 cm in diam- eter. Marquesas Islands ..-....-..,..,...... Pelagodoxa lum over the embryo places the genus 5. Peduncular bract irrserted at the apex of the clearlyin SubtribeIguanurinae of the Are- peduncle: fruit 1.5 crn in diameter or less. ceae,where it appearsmost closelyrelated New Guinea ..--...... ,...... -...Sommieria Io Clinostigma. This relationshipis fur- 6. Flou'ers borne in laterally compressed pits, the long, hairy pedicels; fruit rvith by preliminary cladistic staminate on ther supported ,stigmatic remains lateral in lolver %; seeds studiesof lguanurinae;using a data base ridged and grooved. Southern India and Nico- of 32 characrers, CarpoxyLon and Cli- bar Islands ...... Bentinckkt nostigma are indicated as sister genera 6. Florvers sessile or impressed in the rachillae (Uhl and Dransfieldunpubl.). CarpoxyLon but neither in laterally compressed pits nor the staminate with hair covered pedicels; stig- differs from Clinostigma in lacking stilt matic remains apical to basal; seeds smooth roots, in the stiffiy ascendingrather than the pendulouspinnae of most speciesof Leaflets several-ribbed lvith praemorse apices Clinostigma, in two rather than a single or leaf blades, lvhen not divided laterally. lvith usually inter- bract, in inflorescencebranches toothed rnargins; inflorescences peduncular foliar; triads shallolvly to deeply sunken in stiff and spreadingrather than more or depressions in the rachillae. Malay Peninsula, Iesspendulous, and in a ridged,bony rather Borneo, Java, Sumatra Iguanuru than a thin crustaceousendocarp. Species LeaHet" I -ribbeJ wilh acule or acumitraleapi- of Clinostigmn are poorly known as are ces; inflorescences various; triads super{icial lguanurinae;more thoseof other generaof B. Seed with rurninate endosperm ...-...,, ,...... -...9 field studiesare neededand may change B. Seed rvith homogeneous endosperm ...... ,,... 12 the circumscription of Carpoxylon. A 9. Leaf sheaths splitting opposite the petiole, not "Key revised to the lguanurinae"' with forming a prominertt crorvnshaft; inflores- least in bud, sometimes Carpoxylon now included follows: cences interfoliar, at infrafoliar at anthesis or in fruit; the peduncle elongate, prominent, usually as long as the rachis or longer. Philippines to Micronesia, KEv ro rHE GENERAoF New Guinea, Solomon Islands ..,. Heterospathe q. Leaf sheath. tubular. forming a prominenl THE IGUANURINAE crownshaft; infl orescence infrafoliar ; peduncle l. Prophyll completely encircling the peduncle usually much shorter than the rachis ...... ,...,.l0 at insertion, leaving a circular scar when I 0. Inflorescence lackirrg branches adaxially except caducous: stamens 6 or more ..-,., 2 at the apex, branched to I order only and the l. Prophyll incompletely encircling the peduncle lolver branches t ascending, not divaricate at insertion, open abaxially, leaving an incom- from the rachis at a 90o angle; fruit black at plete scar upon falling; stamerrs always 6 ... 20 maturity. Mascarene Islands,.... Dictyosperma 72 PRINCIPES [Vor. 33

10 Inflorescence with branches spirally arranged, 20. Leaf sheaths with minute scales, split opposite the lower branches abruptly divaricate at about the petiole and not forming a crownshaft; a 90'angle from the rachis and again once- peduncle short; fruit small, 1.4 1.6 cm in or twice-branched; fruit yellow, orange, or red diameter, with tannin cells interior to sclereid I1 layer. Clinosperma ll. Stamens 6-9; pistillode prominent. Nicobar 20 Leaf sheaths densely scaly, tubular and form- Islands, Malay Peninsula, Moluccas, New ing a prominent crownshaft; peduncle elon- Cuinea to the Solomon Islands ...... gate; fruit large, ca. 3.2 cm in diameter, lack- R hopaloblaste ing tannin cells. New Caledonia ...-..---..--Laaoixia ll. Stamens 15 30 or more: oistillode minute or 2r. Seed terete or 2lobed in cross-section, ovoid, lacking. New Guinea, Solomon Islands ..-...-...--.. ellipsoidal, globose or rarely kidney-shaped in Actinorhytis outline....-...... , .-.--.....,...... 21 12. Staminate flowers mostly larger than the pis- 2r.Seed irregular in cross-section, externally tillate; stamen filaments inflexed at the apex angled or intricately ridged, furrowed, and in bud; anthers dorsifixed, with elongate con- sculrtured ...... 23 nective, not didymous -.,.-.,.-..,13 22. Fruii with apical stigmatic remains ,-... ., .....22 12. Staminate flowers mostly smaller than pistil- 22. Fruit with lateral stigmatic remains. New Cal- late; stamen filaments erect in buds; anthers edonia ...--...-,...-,...-,,, ...... Basselinia didymous . ,-.,,-..,.,,,...... -...-...... -..... 19 23. Stilt roots prominent and stout; pistillode of 13. Stamens 12; fruit with basal stigmatic remains, staminate flower shorter than stamens; fruit lacking a shell of sclereids. New Caledonia .., often curved apically. New Caledonia -,,-.,,--..,,--. c'o'::r?"';Z ...... Cam pecar pus 13.$;;; 6;ili;;;i;;. ::::::::: Stilt roots not prominently developed; pistil- 14. Endocarp minutely pitted; seed with lateral lode of staminate flower longer than stamens, embryo. New Caledonia Alloschmidia columar; fruit straight. New Caledonia and 14. Endocarpnot pitted; seedwith basalembryo Loyalty Islands l) 24. Leaf sheaths split opposite the petiole in bud, TJ Leaf sheaths split opposite the petiole; inflo- not forming a prominent crownshaft; inflo- rescence interfoliar. Lord Howe Island ..-...-...... rescence among the leaves in bud, becoming ...... Lepi d or r h ar h i s infraoliar in fruit; peduncle elongate, much l5 Leaf sheaths forming a prominent crownshaft; exceeding the rachis; prophyll and peduncular inflorescence infrafoliar .--...--..-....-...-.....--...-...-....-...... ,I 6 bract more or less persistent, at length mar- l6 Inflorescence densely tomentose; fruit with cescent; inflorescence branches with long bare apical stigmatic residue. Ryukyu Islands .,.-..,,- basal portions, prominently swollen at the ...... Satakentia insertion, stifiy and divaricately l-branched 16. Inflorescence glabrous or at most minutely or the distal undivided. Fiii and New Caledonia hairy; fruit with subapical to nearly basal stig- Cyphosperma matic remains ...... -...-...-.17 24 Leaf sheaths forming a prominent crownshaft; 17.Complete peduncular bracts two; endocarp inflorescence infrafoliar; pedmcle shorter than hard, moderately thick, with longitudinal the rachis; prophyll and peduncular bracts ridges. Vanuatu -... Carpoxylon caducous; inflorescence branches without a t7 Complete peduncular bracts one; endocarp long basal bare portion, nor swollen at the thin or thick and prominently sculptured -... lB insertion -...---.--....-..--...... 24 18. Stilt roots usually developed; staminate flow- zJ. Staminate flowers symmetrical; pistillode thick, ers markedly asymmetrical, with short, trifid columnar, longer than the stamens in bud, pistillode and acute sepals and petals; fruit expanded into a broadly capitate apex; fruit lacking sclereids but with prominent, often subglobose with stigmatic remains lateral in greatly thickened fibers. New Ireland to Samoa upper third, the surface minutely granular- papillate. New Caledonia Veillonia I8. s;ii;;;," ;;; ;;;"r,P"il;;;;i; ^?J' T:::::" t< Staminate flowers slightly to markedly asym- symmetrical, with pistillode as long as stamens metrical, the pistillode elongate-conic to angled- in bud and rounded sepals and petals; fruit columnar, shorter than the stamens in bud, with a layer of short sclereids beneath the not broadly capitate; fruit smooth or drying exocarp. New Caledonia Moratia pebbled but not granular-papillose .-...-...... 25 I9. Fruit ellipsoidal, with basal stigmatic remains: 26. Bracteoles surrounding the pistillate flower sclereids lacking in mesocarp but tannin cells senallike: anthers with locules rror coulrnuous present. New Caledonia ...... Brongniartikentia bui interrupted by sterile connectivelike areasl l9 Fruit globose or nearly so,.with lateral stig- fruit drying den.ely pebbled and shouldered: matic remains; mesocarp with a shell of short mesocarp not readily separable from the stony, sclereids beneath the exocarp .,,-.,,,,,,,-.-,...... -...-...l9 intricately sculptured, 4-angled endocarp with le89l DOWE AND UHL: CARPOXYLON 73

dorsal groove, flanked by 2 ridges. New Cal- John Dransfieldfor critically readingthe edonia ,...-.....-...... -. manuscript. 26. Bracteoles surrormding the pistillate flower very narrow, rarely (P. dennisii) with a slender processbut never sepallike;locules of anthers LrreRerunn CItBo continuous;fruit globoseor subgloboseor col- Down, J. 1989. The unexpected rediscovery of lapsing and drying wrinkled but not pebbled; Carpoxylon. Principes 33(2): 63-67 . mesocarp with a shining inner layer adjacent Unr, N. W. AND J. DReNsrrero. 1987. Genera to and readily separated from the endocarp, Palmarum: a classificationof palms basedon the endocarpsharply -angledto variously ridged work of^Harold E. Moore, Jr. L. H. Bailey Hor- and sculptured but always with a dorsal ridge. torim afid The International Palm Societv. Allen New Britain to Fiji ...-..-..--.-..--.-..-...Physokentia Press. Lamence. KS. Acknowledgments The authors thanl

CLASSIFIED

SEEDS WANTED: Seeds,germinated seeds,and seedlingsof palms, ferns, cacti, tropical and subtropical shrubs, trees, flowering and ornamental wanted in commercial or semi-commercialquantities. LOTHAR SEIK, Pfalzgrafenring 2,D-7403 Ammerbuch 3, W'est Germany.

DU/ARF RHAPIS EXCELSA, Seven green and variegated varieties available. NEW 'oSecret BOOK, of the Orient," ti comprehensiveguide to Rhapis palms-52 pagesfully illustrated. Catalogue $1. Book and catalogue $5 ppd. ("Secret of the Orient" is also available from The Palm Society Bookstore). RHAPIS GARDENS-PS, P.O.D' 287' GREGORY,TX 78349,

PALM SEEDTINGS. 4" , gallon, and 2 gallon palms shippedcountrywide, barerooted,in moist sphagnum. Ran:enea,Neodypsis Lostelliana, Coccothrinax crinata, rare ptycho- spermas,solitary caryotas, Zornbia, Kerriodoxa, Johannesteijsma'nnia m,o'gnif,ca,rare Chrysalidocarp&s spp., Oraniopsis, Hyphaene, Gulubia, Cryosophila, unusual liviston- as, Arenga undulntifolia. Please send stamped envelope for listing. CAROT GRAFF, 6600 S.S/. 45 St., Miami, FL 33155. (305) 666-1457.