19721 UHL: FLORAL ANATON,IY B9 Floral Anatomy of , , and ltaya (Palmae)

Nlrar,rs W. Unr-* L. H. Bailey Hortorium, Cornell Uniuersity, Ithaca, New York 14850

This paper presentsthe floral anatomy Descripfions ol the Chelyocctrpus alliance to accom- CuBlvoc.q.npus(Fig. 1-7) . pany a current assessmentof the group including the descriptionof a new genus Chelyocarpwsulei is described from t'Moore, L972). Although reports of Moore anil Salazar 9494. Flowers, each floral anatomy in palms are few, those 4-5 mm. long and 2 by 4 mm. wide, completed have been valuable in deter- have two broadly ovate sepals, 2 rrrm' mining functions of floral organs and long by 2 mm. wide, which are distinct relationships among genera, and have or slightly joined at the base forming a provided new information on floral shallow cup around two distinct ovate structure in angiosperms (Uhl and petalsof aboutthe samesize. The androe- Moore, 1971). As the accompanying cium consists of seven (five-eight) paper (Moore, 1972) explains, the in a distinctive arrangement. genera considered here are of special One is opposite and sheathedby interest becausethey may form a primi- each sepal and the others form two rows tive alliance within the palms, and of two to three stamenseach, one row becausetwo species,ClrcIyocarpus dia- opposite each petal (Fig. 6). The nuerus and C. zrlel possessflor,al plans flower is thus wider along the axis of that are unique in rthefamily. petal insertion. Filaments of the stamens (Fig. 3a, b) are 2.5 mm.long, ventrally Mqteriqls qnd Methods expanded, and tightly encase the lower two-thirds of two (three, four) carpels. Material examinedconsisted of flowers Each carpel is 1.5 mm. long with a wide at or near anthesis. fixed in formalin- ovarian part I.0 mm. long which narrows aceticacid-ethanol in the field or garden, abruptly to a short wide style 0.5 mm. and subsequentlystored in glycerine alco- long (Fig. a). The style is slightly hol (.LU/o glyoerine, 70/o ethanol). At reflexed and ends in a slanted, distal least 25 flowers of each collection were stigmatic opening (FiS. 7). The single cleared,and six or more were sectioned ovule (Fig. 7) is borne in a locule serially, stained, and filmed as described occupying two-thirds the height of the in previous reports (Uhl, 1966; 1969a ovarian part of the carpeland terminated & b). The reader is referred to the by a large locular canal. The locular accompanyingpaper (Moore, 1972) for surface of the full descriptions and illustrations of the canal and inner ventral flowers; only brie{ descriptions,dimen- locule are lined by a glandular epithe- sions of floral organs, and collection lium. Stigmatoid tissue consisting of numbersare included in this paper. large uniseriate one- to three-celledtri- chomes with large basal cells is borne + From work supported by National Science on the margins and adjacent outer sur- Grant GB-20348X. H. E. Moore. Foundation the Jr. principal investigator. faces of the style. The tissues of 90 PRINCIPES [Vol. 16

I-5. Chelyocarpus ulei. I, sepal, 1 10; 2, petal, X 10; 3, stamen: a, ventral view; b, lateral view, XyB;4, carpel, y 25; 5, subtending bract, t 10. Details: r/, dorsal bundle; st, stamen rtace) u, ventral bundle.

style and extreme distal part of the ovary bundles of the rachilla. These bundles are more mature than the basal part of branch and the branches unite in a the carpel. Large tanniniferous cells are circular complex at the base of the scattered in the style (Fig. 7) but are floral organs. Fibrous sheathsare absent rare below. from bundles of the floral axis, stamen, An attenuatebract (Fig. 5), 5 mm. and carpellary supplies, but are present long, subtendseach flower and is vascu- on traces to sepals and petals (Fig. l, larized by eight (7-L4) traces from 2.6). outer bundles of the rachilla. One or All perianth traces arise from the two of the larger traces is a vagcular vascular complex opposite the appro- bundle and the others are fibrous strands. priate organ. Each sepal receives 12 The vascular supply to the base of each (10-20) traces and each petal 10-12 flower is seven to eight large and about traces. No fusion of petal and stamen seven smaller bundles derived from traces was observed. Sepal and petal UHL: FLORAL ANATOMY 91

the middle: 7. Iongisection 6-7. Chelyocarpus'l"iuit. ulei.6, oblique rransectionof {Iower slightly below of iio*"i. , op, untip"i"iJ* .tut"" ; as. antisepaloq.il"-", ; 6. subte-n ding braet : c, carpel; p, petai;s, sepal.For magnifications.scale' 'l'lg' /' equalsu'z mm'

traces are in adaxial rows and many a trace branched in the lower part of a branch. Branching of traces is variable, filament. even in perianth segments of the same A complex of 3-5 bundles enters the flower. base of each carpel. Just below the The arrangement of the stamens, one locule, this supply separates into a opposite each sepal and one to several dorsal, two ventral, severalshort lateral opposit" each petal, is reflected in the bundles (Fig. a), and 3-4 large strands vascular system. The traces to the anti- which enter the funiculus. The ovular sepalousstamens are bundles adjacent supply divides into 5-6 strands which to or near the mid-bundlesof the corre- can be seen in Fig. 6 in the base of the The spondingsepals (Fig. l), and tracesto funiculus and outer integument. carpel is less antipetalous stamens branch from close vascular supply of the those of many palm carpels T to petal traces (Fig. 2). Although both mature than (Uhl and Moore, I97I), and onlY the one- and two-trace stamens occur, one I dorsal and bases of ventral and lateral trace per stamen is somewhat more bundles can be discerned in cleared common in the collection examined. material (Fig. a) . In sections,however, traces, each may When a stamen has two a ring of ca. 30 procambial strands can arise from a separate part of the stelar be observedaround the locule. complex, or a single bundle may branch The hemianatropous ovule is attached near or in the base of the filament. to the ventral face of the carpel (Fig. 6) Figure f showsthe origin of two traces and the micropyle faces dorsally (FiS. to an antisepalous stamen, and Fig. 3a, 4, 6, 7). The nucellus is 3-4 cells thick 92 PRINCIPES lVol. t6 and surrounded by two integuments are distinct where free. As in Cryoso- which are free for about one-third the phila, traces to floral organs can be lengh of the ovule. The outer integu- distinguished in the solid base of the ment forms the micropyle and is ca. 6 flower. The number of sepal traces cells wide. The inner integument consists varies from 9-I5, and petal traces from of 2 (-3) cell layers with an inner layer 7-9. Filaments of the stamensare basafu of very large cells which appear tapetal. expanded as are those of the other two A large aril, reaching about half the species,and most receive one large trace. height of the ovule, surrounds the funic- Three carpels were present in all flowers ulus. The aril is very shallow on the examined, and one carpel was large with ventral side of the funiculus (Fig. 4, 6, the ovule in early seed formation. The 7) and,is not vascularized. other two carpels were abortive. The Some observations on cleared flowers ovule appears hemianatropous with a from the type collection of Chelyocarpus rim of tissue which may represent an d,ianeurus,Archer 2199 (US), and on aril present basally. Chelyocarpuschuco flowers in young fruit of Chelyocarpus resembles Cryosophila in three-parted chuco, Caaalcante s. n, can be added. whorls of sheathing perianth parts which It should be noted that these descrip- closely encasethe . tions are based on a few flowers from herbarium sheets. Flowers oI C. d,iane- CRvosopnru (Fig. &-15). urws appear somewhat less specialized Flowers of Cryosophila algentea, than those of C. ulei. Each flower has (Read,2330 B), C. sp. (Rea.d,605), C. four slightly imbricate sepals, briefly, warscewiczii (Bailey 558), and C. sp. if at all, united basally, and four alter- (Chapman Field, 197I) were examined. nate, distinct petals. Sepals and petals No significant differences were observed are ovate, and ca. 1,5 mm. longl occa- in the four species. The following de- sionally one petal is smaller than the scription and figures (B-15) are of C. other perianth members. Each sepal or drgented. The same collections of C. petal receives 4-5 traces. There is no ar gented and.C. war scewiczii w ere studied definite midvein and the two middle by Morrow (1965). His description of traces are often widely separated. Eight calyx and corolla as united for much of (-9) stamens are usually present, one their length, of three traces to sepalsand opposite each perianth part. Traces to petals, and of ovules as anatropous the stamens arise slightly above the appear incorrect for this genus (see Fig. perianth traces and are large, usually 8,9, 12). double bundles. The one to three carpels, The flower of C. argentea has a short 1.5 mm. long, resemble carpels of ltaya solid base where lower parts of sepals in shape. The single ovule in each and petals and traces to all floral organs carpel, like others ofthe alliance, appears can be recognized (Fig. 13, l4). The hemianatropous and has a large aril. three sepals (Fig. B, 13-15) are lanceo- Flowers oI C. chuco appearmore stable late, 3-5 mm. long and 1.5 mm. wide, in number of parts with three sepals, and connate for 1.5 mm. basally. The three petals, six stamens, and three sepals are, basally adnate to and closely carpels present in all examined. Sepals sheath the slightly imbricate petals (Fig. and petals are similar in shape and size 12) which are 2 mm. wide and 2.5 mm. to those of the other two species-ovate long (Fig. 9, 15). At anthesis the and ca, 2 mm. long. Sepals are connate sepals and petals form a sheath around for one-third their length, and petals the fused linear filaments of the six r9721 UHL: FLORAL ANATOMY 93

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B-I1. Cryosophila argentea. B, sepal, X 10; 9, petal, N 10; 10, stamen, X 10; 11, carpel, ;1 25. Details: d, dorsal bundle; u, ventral bundle. 94 PRINCIPES tVol. 16

I2-I5. Cryosophila argentea. 12, longisection o{ Ilower; 13-15, series o{ transections through flower. 13, transection through base of flower; 14, transection 120 microns above Fig. 13; 15, transection where floral organs are separate,400 microns above Fig. 14. Details: b, subtending bract; g, gynoecial supply intriangulararrangement;p,petal; s,sepal;sb,stamentraces; sr, lusedfilamentsof stamens. Scale, Fig. 15, equals 0.2 mm. r9721 UHL: FLORALANATOMY 95 stamens(Fig. 15) ; and six anthers, I.5 bundles forming the mid and adjacent mm. in length, are exserted forming a lateral traces o{ each sepal' Traces to radiate pattern on the top of the ovate the stamensarise as branchesof, or from sepal-petalsheath. The individual carpel a bundle adjacent to, the mid-veins of is attenuate with an ovarian part I mm. the three sepals and petals (Fig. 14). high and a tapering siyle I.5 mm. long Bundles to antisepalousstamens are free stamens, tFiS. lf ) . No definite stigma is present. below those of antipetalous The carpel is open distally with upper indicating a two-whorled arrangement ventral margins slightly reflexed at an- for the stamens. thesis. Short unicellular trichomes are The derivation of the gynoecial supply borne along the margins and down the can be seen in Fig. f3. Three bundles ventral surface {or a short distance. A extend into the center of the {lower and long stylar canal, lined by a single-layered branch, and at a higher level the epithelium, extendsfrom the basal locule branches form a triangular arrangement to the distal opening. The ovule is at- (Fig. Ia) which divides into three groups tached ventrally near the base of the of three bundles each, one group supply- locule and is described as campylotro- ing each carpel. A dorsal, two ventral pous since the embryo sac is slightly bundles, and several immature lateral curved (Fig. 11, 12). The ovule is turned bundles form the complement of each so that the micropyle faces a lateral waII carpel. Only the dorsal bundle'can be (Fig. of the locule and is thus pleurotropous. distinguished in cleared material The two integuments are separate for 1l), but a ring of ca. 14 Procambial about one-third the length of the ovule. strands around the locule can be seen The outer integument, 3-6 cells wide, in transversesections. One or two large {orms the micropyle and the two-celled traces to the ovule separate from the inner integument has the inner layer ventral bundles at the base of the funic- specialized as a tapetum. A chalazal ulus (Fig. 11). band of tannin is evident (Fig. 15). A Histological features noteworthy in short flat aril is present on the funiculus Cryosophila are several. Scatteredtannin belowthe micropyle(Fig. 1I). cells are frequent in the outer parts of Each flower is subtended by a small sepals and petals (Fig. L2-I4). As in bract (Fig. I2), which may reach 0.5 Chelyocarpus, sepal and petal traces are mm. in length, but has no vasculature. alignedin an adaxialrow (Fig. 14, 15). Three bundles from the rachilla enter Raphides are abundant in outer parts of each floral base where they branch, and sepalsand petals, in filaments of stamens, branches are united in a circular complex and in the styles of the carpels. The (Fig. 13) from which traces to all floral number of tracesto sepalsand petalsand organs arise. Within the flower, fibrous their branching is irregular. Fibrous bundle caps are present on sepal and bundiesmay branch laterally from major petal traces only. Two to three outer traces, and such branchesare more fre- (e.g., lateral traces for each sepal are the first quentin somespecies C. argentea\ bundles to branch from the complex (Fig. than in others. Closely aligned fibrous 13. outermostbundles). Bundlesin anti- or vascular bundles with large fibrous petalous positions divide to form 6-8 caps appear to provide a protective "Ience" central traces to each petal. In contrast {or inner floral organs. Fibrous to Chelyocarpus, two to three outer bundle sheaths in this genus completely lateral petal traces are branches of surround most bundles tFig. 14, 15). PRINCIPES [Vol. 16

I I i I t L il8 16-79. amicorum.. 16, sepals, connate in a triangular sheath, X 10; 17, petal, dotted margins indicate where joined to adjacent petals, X l0; 18, stamen, dotted lines outline connate part of filament, y 10; 19, carpel, y 13. Details: d, dorsal bundle; l,Tateral bundles; a,lenrral bundle.

Ir.tva (Fig. 16-23). ment accentuatesthe triangular shape of the base of the flower. The carpel is Material investigated for this new asymmetric with a large ovarian part, genus consistedof flowers from Moore, 2.5 mm. long, which is distally attenuate Salazar, and.Guti|rrez 9509. Flowers are in a funnel-shapedstyle 2.5 mm. long to funnel-shaped,ca. 6 mm. triangular (FiS. 19). Branched trichomes cover long, and have a short distinct stalk the margins of the distal opening. The subtendedby a small bract 1.5 mm. long ovarian part of the carpels is covered (Fig. 23). Connate sepalsform a basal with short, closely appressed,uniseriate sheath with free tips to 2.5 mm. high, trichomes, 3-4 cells long, and with which is adnate to a wide petal-stamen scattered low hairs, each with a short tube that surrounds the large single carpel stalk, 7 cells wide and 4 cells high, and (Fig. 2l-23). The free parts of the a distal mound-shapedpart about 10 ovatepetals (FiS. 17) are approximately cells across. 3.5 mm. wide and 2.5 mm. high. The The bract subtendingeach flower has filaments of the 19-24 stamensare large, no vasculature. The epidermis of the subulate basally, and attenuate distally short floral stalk is papillose, and the (FiS. 18) . Stamensare insertedwith one cortex, ca. eight cells wide, may have or two opposite each sepal and ca. six in some irregularly distributed fibrous a row oppositeeach petal. This arrange- bundles unconnected to other strands. r9721 UHL: FLORAL ANATOMY 97

20-23. Itaya arnicorum. 20-22, seies of transections through {lower; 20, transection of flower base, sepals partly lree;2I, transection 300 microns above Fig. 20, petal and stamen traces mostly separate; 22-. tra-nseciion, 400 microns above Fig. 21, petals patly separate from stamen traces (without fibrous caps); 23, longisection of flower. Details: ar, aril; as, antisepalous stamen traces, divided in Fig. 22; j, fllament; p, petal; p-st' petal stamen traces partly separated; r, raphide sac; s, sepal; st, stamen trace. Scale, Fig. 23, equals 0.2 mm. 9B PRINCIPES lVol. 16

Twenty to twenty-five immature bundles, tocule (Fig. 21). The ovule is supplied someprocambial, enter eachpedicel from by six large and four to five smaller the rachilla. Outer bundles in the flower bundles. Derivation of the ovular supply base are joined in an incomplete vascu- is variable. Traces to the ovule arise lar complex from which traces to sepals, from a ventral complex composed of petals, stamens,and a few bundles to the ventral, lateral, and stelar bundles, but carpelarise. the exact make-up of the complex varies Sepals in this genus receive fewer from flower to flower. traces than those of Chelyocarpus and The hemianatropous ovule has a large Cryosophila and there is no constant aril (Fig. 19, 23) which is partially supply. A sepal may have two rather adnate to the carpel wall (Fig. 22). large parallel vascular bundles, only a I-ower parts o{ the aril have one to two central vascular bundle, a central and outer layers of large elongate,probably several fibrous bundles, or may lack glandular cells. The outer integument, vasculature (FiS. 16). Frequently a nine cells wide, forms the micropyle and sepal trace branches at about midJength the second layer of the two"celled inner to produce a lateral petal trace. Each integument is tapetal. The nucellus is petal receivesfive or six traces (Fig. 17). three or more cells wide. The micropyle The origin of stamentraces is variable, usually points toward the dorsal wall but basically {ollows the pattern of inser- of the locule (Fig. 22), but the ovule tion with traces to antisepalous stamens is sometimes obliquely oriented so that arising from bundles opposite the sepals the micropyle is directly laterally. and with most antipetalous stamen traces arising as basal branchesof petal traces Discussion (Fig. 2I). One or two antipetalous Studies of floral anatomy support the stamen traces may be derived directly alliance oI Chelyocarpus, Cryosophila, from bundles of the peripheral complex. and ltaya, the designation of Chelyocar- The bundle to a second antisepalous pursas the least specialized and oI ltaya stamen often branches from the trace to as the most advanced within the group, the first (Fig. 21, 22). Tracesto stamens and the resemblancesoI ltaya to members are large single bundles which have oI the Thrinar alliance. narrow fibrous caps near their origins l. Floral vascular systems of the and extend only a short distance into three genera are alike in that a complex the baseof each filament. of vascular tissue is formed in the base The inner bundles of the stele of the of the flower and gives rise to perianth, pedicel and some small branches from stamen, and most gynoecial traces. The the perianth complex enter the base of complexesapparently reflect a shortening the carpel where about 12 large bundles or suppressionof the floral axis so that form a central ring (Fig. 20) which the insertions of floral organs are closely separates into ovular traces and major appressed. In addition, floral organs bundles of the carpel wall. In cleared and their vascular supplies may be material, a dorsal, two ventral bundles, adnatein varying degrees(Cryosophila, and four pairs of smaller lateral bundles Itaya). More elongatefloral axeswhich could be distinguished (Fig. 19). Each lack vascular complexes are present in ventral bundle divides about mid-height other palm groups as described in and the two branches extend into the Rhapis (Uhl, Morrow, and Moore, L969), style. Additional immature lateral bun- Nannorrhops (Uhl, I969a), and some dles provide ca. 24 strands around the ceroxyloidpalms (Uhl, I969b). 19721 UHL: FLORAL ANATOMY 99

2. Sepals and petals of the three 4. Carpels of all genera are unspe' genera are structurally alike. Ground cialized in shape with ovoid ovarian tissue is parenchymatous with scattered parts which taper to distally open styles. tannins (Cryosophila) and raphides (all No definite stigmas are present, but tri- genera). A row of bundlescomposed of chomes are borne on and near the larser vascular bundles with distinct margins of the openings. Ventral sutures fibious sheathsand small fibrous strands, are closed except distally, although in is aligned along the adaxial face of each Cryosophila a ventral notch indicating perianth part. Sepals and petals are the position of the suture is present similar in Chelyocarpus and Cry osophila, above the attachment of the ovule. The although in the latter, petals are some- dorsal and two ventral bundles of the rvhat different in shape (cf. Fig' 2, 9). carpels in all three genera are larger It should be noted that the number and than the lateral bundles which are branching patterns of the adaxial row difficult to follow becauseof immaturity' of bundles is variable. The perianth is Derivation of the ovular supply is from most reduced in ltaya, where sepals,and a ventral complex of bundles with some stamen bases are adnate in a flaring traces direct from the carpellary stele in tube. Sepals are also more distinct from Itaya. Carpels oI ltaya are larger, have petals in ltayo where sepals are smaller a larger vascular supplY, and more than petals, and sepal vasculature is mature bundles. reduced or lacking. The vascular supply 5. Ovules are structurally alike in of petals in ltaya resemblespetal supplies the three genera. Although the ovule in in the other genera except that fewer Cryosophila is considered campylotro- qnly (five to six) tracesare Present. pous, it varies in a slight curvature 3. Within these genera, the androe- of the nucellus from the hemianatropous cium shows more variation than anY ovules of the other two genera. All other floral organ. Two patterns are ovules have funicular arils, two integu- present. Stamens in Cryosophila ate ments that are free for one-third their arranged in two whorls of three, a lower length, and a two-layered inner integu' whorl oppositethe sepals,and an upper ment with a tapetal inner layer. Ovules whorl opposite the petals' ln Chelyo- in Chelyocarpus and, Itaya are crassi- carpus and ltdyd', one (-two) stamens nucellate. In Cryosophila, the nucellus are opposite each sepal, and two-six apparently disintegrates early and was in a row opposite each petal. Vascular not entire in any of the collections suppliesshow the samearrangement with examined. traces to antisepalous stamens derived Among the palms that have been as branches of sepal traces or from studied, floral structure of this group bundles adjacent to sepal traces in the is more like that oI Rhapis and Nannor- complex. Bundles of antipetalous sta- rhops where sepal and petal traces are mens are derived as branches of the many and ventrally aligned, than the bundles supplying the opposing pe'tal, ceroxyloid palms where perianth traces or from bundles in the complex adjacent are fewer and in a median row. Reduc' to petal traces. Both one- and two-trace tion in number of sepal traces was also stamens are present in Chelyocarpus. found in the ceroxyloid grouP. The Itaya, again shows reduction in that adnation of reduced sepals and petals, stamen traces are extremelY short more than six stamenswith short traces, bundles. and the single carpel in Inya seemtrends 100 PRINCIPES lVol. 16 which are further developed in the petals, and thus they are not branches flowers oI the Thrinarc alliance as stated of the same bundles that form the peri- in the accompanying paper (Moore, anth traces, as in Cryosophila and fior 1972). In the geonomoidgroup of palms, most stamens in ltaya. Variability in Asterogyne (Aristeyera), sepalshave a the number and branching of sepal and number of traces in a median position, petal traces and in number of perianth but only a single trace supplies each parts, of stamens,and of carpels should petal. The characters of the perianth alsobe noted for this genus. in the geonomoids, however, may be Several floral characters cannot yet correlated with the submergence of the be assessedwith respect to primitiveness {lowers in pits. or advancementwithin the palms. These Some suggestionsas to the function- include the vascular supplies of sepals, ing of these flowers can be surmised petals, and stamens, wide or ventrally from their structure. Pollinating agents expanded versus slender filaments, pat- for the alliance are not known. The open terns of stamen arrangement within the anthers but extreme immaturity of multistaminate androecia, and some bundles and tissues in the carpels, and aspectsof ovules. Characters that might anthers protruding from young buds in be considered primitive in this alliance one collection of Cryosophila (C. sp.) include variation in number of floral suggest that these genera are protan- pats (Chelyocarpus),distinct and simi- drous. A number of structural aspects lar sepals and petals, carpels of unspe- appear to protect the developing ovules. cialized form-lacking definite stigmas These are the abundant raphides in but with large distally open stylar canals sepals, petals, stamen filaments, and bearing marginal trichomes, and crassi- carpels; and tannins-particularly the nucellate ovules with large funicular large inclusions in the exposed styles of arils. Chelyocarpus, and the scattered tannini- Lrrrn,trunr Crtnn ferous cells in the petals and sepals of Moonn, H. E., Jn. 1972. Chelyocarpusand Cryosophila. The fleshy stamenfilaments its allies Cryosophila and' Itaya (Palmae). in Chelyocarpus and ltaya tightly encase Principes 16: 67-88. the immature lower parts of the carpels. Monnow, L. O. 1965. Floral morphology and (Pal' The adaxial row of fibrous or heavily anatomy of certain Ph.D. Thesis, Cornell University, in sepals and petals, mae). sheathed bundles Ithaca, New York. particularly in Cryosophjla, appears in and anatomy "stockade" Unr,, N. W. 1966. Morphology cleared material like a around of the inflorescence axis and flowers of the stamensand carpels. a new palm, Aristeyerd spicdta. Journal Within the alliance, Chelyocarpus is of the Arnold Arboretum 47 z 9-22. 1969a. Anatomy and ontogenY of the specialized in that perhaps the least cincinni and flowers in Nannorrhops there is less connation and adnation of ritchiana (Palmae). Journal of the floral organs and of vascular supplies. Arnold Arboretum 50: 4ft-431. Sepals may be slightly connate basally 1969b. F"loral anatomy ol luania, (Palmae-Are- but other organs are usually free. Occa- Rauenea, and, Ceroxylon coideae). Gentes Herbarum 10: 394-411. sionally two antipetalous stamens are & H. E. Moont, Jn. 1971. The palm fused. There is no adnation of sepal and gynoecium. American Journal of Botany petal traces which arise from separate 58:945-992. bundles of a complex. Vascular traces 1969. Anatomy of the palm Rhapis to the stamensusually arise from bundles excelsa, YII. Flowers. Journal o{ the adjacent to those supplying sepals or Arnold Arboretum 50: f3B-152.