19721 UHL: FLORAL ANATON,IY B9 Floral Anatomy of Chelyocarpus, Cryosophila, 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 stamens in a distinctive arrangement. genera considered here are of special One stamen 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 gynoecium. 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.