BULL. BOT. SZJRV. INDIA - Vol. 20, Nos. .I-4: pp. 124-132, 1978

EPIDERMAL STUDIES IN EOPHYLLS UWENILE ) OF sC&IE ARECOID PALMS

Botanical Swmy of India, Howrah

ABSTRACT

The eophylls in palms are morphologically very consistent for each palm group and they show reniarkablc distinguishing characters. Epidermal studies on the eophylls - of *&teen Arecoid palms show that in all, five different types of epidermal complexes can be recognised within the group and this information IS of taxonomic importance.

INTRODUCTION Ghosh and Davis (1973) studied stomata The post cotyledonary leaves or the plum- and trichome characters in adult and juve- ular leaves in palms are generally bladeless, nile leaves of Areca catechu Ihn. along sheathing and scale-like and their number with six non-arecoid palms. The present is more or less constant in each species. The study on 16 arecoid palms was undertaken first foliage which develops in palm to place on record further information on seedlings is termed as 'eophyll'. Its shape the epidermis of eophylls and the nature of is varied and constant which in turn offers stomata1 complex. The resulting data have a good diagnostic character. Tomlinson been evaluated in respect of their taxonomic (19%) cited three major shapes of eophylls importance. and on the basis of their relationship with MATERIALS AND METHODS the adult leaves, categorised eleven distinct The eophylls from the arecoid species, types. namely, Areca triandra Roxb., A. macro- The arecoid group palms are composed of calyx Zipp. ex Bl., Archontophoenix alex- 88 ge,nera and 760 species (Moore Jr, 1973) andrae (F. Muell.) Wendl. et Drude, A. and have distribution in the humid tropics cunninghamiana Wendl. et Drude, Benti- of both the eastern and western hemispher- nckia nico barica (Kurz) B.ecc., Carpentaria es. Eophylls of most of the arecoid palms acuminata Becc., Chrysalidocarpus mada- are bilobed, entire in Roystonea or pinnately gascarien.sis Becc., C. rnadagnscariensis var. compound in Howeio and Reinhardtia lrtcu bensis Hort., Dictyosperma a1 bum (Tomlinson 1961). The anatomy of leaves (Bory) Wendl., Norman bya normannbyi in palms and stomata in particular, has been (Hill) Bailey, HydriasEele wendlandiana (F. studied by many workers. Among others, Muell.) Wendl. et Drude, Ptychosperma Tomlinsan. (1961) made a detailed morpho- macarthurii (Wendl.) Nichols., Rltopalo- logical study of stomata, trichomes etc. of blaste augusta (Kurz) Moore, Roystonea bor- twentyfive arecoid spe$ies. Bavappa (1966) inquena Cook, R. regia (H.B.K.) Cook, also made detailed observation on the epi- Veilchia merrillii (Recc.) Moore, were col- dermal characters of two arecoid species, lected from the seedlings raised at the Areca catechu Linn. and A. triandra Roxb. Introduction Nursery of the Indian Botanic Garden, Howrah, hetween the years 1971 to -- * Date of receipt :14.6.77, Date of acceptance : 23,4,79 1975, The seeds of Areca macrocalyx and N-byo normunbyi were dlected by dle, two halves unequal one slightly broad- the senior author from the Central Planta- er, lanctolate, affx acute. tion Crops Research Institute's Experimcnt- Abaxial . epidermis : Stomata intercostal, a1 Starion at Vittal, South Kanara. Seeds of irregular, widely spaced ; latern1 sub-cells .2, wendlcntdiana, Archontophom- polar cells not distinct, slightly ahorter than ix akxandrae and Carpentaria acuminata the epidermal - cells and .arching on the were obtaind from Singapore Botanic Gar- guard cell. Intercostal cells irregular, cell dm. Seeds of Roystonea borifiguena and walls not sinuous. Costal cells uniformly Vcirchio ~rr'!Iiiwere respectively received elongated. Trichon~eaboth costal and in- from Florida and Philippines. Seeds of the tercostal, longer than the stomata, cells mt of the palms -re from the Indian sclerosed. Manic Garden's own colltcti~s.The seeds were sown by the method adopted by Basu and Mukherji (1972) and eophylls from setdlings of each species were collected and preserved in two sets, one set of voucher specimens deposited in the Garden Herbari- um and the other set bearing the same col- lection numbers were fixed in F.A.A. for twentyfour hours and stored in g: I mlu- tion of ethjI aIcohoI and 10% glycerin: Be- fore peeling, the middle portion of each caphyll.was cut into small sectors and tho- roughly washed in running water for five minutes. The clearing technique was followcd from Ghase and Davis (1973) with little modifica- tion; The cleared epidermal sectors were mained in Safrannin and Light green, de- hydrated in 70% ethyl alcohol and mounted on glass slides in glycerin jelly. Thevcover slips were sealed with melted paraffin. The frequency of atomata and trichomes were calculated from field of r mm square ; mea- .. aurcment of guard cells, trichomes, epider- H8te-l :Fig. 1A: Trichomes: 1. PI~~OJP"RMmadurn. 2. flrm~nbprarmanbyi. 3. Hvdrinstrlr wendlondMM. mal cells and camera Iucida drawings and 4. Area )riondra. microphotographs were made with eyepiece objective combination of 15x 20. The st* Adaxial epl2crrni.c: Guard cells of the matal indices were calculated separately for stomata wide. Epidermal cells rhombohcd- adaxial and abaxial aurfaces of all the six- ral, straight walled. Trichomes mostly in- teen arecoid species from a mean of ten ob- tercostal spherical to oval in shape, 9-3 auvations. The description of eophylls was sclerosed basal cells distinct, made from mounted specimens. Material examined: Basu 21 r, 27 I, 29. OBJERVATION A. mcmdp Zipp. ex Blume Arscr trbndnr Roxb. Morphology : Eaphyll broadly bilobed Mo+hology: Eophyll b'dob.d, ]oba so.* about 12.0 cm long, 3:o cm broad at the 13.0 cm long, r .o-I.3 m broad at the mid- middle, prominently 6 nerved, apex obc. i 26 BULLE~N OF THE B&I'ANICAL SVRVEY d~ IND~ [Vol. 20

Abaxial epidermis: Stomata intercostal Adaxial epidermis: Stomata and tricho- in files ; guard cells wide, lateral sub-cells 2, mes absent. Epidermal cells irregular to polars triangular in shape, short, arching on hexagonal, straight walled. the guard cell, stomata a few on the costa Material examined: Basu 201. U withW4lateral subcells, outer subcells double Bentinckia nieobarica (Kurz) Becc. in size of the inner. Intercostal cells trap- Morphology : Eophyll narrowly hilobed, ezoid, straight walled. Costal cells elongated in files, cell walls thick, oblique or straight. lous 14.0-16.0 cm long, I .5 cm broad at Trichomes costal and i,ntercostal, cells scle- the middle apex acuminate and toothed. A baxial epidermis : Stomata intercostal rosed, distinct, costal trichomes much larger not in files ; guard cells with thick outer with multicellular bases. Adaxial epidermis: Stomata and trichom- walls, lateral sub-cells 2, or sometimes more, poIars not distinct. Intercostal cells irregu- es absent. Epidermal tells rhombohedral, lar in shape. Costal cells more or less uni- .Waterials examined: Basu 217, 248. formly rectailpular in files. Trichomes costal Archomtophoenix crurahgbamian~ Wendl . et and intercostal spherical in surface view, Drude cell walls sclerosed. Moliphslogy : ~o~h~ltbilobed, lob& I 7.0 Adaxial epidermis: S.tornata and tricho- I 7.5 m long, I -5-1.6.+cmboad at thq mid- mes absent. Epidermal cells rhombohedra1 dle, prominently pnerved, apex acuminate. and straight walled. A baxial epidermis : Stomata intercostal Material examined : Basu 261. not in files ; lateral subcelln 2, distinct, para- Cap- acumimta Becc. llel to the guard cells; poLars not always Morphology : Eophyll bilobed, lohes 9.5 distinct either spherical tp oval or triangu- cm long, 1.5 cm broad at the middle, promi- lar. 'Jlntercoastal cells irregular to slightly nently 5 nerved, apex obliquely dentate. hexagonal. *&Gal cells*elongated thick wal- A baxial epidermis ; Stomata intercostal led. Tiichomes costal and intercostal ; coshl not in files ; guard cells with sinuous outer trichornes wider at the base, coinpoded of walls, lateral sub-cells 2, rarely 3, polars dis- 6-8 stlerosed cells. 1in.ct. Intercostal cells elongated in files Adaxial epidermis : Stomata and tri- with sinuous walls. Costal cells uniform, chomea absent. Epidermal cells hexagonal, cell walls deeply sinuous. Trichomes in- straight walled. tercostal closely packed, oval in surface Materinls exar,lniwb Basu 266, 210. view, composed of 3 thick walled cells. A. akxandwt (F. Muell.) Wendl. et Drude Adaxial -epidermis : Stomata a few. In- ~orphol@~: Eophyll hilobed ; lobes tercostal cells rhombohedra1 with slightly broadly lanceolate, equal in size, 13.0- sinuous walks. Costal cell rectangular, cell 13.5 .cm long, 1.9 cm broad at the middle walls deeply sinuous. Trichomes on the prominently 6 nerved, apex obtuse. costa a few, cells not differenciated in sur- Ahaxial epidermis: Stomata intercostal, face view. Muterid examined : Basu 272. irregular not in files ; lateral subcells 2, nar- row, yolars ,not always distinct slightly arch- Chryaalidocarpus madagaacariensis Becc. ing on the guard cells ; opposite ends oblique- Morphology: Eophyll bilobed, lobes n& ly or bluntly pointed. Intercostal cells irre- rowly lanceolate, 9.1 cm long, 8 mm broad gular to slightly hexagonal. Costal fells rec- at the middle, prominently 5 nerved, apex tangular straight walled. Trichomes costal acute. and intercos taZ with distinct sclerosed cell Ahxial epide~rris: Stomata intercostal walls* not in files ; lateral sub-cells z, p01ars dis- AND RASW: STt'DfES rN EOPHYLLS OF PAWIS J 9~~1BASU EPIDERMAL ARECOlD 127

fimXf : Fig, S.& :Addl 4-t: 5. ViiW mhIljii 6. Hvii,m+rrlr rtrj~rflrrrtdtm~tn.7. .ibrrr~nsb~nrror~jmmrbyi. 8. ArclWntSphcnir a!txflrrrlrtrr. tin& Intercostal bands wide composed of Matlmirls ~*sriwiinrd: Baal "5 2 I 6. long rectangular cc1b with oblique pani- Cbpdldocarpss mdagmcarhd~ var. hcm tion walls. Trichornes absent. bensi~ Hort. Adaxial epidermis: Stomata intercostal. Morphology : E~phyllbilobed ; lobes I 0.6 widely rparrd, smaller ; guard cells with cm long. i .jcm broad at the middle, pro- sEnuozra outer walls ; lateral subcells 2, pm minently 5 nerved, apex bifid and toothed. lam "distinct, slightly arching on the yard Aha-rinl epidermis: Stomata intercostal cells. Intercostal bands wide composed of irregular ; lateral aubcells 2, po1ars slightly rhombuhcdral to fusifomed cclls in file. amhing on the guard ccIla, oblique at oppo- Costal cells narrowly elonpted. Trich~homen site ends Intemtal cell6 irregular to rham- abscnt. bohedsal, straight walled. Costal cclls elon- 1-28 BULLET& OF THE BOTANICAL SURVEY OF INDIA [Vol. 20 gated in files, shorter wails oblique. Tri- an the guard cells. Intercostal cells rep- chomes absent. larlv rhombohedral, straight walled, Costal Adaxial epidermis : Stomata a few, mostly celfs rectangular, elongated and straight costal, Intercostal cells rhombohedral walled. Trichomes costal, oval shaped in straight walled. Trichomes absent. surface view, cells not distinct. .Materials examined: Basu 22 I, 250 Adaxial epidermis: Stomata a few or ab- sent. Epidermal cells rhombohedral. Tri- Dictymprms album (Bory) Wendl. chomes mostly on the costa, intercostal tri- Mor@ology : Eophyll bilobed, lobes r 2.5 chomes a few. cm long, I .5 cm broad at the middle, pro- Material exemined : Basu 246. minently 5 nerved, apex truncate. Abaxial epidermis : Sromata intercostal Plychbsperma maawthurii (Wendl.) Nichols. not in !3es ; lateral sub-cells 2, nearly ~drpholo~y: Eophyll bilobed ; lobes I e .S double the size of the guard cells. Intercos- cm long, 2.5 cm brdad at the middle, apex tal cells rectangular to hexagonal, cell truncate. walls straight. Costal cells rectangular, Abaxictl epidermis: Stomata in files close elongated in files. Trichomes costal with to the costa; lateral sub-cells 2, much wider multicellular bases, cell walls not distinct. than the guard cells, polars slightly, arching, Adaxial epidermis : Stomata absent. In- not always differentiated . Intercostal cells tercostal cells irregular, cell walls straight. irregular in wide bands. Costal cells elon- Costal cells rectangular. Trichomes absent. gated in files with oblique partiti& walls, Material examined : Basu 236. straight or sometimes sinuous. Trichomes Normanbya nomanbyi (Hill) Bailey oval in surface view, cells sclerosed ; distinct. Morphology : Eophyll bilobed, lobes I 2.5 Adaxial epidermis: Stomata a few near cm long, 2.5 cm broad at the middle promi- the costa. Intercostal cells with deeply nently 5 nerved, apex obliquely dentate. sinuous walls. Costal cells uniformly elon- Abaxial epidermis : Stomata intercostal gated in files, longer walls sinuous. Tricho- in files ; subcells 2, polars distinct slightly mes absent. arching on the guard cells. Intercostal cells Material examined: Basu 245. irregular to fusiformed in shape, cell walls Rh~d,hdeaugustan (Kurz) Moore deeply sinuous. Costal cells rectmgular, Morphology : Eophyll impripinnate, 7.0- elqngated in filgs, narrower than intercostal I 1.0 cm long, leaflets usually 13, terminal cells, walls -deeply sinuous. Trichomes both leaflet bilobed, lobes 5.04.0 cm long, 4.5 costal and intercostal. cm broad. Adaxial epidermis: Stomata a few, guard Abaxt'al epidermis: Stomata intercostal, cells with thick sinuous walls. Intercostal irregular, not in file. Intercostal bands wide, ceUs fusiformed ; longer walls sinuous. Cost- composed of fusiformed cells ; cell walls si- .el cells rectangular in fil* ; longer walls nuous, some central cells in the intercostal deeply sinuous. Trichomes absent. band straight walled. Costal cells rectangu- Material examined: Basu 202, 248. lar to rhombohedral with deeply 'sinuous Hych3netele wdlandismm (F. Muell.) Wendl. longer walls. Trichomes both costal and et Drude intercostal ; cells sclerosed, not distinct. Morphology : Eophyll bilobed, lobes I 2.6 Adaxial epidermis : Stomata absent. In- cm long, 1.5 cm broad at the middle, apex tercostal and costal cells have distinct deeply narrowly truncate. sinuous walls. Trichomes abundant on the Abaxial epidermis : Stomata intercostal costa. in files, aub-cells z, distinct, polars arching material examined : Basu ,247. DASU: IN OF t978I HASU AND EPIDERMAL SIWDlES EOPHYLLS ARECOlD PALMS 12g

matt-I 1I : Fig. 9-1 1. Abaxial epidermis : 9. Nmnn~lbynnort~~mrbi. 1 Q. Awca triandra. I I. Bwho$wmn trmnrthun?.

Raystoms borinquca~Cook Adaxial epidermis: Stomata intercostal, Morplaology : EophylP, solitary, entire, polar ceIls indistinct hut arching on the linear-lanctolate, 21.3 crn Iong, r.1 cm guard cells. Costal and intercostal cella dis broad at the middle ; midnerve prominent, tinctly elongated. Trichornes have multi- apex narrowly truncate. cellular bases. Abaxiol epidermis : Stomata intercostal, -Matariizl examined : Basu 253. widely spaced, in files ; lateral sukclls 2, Veftchh mcMUi Pecc.) Moore wider than the guard cells, polars larger, in- Morphology : Eophyll bilobed, lobes tqual distinct but arching On the guard cells. In- in Iength, r 2.913.0 cm long, 2.2 crn broad tercostal cells mu ch elongated, narrow with at the middle, apex truncate. oblique partition walls. Costal cells reaang- Abaxial epidermis: Stamata intercostal ular and straight walled. Trichornes absent. not in files ; lateral aukella 2, deeply cum- Adaxial epidrrmis: Stomata a few. Epi- 4, polar cell^ small, oval, slightly arching dermal cells rectangular, longitudinally ex- on the guard cells. Intercostal cells rectang- tended. Trichomes absent or a few. ular to rhombohedra1 or fusiforrned, longer Matekls eramined : Basu 213, 249. walls sinuous. Costal cells rectangular in files, ell walls dcepl y sinuous. ~hchomen Roy&oaer regia (H.B.K.) Cook only on the cmta. iWo7phology : Eophyll solitary entire, li- Adaxial epidermis : Stomata absent or near-lanccolare, 22.0 crn long, 1.3 crn broad at the middle, apex arnrminate. a few near the costal bands. Epidermal A haxbl epidermis : Stomata intercoaal cetls rectangular to rhombohedral, cell walls deeply sinuous. Trichomes absent, in files ; lateral subcells wider than the Matertkl examined: Basu zog. pard cdls, polars sometimes triangular, arching on the pard cells. Stomata with STOMATAL FREQUENCIES AND MEASUREMENT a common polar cell frequent. Intercostal OF GUARD CELLS cell irregularly elongated. Costal cells in Stomata1 frequencies from I mm sqwre files with ohliqec prrtition wal1s. Trichom- of the adaxial and abaxial epidermis of es absent eophylls of sixrem arecoid species were =al- Is BULLETIN OF THE BOTANICAL SURVEY OF INDU [Vol. 29

culated with a lens combination of 15x 20, guard cells shows that Dactyosperma a!burn the observations were recorded in table-I. has the smallest guard cells and Rhopalo- The frequency of stomata on the abaxial blaste augusta has the largest. The cutinis- side ranges from 15 to 25 per unit area ex- ed ledges (Tornlinson 1961) are prominent cept in Roystonea borinquena where num- in guard cells. ber of stomata was 9.2 only. The length of

TABLE I - Stomata1 Stomata1 Epidermal cells Trichomes Guard ceNs Sl. indices frequency in micron in micron in micron No. Name - Adaxial Abaxial Adaxial Abaxial Length Width Length Width Length Width

I 2 3(a) 3(b) 4(a> 403) 5(a) 503) 6(a) 6(b) 7(a) 703) -* - + 1. Ar~catriandra 1.8 4.0 1.6 15.62 61.2 15.7 48.3 46.8 19.5 5.2 2. A. macrocalyx Nil 3.5 Nil 15.6 63.7 10.5 37.4 33.8 19.2 5.0 3. Dictyospm album Nil 5.4 Nil 22.5 32.5 10.2 109.2 37.4 18.6 3.2 4, Nwpaloblaste auguskr Nil 5.0 Nil 16.25 52.5 12.8 Nil Nil 30.2 5.2 5. Roystoneu regia 9.0 8.2 6.25 25.0 58.8 11.2 67.6 52.0 26.3 4.2 6. R. boringu~na 0.05 3.4 3.3 9.3 56.7 11.5 Nil Nil 21.2 5.2 7. Car@entariaacuminatu 2.8 8.1 1.1 15.9 62.7 16.2 57.2 31.2 26.8 5.0 8. Normanbya nonnanbyi 0.7 7.7 1.9 22.5 69.8 15.7 67.6 37.4 22.2 5.2 '9. Archontophoenix alwcandrae Nil 5.2 Nil 15.2 40.7 20.5 11.0 10.4 21.5 3.0 10. A. cunninghamiana Nil 5.2 Nil 18.7 36.5 11.6 37.4 11.0 21.6 5.2 11. Ptychospmna macarthurii 1.4 8.6 1.4 26.8 58.5 11.7 62.4 37.4 22.6 5.0 12. Bmfinckia nicobarica 1.7 7.8 6.2 28.1 44.3 15.4 22.4 11.4 26.6 5.5 13. Hydnirstde w~ndlandiana 0.9 3.6 3.7 12.5 76.0 10.4 71.0 47.2 25.7 5.2 14. Vcitchia mGnillii Nil 5.2 Nil 15.6 53.5 11.5 22.4 15.5 23.4 5.1 15. Cluyalidocarpus madogascariensis 2.4 7.3 5.2 18.7 47.8 18.2 Nil Nil 26.0 4.6 --16. C. madagascarimsis var. hcubensis 2.7 11.5 10.5 21.0 58.2 15.6 Nil Nil 25.5 5.2 The stomatal index from the abaxial epi- found in their adult leaves. Areca macro- dermis of arecoid palms shows a range of calyx, Norman by a norman byi, Pty chospsr- 3-5 to 1 I .S and Areca macrocalyx, Royston- ma macarthurii, Hydriastele wendlandiana, ea borinquena, Hydriastele wendlandiana, Roystonlea borinquenn and R. regiu) how- Areca triandra, have relatively low index ever, show stomata in files. Tomlinson (1961) number. The stomatal indices were in the reported that in arecoid genera Archonto- higher range found int Chrysalidocarpus phoenix, Dictyosperma and a few others, madagascariensis var. lucu bensis, Ptychos- stomatal arrangement is in files in the perma macarthurii. Roystonea regia and adult leaves. In our observations an irre- Carpentaria acumdnata. Stomata absent in gular arrangement of stomata was noted in the adaxial side of six arecoid species, the the eophylls of these two species. Ghose rest shows a comparatively low stornatal in- and Davis (1973) have pointed out from dex number. their comparative studies in the juvenile DISCUSSION and adult leaves of eight palm species that Observations from the adaxial and abaxial the number of stomata per unit area increas- epidermis of sixteen arecoid palms show the es manifold in the adult leaves (exception tetracytic type of stomata and a majority only in Hyphnene thebaica). The subsidi- of them are irregularly arranged in the in- ary cells are basically four in the stomata tercostal region. The concentration of stoma- of arecoid palms, the lateral subsidiary cells tp is higher in the abaxial side which is in having walls of various thickness, the polar conformity with the stomatal arrangement cells, two in numbers being not always dis- 39781 BASU AND BASU: EPI~EICMAL~f'lfbx~s IN EO- dp NU?COID PAWS 131 tinct hmthe adjoining epidermal cells but Wc~shingtoniuand others. Their om- variously arching on the guard cells. Steb- tion in juvenile leaf of Phok shows an bin and Khus (1961)observed 2, 4 and 6 increased number of subsidiary cells. Ghose. subidiary cells in palms. According to and Davis (1g74), however, did not find them, four subsidiary cells having two lat- change in the number of subsidiary cells in era!s and two roundish polar cds are the juvenile and adult leaves in Phomh characteristic of the adult leaves of Phomix, sylvestrir. i 32 BULLETIN OF THE BOTANICAL SURVEY OF INDIA [Vol. 20

The adaxial and abaxial epidermal cells equally of significance in the of from the eophylls show two distinct cell- the palms. wall types. The sinuous condition is mostly restricted to the anticlinal walls and found ACKNOWLEDGEMENTS in Carpentaria acuminata, Normanbya nor- We express our thanks to Dr. T. A. Davis, manbyi and Rhopalobalaste augusta. In Coconut Agronomist, F.A. O., L.P.T.I., Pty chosperrma macarthurii and Veitchia Jalan Siswa Barat, North Sulawesi, Indone- merm'Ilii, only the adaxial epidermal cells sia ; Dr. N. M. Nayar, Joint Director, Cen- show sinuous anticlinal walls. These sinu- tral Plantation Crops Research Institute, ous epidermal cells are a characteristic of Vittal, South Kanara, Karnataka and the the Lepidocaryoid, Coryphoid and Borassoid Director, Botanical Gardens, Singapore for palms and to some extent of the Phoenicoid supply of arecoid palm seeds. We are also palms. The si~iuousnature is due to cuti- grateful to the Deputy Director, Indian cular deposits on the anticlinal walls (Torn- Botanic Garden, Howrah for providing faci- linson 1961). Anlong the straight-walled lities in the department and to Shri C. C. types both Roystonea regia and R. borin- Mukherji for solving photographic prob- querta have the characteristic long narrow lems. We express our gratitude to the epidermal cells which isolate this genus in Director, Botanical Survey of India for his the arecoid group. On the basis of the na- encouragement on palm studies. ture of the adaxial epidermal cells the palms studied can be dis;nguished into the fol- REFERENCES lowing categories : BASU,S. K. AND D. P. MUKHERJI.Studies on the germi- nation of plam seeds. PrinciH 16(4) :136-137. 1972. A. Adaxial epidermal cells rect- Rhopaloblaste augusta, BAVAPPA,K, V. A. Morphological and anatomical angular, longitudinally extended Ptychos@rma macar- studies in Areca catechu Linn. and A. triandra Roxb. with sinuous anticlinal walls. thurii, Vcitchia medlii. Phytomorphology 16 :436-4343, 1966. B. Adaxial epidermal cells rhom- Carpantaria acumi- COOK,0. F. Juvenile characters of Royal palms. Science bohedral, anticlinal walls sinu- nata, Normanbya nor- 81 : 590. 1935. 011s. manbyi. EAMES,A. J. Neglected morphology of the palm leaf. Phytomorphology 3 : 172-289. 1953. C. Adaxial epidermal cel!s rhom- Bentinckia nicobarica, GHOSE,M. AND T. A. DAVIS. Stomata and trichome bohedral, straight walled. Ch~ysalidocar~trsmada- in leaves of young and adult palms. Ibid. 23(3-4) : gascariensis var. lucu- 216-229. 1973. bensis, Areca trhndra: -- Ontogeny and structure of stomata in Cocosnuci- A. nracrocalyx, Dictyo- fera L. J. Indian bot. Soc. 53 : 19-32. 1974. sperma album. Hydria- MOORE,Jr. H. E. Synopsis of various genera of Arecoi- stele wendlandiana. deae. Gentes Herb. 8 :481-536, 541-576. 1957. D. Adaxial epidermal cells hexago- Archontophoenix. - The major group of palms and their distribution. nal, straight-walled. Ibid. 11 (2) : 27-141. 1973. E. Adaxial epidermal cells rectan- Roystonea regia, STEBBIN,G. L. AND G. S. KRUS.Variation in the organi- gular, longitudinally extended. R. horinquena. sation of the stomata1 complex in the leaf epidermis of and 1ts bearing on the phylogeny. Amer. J. bot. 48 : 51-59. 1961. Eophyll characters have long been recog- TOMLINSON,P. B. Seedling leaves in palms and their morphological significance. J. Arnold Arbor. 41 : nised in the diagnosis of palm taxa (Cook, 414-427. 1960. 1935; Eames, 1953 ; Moore, 1957, 1973 ; Wes- - Anatomy of the monocotyledons 11 Palms. Oxford 1961. sel Boer, 1965). As shown above, the epi- WESSEL BOER, J. G. Indigenus Palms of Srrinam. dermal characters of the eophylls seem to be Leiden. 1965.