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Floral Morphology of a Few Species of Euphorbiaceae by N

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Floral Morphology of a Few Species of Euphorbiaceae by N FLORAL MORPHOLOGY OF A FEW SPECIES OF EUPHORBIACEAE BY N. C. NAIR AND V. ABRAHAM (Botanical Survey of India, Dehra Dun and Botanical Survey of India, Madan Street, Calcutta) Received April 23, 1962 (Communicated by Dr. V. Purl, r.A.sc.) INTRODUCTION OUR knowledge on the floral anatomy of Euphorbiace~e is inadequate. Haber (1925) and SchoiJte (1937) have described the morphology of cya- thium. Saunders (1939) gave an account of the floral anatomy of a few species of Euphorbia, Spurge mercurialis and Ricinus communis. The present account deals with Jatropha curcas Linn., J. gossypifolia Linn., Croton bonplandianum Baill., Codiceum variegatum Blume., Hevea braziliensis Muell Arg. and Breynia rhamnoides MueU Arg. The material for the present study was collected from Changanacherry, Kerala State and fixed in formalin-acetic acid-alcohol. Customary methods of dehydration and imbedding were followed. Sections cut at 8-12tz were stained with safranin fast green. OBSERVATIONS Jatropha.--The flowers of J. gossypifolia are provided with glandular as well as unicellular hairs (Fig. 1, R, S) on the pedicel and outer surface of the calyx. These are absent in J. curcas. The gamosepalous calyx is campanulate having valvate and persistent sepals (Fig. 1, C, E) whose margins are toothed and glandular in J. gossypifolia and entire in J. curcas. The petals are slightly united at the base (Fig. 1, D) and show twisted aestivation (Fig. 1, H). Within the perianth whorl and alternating with the petals are five nectariferous glands composed of dark staining parenchyma (Fig. 1, D). These are very prominent in the female flower. While the male flower of J. curcas has ten stamens in two whorls of five each that in J. gossypifolia has eight stamens in two whorls of five and three respectively (Fig. 1, F-H). The inner stamens are longer than the outer. All of them are united by the bases of their filaments to form a staminal column (Fig. 1, E). The 1 BI 2 N.C. NAIR AND V. ABRAHAM lobes of the dithecous and extrorse anthers are divaricate and the connective protrudes slightly above the level of anthers. The anther wall consists of epidermis, fibrous endothecium, two middle layers and a secretory two- to four-nucleate tapetum. There are only very few female flowers in an inflorescence. Generally the central flower of a dichasium is female. There are two whorls of five minute staminodes each at the base of the ovary. Sometimes they develop into functional stamens whereby the flower becomes bisexual. The pedicel has a siphonostele. Each sepal trace while leaving the receptacle divides into three (Fig. 1, A). In the higher regions of the sepal there are five to seven vascular bundles. Each of the second whorl of five traces before entering the petal gives a trace on either side which ramifies in the alternipetalous glands (Fig. 1, B, C). A petal in transection shows five to nine bundles. The remaining stele breaks into two rings of vascular bundles. The outer whorl consists of five bundles in both the species while the inner consists of three in J. gossypifolia and five in J. curcas. They continue in the staminal column and supply their respective whorls of stamens (Fig. 1, E-G). Each staminal trace in the region of the anther divides into three branches (Fig. 1, G, H). While each lateral branch supplies a theca, the median branch fades out as the tip of the connective is approached. After the departure of the petal traces, two whorls of five traces each depart from the siphonostele of the female flower to supply the staminodes (Fig. 1, I). Above this level the stele is triangular. From each of the angles there diverges out a trace that forms the dorsal trace of a carpel (Fig. 1, J, K). Each dorsal bundle branches profusely in its upward course through the ovary wall (Fig. 1, L-O). The remaining stele breaks up into three strands and the locules of the ovary appear at this level (Fig. 1, K). These bundles, which are normally oriented, take their place in the septa (Fig. 1, L, M). Each carpel bears an anatropous bitegmic and crassinucellate ovule. As in many Euphorbiace~e there is a well-developed obturator (Fig. 1, P) which in J. gossypifolia is more prominent than in J. curcas. As the ovular region is approached each of the septal strands becomes con- centric and ultimately breaks up into two inversely oriented ventral bundles from each of which is given off an ovular trace (Fig. 1, N). Traces from ventrals of the same carpel supply an ovule. Thus each ovule receives two traces which fuse after entering the funicle in J. gossypifolia and before entering the funicle in J. curcas. During post-fertilization stages the funi- cular strand divides at the chalazal end of the ovule into a large number of bundles which traverse the inner integument (Fig. 1, M)almost to the tip. Floral Morphology of a Few Species of Euphorbiaceae . LL>~:J (' J . ~ - . ,t ,~( ~do d~ ~,. 8t J J 14 oh..- @ "5mm7 Ftao I 4 N.C. NAIR AND V. ABRAHAM Similar cases are known in other members of the family, e.g., Riclnus corn~ munis (Singh, 1954). The ventral bundles fuse and fade out as the top of the ovary is approached. The dorsal bundles alone continue up to the tip of the reflexed and glandular stigma (Fig. 1, O-Q). Breynia.--The short pedicelled male flower has a gamophyllous fleshy perianth which breaks up into six parts above. These are arranged in two whorls of three valvate tepals each (Fig. 2, I-K). The stamens are united to form a column bearing usually three and rarely five dithecous extrorse anthers (Fig. 2, G-I). The column continues above into a sterile three- lobed region which ultimately breaks up into three parts. The anther wall is similar to Jatropha. The hypodermal cells toward the sterile tip of the column bears fibrous thickenings similar to those found in the endothecium of the anthers. In the lower region of the pedicel there are two collateral vascular bundles which divide repeatedly to form eight to nine bundles (Fig. 2, A, B) and enter the triangular receptacle. One bundle diverges towards each angle of the receptacle (Fig. 2, C), followed by three more traces on alter- nating radii (Fig. 2, C, D). They constitute the supply of the perianth and each of them divides into three branches before the perianth gets split up (Fig. 2, H, I). Three bundles are left in the centre which become concentric (Fig. 2, H) and extend up to the middle of the staminal column (Fig. 2, F, G). After the perianth supply has separated, the receptacle of the female flower is again triangular. A trace diverges out towards each angle to become the dorsal bundle of the carpels (Fig. 2, L-P). In their upward course they give a few branches to feed the ovary wall. The central stele breaks up into six bundles which get arranged in three groups of two each on septal radii (Fig. 2, M). The locules appear at this level. At first these bundles are normally oriented but become inversely oriented at about the middle of the ovary. The trilocular ovary contains two ovules in each loculus. Each of the inversely oriented bundles after giving out a trace to an ovule on the corresponding side (Fig. 2, N) continues upward and fades out as the stylar portion is approached (Fig. 2, O, P). The dorsal bundles alone continue up to the stigma. Hevea.--The flowers are arranged in loose panicles. They have each a thick and hairy gamophyllous perianth which breaks up into five valvate tepals above. Within the perianth, in the female flower, there are five- minute staminodes in radii alternating with the tepals. In the male flower there are ten stamens, in two whorls of five each, united by their filaments Floral Morphology of a Few Species of Euphorbiaceae 5 e • ,~e ~ ~ ~'~ ¢~;, IIfI~,fl "', \ E / --- r, N "Smm. \ 0 \ ~ " / R FIO. 6 N. C. NAIR AND V. ABRAHAM to form a staminal column. The wall of the anther is similar to that of Jatropha. The siphonostele of the pedicel expands in the receptacle and ten traces diverge out in two whorls of five each (Fig. 2, Q). Those which arise first are the midrib traces of the tepals while the rest are commissural traces. The latter fork into two each to form the marginal bundles of the tepals (Fig. 2, Q, P, W). Of the next two whorls of five traces each, the antipetalous ones in the female flower fade out in the cortex while the inner whorl enters the staminodes (Fig. 2, Q-S). The stele becomes triangular and gives rise to three dorsal traces (Fig. 2, R). The remaining stele breaks up into six bundles which become inversely oriented and fuse in pairs in the septal radii (Fig. 2, S, T). Further behaviour of these bundles as well as the dorsals is similar to that of Jatropha (Fig. 2, T-W). In the male flower the central stele organises into two rings of five bundles each above the level of perianth. These continue in the staminal column and each bundle ultimately becomes the supply of a stamen (Fig. 2, X, Y). Croton.--The racemose inflorescence bears three to four female flowers in the lower nodes while in the upper nodes only male flowers are generally present. The inflorescence axis as well as pedicel and outer surface of the calyx lobes bears trichomes of various kinds (Fig. 3, A-C).
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