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Pollination Ecology of <Emphasis Type Proc. Indian Acad. Sci. (Plant Sci.), Vol. 100, No. 3, June 1990, pp. 195-204. Printed in India. Poilination ecology of AlangŸ lamarkii (Alangiaceae) T BYRAGI REDDY and CH. ARUNA Department of Environmental Sciences, Andhra University, Waltair 500 003, India MS received 70ctober 1989 Abstract. Alan#ium lamarkii flowers during February-April. Buds are opening around the clock and offer pollen and nectar as the reward to their insect visitors. The breeding system incorporates both geitonogamy and xenogamy. Flowers are large, hermaphrodite, essential organs are centrally situated, stigma Ionger than anthers. Nectar contains glucose, sucrose and fructose. The glucose being dominant. Sugar concentration ranges from 25-29%. Protein and amino acids were present. Pollen-ovule ratio is 27300: 1. Altogether 18 insect species were found foraging at the flowers. Of the 18 species of insect foragers bees (Apis florea, Trigona, Amegilla, Ceratina, X ylocopa latipes, X ylocopa pubescens) and wasps (Delta sp., Rhynchium) promote both geitonogamy and xenogamy. The bees collected pollen as well as nectar. The wasps while foraging for nectar their dorsal side touches the anthers and causes nototribic pollination. The butterflies visit the flowers for nectar only, but the contact between proboscis and the essential organs is unlikely. Keywords. Pollination; geitonogamy; xenogamy; nototriby; nectar; Alangium lamarkii. 1. Introduction Pollination ecology has entered into a new phase of synthesis and correlation. A plant species in bloom needs more than one insect species for its pollination purposes. Likewise, an animal requires a series of plant species in bloom to provide continuous nourishment, then it becomes necessary as pointed out by Baker (1963) to consider the pollination relationships on a community basis. Studies of Baker (1970, 1973), Baker et al (1971), Frankie et al (1974) and Macior (1974a, b) reinforced the need for synecological study of pollination relationships. Heithaus (1974) and Frankie (1976) showed how plant-pollinator interactions are crucial in determining community structure and its functioning. The available information on pollination ecology is based mostly on the studies carried out in the temperate areas of Europe and north America. Only recently studies have been undertaken in the tropics (see Percival 1965; Proctor and Yeo 1972; Janzen 1975; Faegri and Pijl 1979). These studies pointed out the need for a synecological approach in the study of pollination (Baker et al 1971; Frankie and Baker 1974). In India, although some information on pollination of some tropical plants is available (Kapil 1970; Deodikar and Suryanarayana 1977; Kapil and Jain 1980; Mahrotra et al 1983; Reddi 1987) ir is highly deplorable that even basic data on the pollination of any of the varieties of the plant species occurring is totally lacking (Mohan Ram 1980). Information on pollination of wild plants is undesirably meagre. Only recently there have been studies initiated and completed (Subba Reddi et al 1979, 1981, 1983; Ananthakrishnan et al 1981; Gopinathan et al 1981; Pant et al 1982, 1983; Subba Reddi and Reddi 1982, 1984; Reddi and Subba Reddi 1983, 1984, 1985; Birbahadur aad Ramaswamy 1984; Mathur and Mohan Ram 1986; Panth and Chaturvedi 195 196 T Byragi Reddy and Ch. Aruna 1986; Velayudhan and Annadurai 1986; Meera Bai 1987; Soloman Raju 1987; Byragi Reddy 1988; Rama Devi et al 1989; Jyothi et al 1990). The need to understand some interactions, especially in the species rich tropical ecosystem(s) is outstanding. The present study describe the interaction of 18 insect species with the flowers of Alan#ium lamarkii (Alangiaceae), a large tropical tree and also a medicinal plant. 2. Materials and methods A. lamarkii growing at Visakhapatnam (17~ and 82~ was used for observations. Pollen output per anther was assessed by counting aU the pollen grains in a sample obtained by gently crushing and tapping the anther on a clean microscope slide spreading the pollen mass uniformly. The longevity of pollen and stigma was based on the fruit set success from hand-pollinations at regular intervals. Nectar produced in flowers, protected from insects by butter paper bags for 2 h, was measured using disposable micropipettes. Refractometer was used to determine nectar sugar concentration. Paper chromatography was used to determine nectar sugar composition (Horborne 1973). Proteins and amino acids were identified by the method of Baker and Baker (1973). The flowers tO be hand- pollinated were emasculated p¡ to anthesis and then bagged. Test for apomixis/ autogamy, geitonogamy/xenogamy were conducted through controlled pollinations. Apomixis was tested by bagging the emasculated flowers free of pollen, autogamy by pollinating flowers with the pollen of the same flowers. Geitonogamy by pollinating flowers with the pollen of the different flowers of conspecific plant, for xenogamy with the pollen of the different conspecific plant. Insect visitors collected during the study period at all 3 study sites (LIC quarters, Pedagadili and Simhachalam) were identified through the courtesy of Commonwealth Institute of Entomology, London and Zoological Survey of India, Calcutta. Butterflies were identified after Wynter-Blyth (1957) and their nomenclature used is that of Varshney (1983). The behaviour of visitors, the length of a visit and number of flower visits in a unit time were carefully recorded. The more frequent visitors were caught and their bodies were examined under a stereomicroscope for the pollen adhering to body arcas and then washed off with alcohol. To assess pollen amounts transferred on to the stigma in a single visit by a particular kind of insect, bagged flowers just before anthesis were opened one by one for the insects to visit. When such exposed flowers received the first visit, their stigmas were examined for pollen. Similarly the pollen deposited on the stigmas was assessed at regular intervals. 3. Resuits The plants begin to bloom soon after the cold season. The blooming season extends from February to April every year. Flowers are white or yellowish white, hermaphrodite in axillary fascicles bearing 2-3 flowers. The root bark is anthelmintic and purgative. It is useful for fevers and skin diseases, and is generally administered in the forro of powder. Pollination ecology of A. lamarkii 197 3.1 Phenology of anthesis Opened flowers are evident throughout the day and night with a higher frequency during 0500-1000 h. A bud takes 20-30 min to become fully open. Concomitant with the gradual opening of flower the process of nector secretion is also started. Petals covered the stamens and stigrna, but are yellowish at the time of anthesis. The petals were excited by this time and are consequently reflexed (deflexed) downwards asa sudden mechanism. They, then expose the anthers and stigmas which happens to be longer than the former to the visitors. 3.2 Flower morphology Flowers are considerably large and hermaphrodite. Calyx tube is adnate to the ovary; limb is truncate or 4--10 toothed. Petals 4--10, linear, light green, polypetalous, valvate, thickened and recurved in flower. Flower length ranges from 1.6-2.2 cm (av. 1.8) and is 0-5 mm wide, stamens ranging from 20-30. The anthers are dithecous and introrse. Stigma is large, capitate, projecting beyond the anthers. Ovules are solitary and pendulous. 3.3 Pollen characters Anthers open immediately after anthesis. Pollen grains are freed through longitudinal dehiscence. The number of pollen grains range from 21880-32820 (x=27300) per flower. The pollen grains are spherical and small i.e., 15/~m. Exine has sculptured ornamentation, cytoplasm is granular, 2-5 colporate. Pollen grains remain viable for 40 h from the time of anther dehiscence. Pollen ovule ratio came to 27300: 1. 3.4 Stigma receptivity Stigma remains receptive for 36 h beginning with anthesis. On hand-pollination 0, 6, 12, 24 and 36 h old stigrnas gave 80, 70, 60, 40 and 30% fruit set respectively and still older stigmas were not receptive. 3.5 Nectar dynamics Nectar is situated around the ovary base. Nectar volumes measured at 2 h intervals indicating that the rate of production is not consistent throughout flower life. Sugar concentration ranges from 25-29%. The sugars present were glucose, sucrose and fructose, the glucose being dominant. Proteins and amino acids too were present. The score on histidine scale was 3. 3.6 Flower visitor activity dynamics 3.6a Composition abundance: Du¡ the study pe¡ 18 insect species were found foraging at the flowers (table 1). Of these, 12 are Hymenoptera (3 Apidae; 4 198 T Byragi Red@ and Ch. Aruna Table 1. Particulars of flower-visitors on A. lamarkii. Forage type Body region of poUen Visitor species Poilen Nectar deposition Hymenoptera Apidae Apis cerana indica + + Head ventral side A. florea + + ~~ Trigona sp. + + Anthophoridae Amegilla sp. + + ~~ Ceratina sp. + + ~~ Thyreus histrio + + Pithitis bingami + + --~ X ylocopidae Xylocopa latipes + + Head dorsal ventral side X. pubescens + + Eumenidae Delta sp. - + Dorsal side head Ropalidia spatulata - + Rhynchium metallicum - + --~ Diptera Muscidae Musca sp. - + Ventral sidr Lepidoptera Sphin#idae Macroglossum gyrans - + Proboscis legs Danaidae Danaus chrysippus - + --~ Euploea core - + ~~ Pieridae Catopsilia pyranthe - + ~~ Hesperiidae Pelopidas mathias - + Anthophoridae, 2 Xylocopidae, 3 Eumenidae), 1 Diptera and 5 Lepidoptera (1 Sphingidae, 2 Danaidae, 1 Pieridae and 1 Hesperidae). Of the 18 visitor species, 11 were encountered at all the 3 study sites. These species include A. florea, Trigona sp., Amegilla sp., Ceratina sp., Xylocopa latipes, X. pubescens, Delta sp., Rhynchium metallicum, Musca sp., C. pyranthe and Pelopidas mathias. The total number of species caught at each of the study sites was 14. The number of visits made by different insect species at 3 study sites are given in table 2. On the whole, bees frequented most and shared 77% of the total visits followed by wasps (12-6%) and butterflies (7%). Among each group of insects, the visits of individual species varied numerically from site to site (figure 1). Among the br visits Xylocopa latipes shared 24-3% of total bee visits, foilowed by X.
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