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Distribution and Pollen Morphology of Some Species of Grewia Linn

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Distribution and Pollen Morphology of Some Species of Grewia Linn DISTRIBUTION AND POLLEN MORPHOLOGY OF SOME SPECIES OF GREWIA LINN. IN NIGERIA. 1*Shokefun, E. O., Orijemie, E. A.2 and Ayodele, A. E.3 1Department of Botany, University of Ibadan, Ibadan, Nigeria ABSTRACT The distribution and pollen morphology of some Grewia species in Nigeria were studied. These species occupy a wide range of habitats; six species (G. malacocarpa Mast., G. barombiensis K.Schum., G. brunnea K.Schum., G. coriacea Mast., G. hookerana Exell & Mendonca. and G. oligoneura Sprague) are found in the lowland rain forests of the Southern parts of the country, eight species (G. barteri Burret, G. bicolor Juss., G. cissoides Hutch&Dalz., G. flavescens Juss., G. lasiodiscus K.Schum, G. venusta Fresen., G. tenax (Forsk) Fior and G. villosa Willd) were present in the Guinnea savannah dry land of the Northern part of the country while two species (G. carpinifolia Juss. and G. mollis Juss.) occur in both ecological zones in the country. Grewia bicolor Juss. is the only species restricted to the high mountains of the Northern part of the country among the Savannah species. The pollen results reveal two pollen types: the Microcos and Grewia types. The Microcos pollen type consists of the following characters: tricolporate with short colpi, small-sized pollen of 17.5µm-21.0µm x 15.0µm-20.0µm (PA x ED); sub-prolate to prolate shape; exine is thin (0.5 µm -1.50µm) and fine reticulations; lumina shape range from round, angular and elongated. Parent plants with this pollen type are natural to the lowland rainforest zone. The Grewia type consists of pollen, parent plants of which are natural to southern Guinea savannah, as well as G. mollis and G. carpinifolia which occur in both ecological zones. The pollen grains of the Grewia type are mainly tricolporate but G. venusta is three to four colporate; they possess long colpi; are medium (30.0µm) to large (55-67.5µm x 57.5-82.5µm) in size. Pollen shapes include sub-prolate, prolate and oblate-spheroidal; exine is generally thick (0.87µm-3.0µm), exine pattern is reticulate, coarse reticulate and sometimes striato-reticulate; lumina shapes are distinctly polygonal or irregular, and contain bacules. These characters are discussed in relation to the taxonomy of the genus in Nigeria. Keywords: Grewia, Distribution, Palynology, Nigeria. INTRODUCTION Grewia Lin. (Tiliaceae) consists of about 280-300 species well represented in Tropical Africa, Asia and Australia (Cronquist, 1981; Chung et al., 2003). In West Africa, 17 species occur; sixteen of these are recorded for Nigeria (Hutchinson and Dalziel, 1954). Nigeria is a major centre of diversity for Grewia in West Tropical Africa (Czarnecka et al., 2006). Six species are known to occur in lowland rainforest, eight in dry woodland savannah while two species occupy both ecological zones (Hutchinson and Dalziel, 1954). The lowland rainforest species are distinguished by presence of panicle inflorescence while the savannah species are distinguished by the presence of cymose inflorescence (Burret, 1926; Hutchinson and Dalziel, 1954; Bayer and Kutbitzki, 2003; Cowie et al., 2011). Taxonomic studies of the genus have been carried out by many authors using morphological, anatomical, palynological and molecular data (Burret, 1926; Chattaway, 1934; Cronquist, 1981; Chang and Miau, 1989; Watson and Dallwitz, 1992; Alverson et al., 1997; Thorne,1998; Bayer et al., 1999; Judd et al., 2000; Bayer and Kutbitzki, 2003; Chung et al., 2003). In particular, morphological data have provided taxonomic information at family, subfamily, generic, subgeneric and species level for different groups of plants (Stuessy, 1990). Perveen et al. (2004), highlighted the morphological features of the pollen grains of Grewia in the study of the family Tiliaceae as follow: 3 colporate aperture; rugulate/finely reticulate to coarse reticulate tectum and prolate or subprolate pollen shapes. El-Husseini (2006) reported the importance of the colpi ends, pollen size, muri ridge shape, equatorial view and size of lumina and stated that spheroidal pollen grains are peculiar to the Grewia type. Perveen and Qaiser (2007) recognized two pollen types: Corchorus depressus type and Corchorus tridens type in their study of the subfamily Grewioideae. Pollen data have been used effectively at generic and subgeneric or sectional levels on numerous occasions within tribes (Vezey et al., 1994). Although considerable works have been done on the species of the genus, data on Nigerian species are sparse. The aim of this study was to investigate the ecological distribution and pollen morphology of some species of Grewia in Nigeria with a view to providing additional information for the delimitation of the species. The results would provide a better understanding of the relationship between and among the species in Nigeria. MATERIALS AND METHODS Specimens of Grewia species were collected during field trips to different locations within Nigeria. Flower buds and floral specimens were preserved in 50% ethanol. Voucher specimens were made for all collections using standard herbarium procedures and deposited in the Herbarium of the Department of Botany, University of Ibadan, Ibadan, Nigeria (UIH). The various locations visited are shown in Figure 1. The herbarium specimens were studied for their distributional data in the Forest Herbarium Ibadan (FHI), Ibadan, and the University of Ibadan Herbarium (UIH). DISTRIBUTION AND ECOLOGY Data obtained during the study of specimens in the herbarium and notes taken during field trips provided useful information for the distributional ecology of the genus in Nigeria. Pollen Morphology: Flower and floral buds preserved in 50% ethanol from the field collections and in the case of species that were not collected from the field, flower and floral buds from well identified herbarium specimens were assessed for pollen morphology using acetolysis method according to the procedures described by Erdtman (1952, 1960) and Ayodele (2005). The floral buds were crushed with a glass rod in centrifuge tube. Three millilitre of freshly prepared acetolysis mixture (9 parts acetic anhydride to 1 part concentrated Tetraoxosulphate VI acid) was added to the content in the tubes. The content was heated in a water bath from 700C to boiling point, stirring occasionally. The centrifuge tubes and content were left in boiling water for 3 minutes and then centrifuged at 4,000 r.p.m. for 5 minutes while still hot. The supernatant was decanted into Acetolysis waste bottle. Some water was then added to the sediments in the tubes and shaken vigorously using a whirl mixer. Few drops of methylated spirit were added to remove the foam formed and centrifuged again. The supernatant was decanted. The washing with water and centrifuging were repeated four times. Fifty per cent glycerine was added and left to stand for two hours. The tubes were shaken vigorously using a whirl mixer and centrifuged at 4,000 r.p.m. for 10 minutes. The supernatant was decanted off finally and the tube was inverted over filter paper and left overnight. One hundred per cent glycerol was added to the tubes and shaken. This was then poured into labeled storage vials. The pollen grains were mounted in unstained glycerin jelly and observations were made with a Fisher scientific illumination microscope at (E 40; 0.65) and oil immersion (E 100; 1.25) using 10x eye piece. The measurement was based on 20 readings from each specimen. Photomicrographs were taken using Leica CME with Digital Microscope Eyepiece attachment and Photo Explorer 8.0 SE Basic software. Terminologies used were in accordance with Erdtman (1952), Moore et al. (1991) and Perveen et al. (2004). All slides are deposited in the Herbarium of the Department of Botany, University of Ibadan, Ibadan, Nigeria. RESULTS Distribution and Ecology Nigerian lowland rainforests and Guinea Savannah provided suitable habitats for the sixteen species in the genus Grewia. Six species, G. malacocarpa, G. barombiensis, G. brunnea, G. coriacea, G. hookerana and G. oligoneura are confined to the high rainforest of the Southern part of Nigeria while eight species (G. barteri, G. bicolor, G. cissoides, G. flavescens, G. lasiodiscus, G. venusta, G. tenax and G. villosa) are found in the Guinea Savannah in the Northern part of Nigeria. Two species (G. carpinifolia and G. mollis) occupy both ecological zones (Fig.1). Some of the Nigerian species have narrow distributional range, e.g., G. villosa and G. tenax occur in the drier regions of Northern Guinea Savannah while G. hookerana and G. oligoneura are found only in the South-West and south-East high forests, respectively (Table.1). The 16 species studied are tentatively divided into three groups based on their ecological preferences viz: Group A: G. malacocarpa, G. barombiensis, G. brunnea, G. coriacea, G. hookerana and G. oligoneura all distributed in the rainforest ecological zone; Group B: G. barteri, G. bicolor, G. cissoides, G. flavescens, G. bicolor, G. lasiodiscus, G. tenax, G. venusta, G. villosa distributed in the savannah zone and Group C consisting of G. carpinifolia and G. mollis found in both ecological zones. Pollen analysis: Two pollen types and 5 subtypes are recognized for the Nigerian species of Grewia based on light microscopy (Fig. 2.). Grewia type This is typical of the savanna species and the two species occupying both ecological zones. The pollen grains are large to medium in size, isopolar and radially symmetrical; Polar view (30.0 - 67.5 µm); Equatorial diameter (18.8 – 82.5µm). Polar/Equatorial diameter ratio: 89 – 175% (Table 3); oblate spheroidal, subprolate and prolate; Ambient circular, trilobed or triangular; 3-colporate except G. venusta 3-4 colporate (Table 2); Colpi long (20.0 – 45.0) µm, colpi ends acute, open or acute/open; Ora(endoaperture) lalongate, 6.0 -15.0µm broad, rectangular; Sexine thicker than nexine; Nexine 0.50 – 2.5µm; Exine 1.06 – 2.75µm thick. The pollen grains are elliptic, circular, oblong and oval in equatorial view; striate, reticulate or coarse reticulate sculpture with bacules; lumina small or large (0.90-13.1)µm in diameter, distinct; lumina shape polygonal to irregular, muri ridge entire or wavy, with or without bacules (Table 2& 3).
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