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Azeez and Folorunso (2).Cdr Ife Journal of Science vol. 16, no. 2 (2014) 171 PHENOLOGY AND POLLEN STUDIES OF SOME SPECIES OF ANNONACEAE IN NIGERIA. Azeez, S.O.* and Folorunso, A.E. Obafemi Awolowo University, Department of Botany, Ile-Ife, Osun State, Nigeria. Corresponding Author; email: [email protected]. (Received: 17th April, 2014 ; Accepted: 27th May, 2014) ABSTRACT Phenology and pollen studies were investigated in some species of Annonaceae in Nigeria to help in providing pollen data and record of flowering patterns of the species studied. Phenological data for flowering and fruiting were tracked once a week for both rainy and dry seasons. Pollen grains from fresh anthers of the different species of Annonaceae were collected for pollen grain stainability and morphological studies. The peak flowering period in all the species studied was between March and April. Flower production took place before flushing of leaves in M. tenuifolia, however, in A. reticulata and A. squamosa, flowering and flushing of leaves occurred at the same time. All the species have predictable annual fruiting periods; the harvesting of their fruits and seeds, and sustainable management is bound to be relatively easy. The percentage pollen stainability was higher than percentage pollen germination in the species studied. Low fertility was observed in A. reticulata and C. patens. Tetrad pollen observed in Annona, Monodora and Xylopia is an advanced character in Annonaceae and could have an advantage over the monad. The genus Cleistopholis now appears to be one of those genera which have retained the largest number of the characteristics of a primitive member of Annonaceae. The evolutionary development is from Cleistopholis (monad) to Xylopia (tetrad) to Monodora (tetrad) to Annona (tetrad). Keywords: Phenology, Pollen, Annonaceae, Evolution, Monad, Tetrad. INTRODUCTION monosulcate pollen as in Anaxagoreoideae and The dearth of phenological studies in tropical Ambarioideae, was ancestral in the family. Because plants has been pointed out by various workers columellar taxa diverged earlier than previously (Morellato and Haddad, 2000; Keatley, 2000). thought, columellae may have re-originated in the Phenological studies of tropical plants aim for a common ancestor of Malmeiodeae and better understanding of the intricate relationship Annonoideae, which make up most of the family, of species maintenance through the processes of with several later reversals to granular structure pollination and seed dispersal. Phenological data (Chatrou et al., 2012). are of great importance for the correct use and protection of the ecosystems (Machado et al., This work therefore studies the phenological 1997). Phenological parameters could facilitate parameters of Annona senegalensis, Annona squamosa, data collection from these taxa in Annonaceae if Annona muricata, Annona reticulata, Cleistopholis well documented. Farmers and fruit growers can patens, Monodora tenuifolia and Xylopia aethiopica to use phenological data as a tool in knowing when to facilitate data collection in them. Pollen plant and harvest crops and in some cases when to parameters in these species were also studied to apply herbicides and insecticides. It could also aid resolve the problem of classification in the family. in resolving the confusion arising from the classification of this family. MATERIALS AND METHODS The classification of the Annonaceae has been a Phenological Studies herculean task to workers on this family, the largest The flowering and fruiting periods of some of the family of primitive Angiosperms (Chatrou, 2005). species of Annonaceae studied in Obafemi Palynology has played a major role in attempts to Awolowo University Ile-Ife, Nigeria Campus clarify higher-level relationships in the (longitude 7° ̎ 50.70'' N and latitude 4° 58.44'' E) Annonaceae. Four major clades including were observed and documented. These plants Annonoideae, Anaxagoreoideae, Ambarioideae were surveyed once a week for both rainy and dry and Malmeiodeae have been reported for seasons, the trend of the flowering, fruiting and Annonaceae (Chatrou et al., 2012). Granular shedding of leaves were then recorded. The 172 Azeez and Folorunso: Phenology and Pollen Studies of some Species of Annonaceae in Nigeria. phenology was first studied between year 2003 and rainy season. All the species studied are deciduous 2005; after three years, the phenology was studied because, at one time or the other, there was again which covered between 2008 and 2010. effoliation and leaf flushing. Shedding of leaves occurred mostly during the dry season around Pollen Studies November to January. In Monodora tenuifolia there Pollen grains from fresh anthers of the different was flowering after defoliation followed by leaf species of Annonaceae studied were collected for flush. In A. reticulata and A. squamosa, flowering pollen grains stainability and pollen and flushing of leaves occurred simultaneously. morphological studies. For each species, pollen Flowering and fruiting occurs throughout the year grains were shaken onto a clean slide and stained in A. muricata while in A. senegalensis and A. reticulata with Cotton Blue in Lactophenol. Four slides were only flowering occurs throughout the year (Figure prepared for each species. The pollen grains were 1). Cleistopholis patens flowers between January and examined and scored for stainability at 100 July. A. squamosa flowers between February and magnification and percentage stainability was June; M. tenuifolia flowers between February and calculated. Well formed, intact and uniformly- August and Xylopia aethiopica flowers between May stained pollen grains were considered viable while and December. During the latter part of the those that were only partially stained or not stained flowering period, only the younger branches at all and with collapsed outline were scored as towards the top produce flowers. The peak non-viable (Olorode and Baquar, 1976). Pollen flowering period for all the species studied except germination was carried out using the method of Xylopia aethiopica was between March and April Nurhan (2003). Fresh matured pollen grains were while the peak flowering period in Xylopia aethiopica dusted on the sterilized culture medium prepared was July. from 25ml distilled water, 0.5g agar and 6.25g saccharose. The culture medium was left for 12 The fruiting period in A. senegalensis falls between hours at 18OC for pollen tubes to germinate in vitro. December and September with peak period The pollen grains were then fixed in 1:3 acetic- between May and July (Figure 2). A. reticulata alcohol. 1 drop of this suspension was put on the produces fruit between June and December slide and then stained with Cotton Blue in having a peak period between November and Lactophenol. The slides were viewed under oil and December. Cleistopholis patens has its fruiting peak phase contrast illumination and then period between April and May; Xylopia aethiopica photographed under Lietz Dialux research produces fruits between June and January with the microscope. peak fruiting period between August and September while that of Monodora tenuifolia occurs The pollen grains were acetolysed following the between May and June (Figure 2). The peak procedure of Erdtman (1943). The pollen grains fruiting period in A. squamosa is in May. The peak were mounted in the dilute 25% glycerine and flowering and peak fruiting period of A. muricata photomicrographs of the acetolysed pollen grains occurred three to four times in a year. It was noted were taken with Leica camera, mounted on a Leitz that there was variation among years in time, Dialux research microscope. length and abundance of flowering and fruiting. RESULTS Compound pollen grains in form of tetrad It was observed that A. senegalensis and Cleistopholis occurred generally in all the species of patens flowers during the dry season to the Annonaceae studied except in Cleistopholis patens beginning of rainy season between January and (Figure 3). A. muricata has the largest pollen April. A. squamosa, A. reticulata, Monodora tenuifolia diameter and C. patens has the smallest diameter flower during the early rainy season between May (Table 1). It was observed that the tetrad structure and August. The flowering period of A. muricata of Xylopia aethiopica could be easily broken up and Xylopia aethiopica covers both the early and the when it was acetolysed. Clestopholis patens has oblate late rainy season, March to October (Figure 1). pollen structure and it is surrounded by a thin wall. The fruiting periods of the species is generally This is a pollen grain in which the polar axis is from the beginning of the rainy season to the late shorter than the equatorial axis. The pollen Azeez and Folorunso: Phenology and Pollen Studies of some Species of Annonaceae in Nigeria. 173 structure of A. squamosa has a tetragonal tetrad in also perforated. The pollen forms of Monodora which the centres of the grains define a tenuifolia (Figure 3) and A. reticulata are similar to tetrahedron, it is transparent and surrounded by a that of A. senegalensis, though malformation was thin wall. The pollens of A. muricata are also observed in some pollens of A. reticulata. A tetragonal tetrads like A. squamosa, however it is spheroidal pollen grain structure was observed in perforated, not transparent and surrounded by a Xylopia aethiopica and it is perforated. In this pollen thick wall (Figure 3). type, the polar axis and the equatorial diameter are approximately equal (Figure 3). The pollen is Annona senegalensis has tetragonal tetrad, a unipolar tetraporate. However, when the pollen had not tetrad in which all four members are in contact at broken up, it had oblate pollen grain structure. the centre of the tetrad, so that in the correct orientation, the adjacent walls form a cross. It is Table 1: Pollen Stainability and Germination in Relation to Pollen Diameter. Species Pollen stainability Pollen germination Mean (µm) diameter % % A. muricata 81.56 37.21 23.43 A. reticulata 43.64 12.45 6.30 A.senegalensis 66.26 21.32 10.50 A. squamosa 74.38 12.88 7.00 C. patens 38.53 0.94 1.75 M. tenuifolia 92.24 65.83 10.15 X.
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