American Journal of Plant Sciences, 2011, 2, 733-743 doi:10.4236/ajps.2011.26088 Published Online December 2011 (http://www.SciRP.org/journal/ajps) Copyright © 2011 SciRes. AJPS 733 Ecological Structure and Fruit Production of African Fan Palm (Borassus aethiopum) Populations Christine Ouinsavi, Charlemagne Gbémavo, Nestor Sokpon Laboratoire d’Etudes et de Recherches Forestières (LERF), Faculté d’Agronomie, Université de Parakou, Parakou, Benin. Email: {ouinsch, cgbemavo, nsokpon}@yahoo.fr Received July 6th, 2011; revised August 22nd, 2011; accepted September 9th, 2011. ABSTRACT Ecological structure and fruit production of African fan palm stands in Benin were studied on the basis of surveys laid out in ten (10) populations located in three ecological zones. Dendrometric and fruit production parameters such as: diameter at breast height (dbh), total height (TH), number of fruit bunch per tree, number of fruits per tree, number of seeds per fruit, fruit length and width were measured on all African fan palm trees within thirteen (13) plots of 1,000 m2. Diameter and height classes distribution of the palms in each ecological zone adjusted to Weibull distribution showed a bell shaped curve with left dissymmetry, characteristic of young stands (form coefficient between 1 and 3.6). African fan palm stem number was 156 trees/ha. Variation of dbh, TH, number of fruit bunch per tree were highly significant between populations and ecological zones (P < 0.05%). Trees from Sudanian zone were highly grown (mean dbh = 52.2 cm) while those of Sudano - Guinean transition zone were less (mean dbh = 32.73 cm) and dominated by female trees. African fan palm stands of Pahou is suitable for seed collection for in situ conservation of the species while those of Ouorghi and Akomiah could provide fruits for edible hypocotyls production. Keywords: Ecological Structure, African Fan Palm, Fruit Producti on, Population 1. Introduction Tropical forests are an important reservoir of biodiversity and they play a fundamental role in giving satisfaction to many needs of the people. Nowadays, the non timber forest products, exploited and consumed by local people are becoming very scarce due to high human pressure, over exploitation, non sufficient silvicultural data, cli- mate changes highlighted by the recurrent dry seasons, all situation that compromises local people food security and income. It is the case of African fan palm (Borassus aethiopum) which provide people with a wide usefulness on ecological, nutritional, therapeutically, small scale producing and culturally plans [1-5]. African fan palm roots are used as anti asthmatic. Its leaves are used in many craftsmen products. The petioles are used as fenc- ing, firewood and the hypocotyls are edible [4]. The male flowers are used as soils fertilizers and as fodder [6]. They are considered as an excellent fodder and their nu- trients content are similar to those of ground nut and cow- pea haulm [7]. The bole of the palm is used in carpentry for house and bridge building. Its wood is very resis- tant to termites and fungi attacks and to climate varia- tions. In Benin, a research work carried out by [3] and [4] revealed that from the sampled population, 90% of the households eats African fan palm hypocotyls, 16% eats the fruits and 13% consume the almonds. The same au- thors showed that African fan palm contributes to cure 33 diseases and is of great importance to Lokpa and Yom ethnic groups in the Northern Benin. African fan palm products are sold and can allow the traders to draw month- ly gross margin ranging from 20,920 to 112,975 FCFA for hypocotyls, 332 to 1925 FCFA for fans selling, average of 6500 FCFA for hats and 1616 to 3264 FCFA for sieves selling. According to [3], more than 88% of the organs of African fan palm are useful (hypocotyls, fruits, almonds, leaves, petioles, blades, boles) and harvested from natural forests. Meanwhile, African fan palm can ge- nerate a lot income to the rural population and contribute to poverty alleviation only few works were carried out on the structure and dynamics of its populations. This paper aims to assess ecological structure and pa-
Ecological Structure and Fruit Production of African Fan Palm (Borassus a ethiop um) Populations 734 ttern of fruit production of Borassus aethiopum for a better management of the tree populations in Benin. 2. Material and methods 2.1. The Study Area This study was carried out in the Republic of Benin located in West Africa between 1˚ and 3˚40'E and 06˚30' and 12˚30'N. It is bordered by Niger in the north, Bur- kina Faso in the northwest, Togo in the west, Nigeria in the east and the Atlantic Ocean in the south and covers an area of 112,622 km2 (Figure 1). Benin is located in the Dahomey gap which is charac- terized by numerous physical constraints. The mean an- nual rainfall is low (900 - 1300 mm) compared with neigh- bouring countries in the Guineo-Congolean zone such as Ghana and Côte d’Ivoire (average of 1500 mm) and Ca- meroon (1800 - 3000 mm). Reference [8] distinguished four major types of vege- tation in Benin which are described as follows: The coastal and sub-coastal zone, which extends from the coast (6˚20'N up to 7˚N), is composed of beach ve- getation with an abundance of Remirea maritima and Ipomea pescaprae, anthropic vegetation of mangrove swamp forests characterized by Rhizophora racemosa, Avicennia germinans and Dalbergia ecastaphylum, ri- parian forests, oil palm and coconut plantations. This zone has a subequatorial climate type with two rainy seasons (from mid-March to mid-July and from mid-September to mid-November) which alternate with two dry seasons (from mid-July to mid-September and from mid-Nove- mber to mid-March). The Guineo-Congolean zone composed of relict semi- deciduous forests characterized by species such as: Dia- lium guineense, Triplochiton scleroxylon, Strombosia glau- cescens, Cleistopholis patens, Ficus mucuso, Cola cordi- folia, Ceiba pentandra, Trilepisium madagascariense, Celtis spp, Albizia spp, Antiaris toxicaria, Diopyros mespiliformi s, Drypetes floribunda, Memecylon afzelii, Celtis brownii, Mimusops andogensis, Daniellia oliveri, Parkia spp and Vitellaria paradoxa [9-11], some savan- na and plantation areas. The Guineo-Sudanian Transition zone with a mosaic of forest woodlands, sometimes dry forests and wooded savanna crossed by gallery forests. The most abundant species found in this zone are: Isoberlinia doka, I. tomen- tosa, Monotes kerstingii, Uapaca togoensis, Anogeissus leiocarpa , Antiaris toxicaria, Ceiba pentandra, Blighia sapida, Dialium guineense, Combretum fragrans, Entada Africana, Maranthes polyandra, Pterocarpus erinaceus, Terminalia la xiflora and Detarium microcarpum. These two zones are characterized by a Guinean cli- mate type with two rainy seasons and two dry seasons distributed as above but the short rainy season of this region is sometimes absent. The total annual rainfall rea- ches 1100 mm. The Sudanian zone composed of dry forests, savanna woodlands, shrub savanna and savanna grassland. The gallery forests are characterized by species such as: Hae- matostaphis barteri, Lannea spp, Khaya senegalensis, Anogeissus leiocarpa, Tamarindus indica, Capparis spi- nosa, Ziziphus mucronata, Combretum spp and Cissus quadrangularis. This zone has a Sudanian climate type with one rainy season (from April to October) followed by one dry season (from November to March). The aver- age rainfall varies from 1100 mm in the southern part of this zone to 900 mm in the northern part. According to the FAO world classification of soils, the major types of soil distinguished in Benin are: Leptosols in the coastal zone, Acrisols, Ferralsols or Lixisols and Vertisols in the Guineo-Congolean zone, Luvisols, Lixisols, Alisols, Plinthosols and Arenosols in the Sudano-Guinean transition and the Sudanian zones. Human pressure on land for agricultural purposes and forest exploitation is consi- derable. In southern Benin where population density varies from 200 to 450 inhabitants/km2, average farm- land area is 0.5 ha and the fallow system is more or less absent. In the north, where mean population density is 14 inhabitants per km2, average farmland area is 5 ha and fallow length is about 5 years. Forest over-exploitation has led to declines in the num- ber of some trees species per hectare. Species such as Mi- licia excelsa, Khaya senegalensis, Afzelia Africana and Pterocarpus erinaceus have become scarce in their natural habitat [12]. Human pressure on forest ecosystems has also reduced the occurrence of relict forests, mainly sacred groves [13] and individual trees of some species, such as Milicia excelsa, Triplochiton scleroxylon, Borassus aeth io- pum, protected by traditional ethno botanic conservation practices [14]. 2.2. B. aethiopum Description and Populations Sampling Borassus aethiopum is a common palm in West African humid savannas. It is found in drier areas but is then re- stricted to riversides [15]. It is a savannah tree of 20 to 30 m high and 0.30 to 0.-0 m of diameter. It is a dioe- cious, tall, solitary and pleonanthic palm of the Bora- sseae tribe [16]. Male and female flowers appear on se- parate trees. Leaves are fan-shaped, induplicate and cos- tapalmate with petioles of 1.5 to 3 m long. The stem is massive and covered with leaf bases abscising clearly in older individuals. The male flowers are abundant and small, inserted at the base of a bract. The female flowers are larger than the males. Female bear 50 - 100, 1.0 - 1.5 Copyright © 2011 SciRes. AJPS
Ecological Structure and Fruit Production of African Fan Palm (Borassus a ethiop um) Populations Copyright © 2011 SciRes. AJPS 735 Figure 1 . Distributio n of African fan palm populations ac c ording to the ecological zones of [17]. kg fresh weight fleshy fruits [15]. Fruits are globulous or ovoid, yellowish when mature with one to four seeds. The sap is used to make palm wine [18]. Stems and leaves are used to construct houses. The tree’s roots are used in medicine and the bowl is used in bridge construction and carpentry. Fruits and young seedlings are edible [19]. Seed germination is remote-tubular [16]; [20]: the cotyle- donary axis extends downward into the soil and carries the seedling to a depth of 40 cm, where the first roots and leaves develop. African fan palm is originated from Sahelian Africa and its distribution area ranges from Senegal to Tchad. The species is found in Benin, Niger, Togo, Ghana, Côte d’Ivoire, Gambia, Senegal, Burkina-Faso, Mali, Mauri- tania, Nigeria and Tchad. In Benin, African fan palm populations were sampled according to three ecological zones as follows: Goroubi, Loumbou-loumbou, Niaro and Kongourou in the Suda- nian zone; Akomiah and Ouorghi in the Sudano-Guinean Transition zone; Se, Grand-Popo, Pahou and Ouidah in the Guinean zone (Table 1). 2.3. Data Collection Rectangular plots of 1000 m2 were laid in each sampled population of African fan palm. The plot number varies according to the area covered by African fan palm popu- lation. Diameter at breast high and the total height of all the fan palm trees contained in each plot were measured. In total, 203 palm trees covering 13,000 m2 were mea- sured (Table 2). The sex of each tree, the number of fruits bunch and the number of fruits per female tree were recorded. The size (length and width) of each fruit and the number of seeds per fruit were also recorded. In total 82 males and 121 female trees were sampled to estimate tree fruit production. 2.4. Data Analysis African fan palm trees distribution within population and according to the ecological zones was adjusted to the Weibull function, commonly nowadays used in Forestry surveys [21-23]. The Weibull distribution is as follows: 1 () exp cc cxa xa fx bb b a = position parameter; a = 20 cm (diameter); a = 5 or 7 m (Height according to the sites); b = scale parameter or size parameter; c = form parameter linked with the observed structure. Trees diameter or trees height distribution were ad- justed to Weibull function with Minitab 14 Software. A one way Analysis of Variance was performed to test for statistical difference between different ecological zones
Ecological Structure and Fruit Production of African Fan Palm (Borassus a ethiop um) Populations 736 Table 1. Distrib ution of B. aethiopum populations acco rding to the three e cological zones. Ecological zones B. aethiopum populations Latitude Longitude Rainfall gradient Goroubi 11˚55.055 003˚15.484 Lombou-loumbou 12˚14.217 002˚54.218 Niaro 10˚23.433 002˚25.039 700 to 900 mm Sudanian Kongourou 10˚22.571 002˚50.308 Akomiah 08˚06.241 002˚32.432 Sudano-Guinean Transition Ouorghi 08˚10.201 002˚37.221 1000 to 1200 mm Se 06˚29.037 001˚50.152 1100 mm Pahou 06˚23.177 002˚14.069 1200 mm Grand Popo 06˚17.484 001˚49.278 900 mm Guinean Ouidah 06˚23.429 001˚58.538 1200 mm Table 2. Descript ion of B. aethiopum studied pop ulations . Ecological zones B. aethiopum populations Plot number Trees number Total plots number per ecological zones Total trees number per ecological zones Goroubi 1 3 Lombou -loumbou 1 21 Niaro 1 21 Sudanian Kongourou 1 17 4 62 Akomiah 2 30 Sudano-Guinean Transition Ouorghi 2 30 4 60 Se 1 20 Pahou 1 18 Grand Popo 2 22 Guinean Ouidah 1 21 5 81 Total 13 203 13 203 for variable such as: dbh, total height, fruits number per tree, fruits cluster number per tree, seeds number per fruit, fruit size (length and width) using SPSS for Win- dows V. 16 Software. Turkey Homogeneity test allows us to separate the site and ecological homogeneous groups. The correlations between growth parameters and fruits production were performed with Minitab 14 Software. 3. Results 3.1. Diameter Classes’ Distribution of B. aethiopum Trees Figure 2 showed the class distribution of the fan palm trees diameter according to the different ecological zones. The Weibull distribution of fan palm trees per diameter classes shows a left dissymmetric bell shaped curve with a c form coefficient comprised between 1 and 3.6 for the three ecological zones. This value of c means that the stands were dominated by young trees. In the Sudanian zone, trees of diameter class centre 35 and 65 were do- minant in the stand while in the Sudano-Guinean Tran- sition zone and Guinean zone, trees of diameter class centre 25 were dominant. 3.2. Total Height Classes Distribution of B. aethiopum Trees Figure 3 showed the total height class distribution of trees of African fan palm according to the three ecologi- cal zones. This distribution well adjusted to the Weibull function showed a bell shaped curve with a left dissym- metry (1 ≤ c ≤ 3.6). Like the diameter class distribution, the stands were dominated by young trees. In the Su- danian zone, trees of class (8 - 12 m) were dominant in the population while in the Sudano-Guinean Transition zone and the Guinean zone, trees of height class (12 - 14 m) were dominant in the stands. 3.3. Structural Characteristics of B. aethiopum Populations Table 3 showed the number of tree per stand and per ecological zones of the studied African Fan Palm popu- Copyright © 2011 SciRes. AJPS
Ecological Structure and Fruit Production of African Fan Palm (Borassus a ethiop um) Populations737 Center of diameter classes Frequency ( trees/Class) 85756555453525 20 15 10 5 0 Shape 2,235 Scale 37,08 Thresh 20 N6 3-Parameter Weibull Sudanean zone populations 2 2.235 37.08 Center of diameter classes F re quenc y (trees/class) 55453525 40 30 20 10 0 Shape 2,031 Scale 14,71 Thresh 20 N60 3-Parameter Weibull Sudano-Guinean transition zone populations 2.031 14.71 C enter of diam eter classes F re q uency ( trees/class) 756555453525 40 30 20 10 0 Shape 1,498 Scale 17,26 Thresh 20 N81 3-Parameter Weibull Guinean zone populations 1.498 17.26 Figure 2. Diameter classes’ distributio n of trees of African fan palm population s. Center of height c lasses Frequency ( trees/Class) 242016128 20 15 10 5 0 Shape 1,659 Scale 7,087 Thresh 7 N6 3-Parameter Weibull Sudanean zone p o pulat ions 2 1.659 7.087 Center of height classes Frequenc y (tree s/Classe) 18161412 40 30 20 10 0 Shape 2,347 Scale 3,307 Thresh 11 N60 3-Parameter Weibull Sudano -Guinea n transitio n zone populations 2.347 3.307 Ce nter of he ight classes F re que ncy (tree s/Class) 181614121086 40 30 20 10 0 Shape 3,530 Scale 8,290 Thresh 5 N81 3-Parameter Weibull Guinean zone populations 3.530 8.290 Figure 3. Total height classes distribution of trees of African fan palm populations . Copyright © 2011 SciRes. AJPS
Ecological Structure and Fruit Production of African Fan Palm (Borassus a ethiop um) Populations 738 Table 3. Trees number per st and and per ecological zones of the stu died B. aethiopum populations. Ecological zones B. aethiopum populations Sampled area (m2)Number of trees Number of trees per ha Number of trees per ecological zones (trees/ha) Goroubi 1000 3 30 Lombou-loumbou 1000 21 210 Niaro 1000 21 210 Sudanian zone Kongourou 1000 17 170 155 Akomiah 2000 30 150 Sudano-Guinean Transition zone Ouorghi 2000 30 150 150 Se 1000 20 200 Pahou 1000 18 180 Grand Popo 2000 22 110 Guinean zone Ouidah 1000 21 210 162 Total 13,000 203 - - lations. The average number of trees of African fan palm was 156 trees per hectare across the study area. The den- sity varies from 150 trees per hectare with stands of Su- dano-Guinean Transition zone to 162 trees/ha when con- sidering populations of Guinean zone. The lower number of trees /ha were recorded in populations of Goroubi (30 trees/ha); Grand-Popo (110 trees/ha), Ouorghi and Ako- miah (150 trees/ha). Table 4 showed the structural charac- teristics and fruit production of African Fan Palm trees. The mean diameter at breast high of the palm trees ranged from 25.56 cm (Population of Se) to 62.03 cm (Population of Niaro) and Kongourou (62.05 cm). For the populations of Ouorghi, Akomiah, Goroubi, Grand Popo and Loumbou-loumbou, the mean diameter was respectively 31.55 cm; 33.92 cm; 34.43 cm; 33.88 cm and 36.32 cm. When considering the populations of Ouidah and Pahou, the mean diameter of the fan palm trees was as follows: 42.99 cm and 39.1 cm. There is a highly significant difference within and between the trees populations as far as the trees diameter is concerned (P < 0.001). When considering the ecological zones, the mean tree diameter was 52.20 cm with the Sudanian popula- tions; 32.73 cm in the Sudano-Guinean Transition zone and 35.34 cm in the Guinean zone. The mean diameter of the African fan palm trees highly varied according to the ecological zones. The mean total height of the trees varied from 9.77 m (Population of Kongourou) to 17.52 m (Population of Loumbou-loumbou). When considering the population of Akomiah, Goroubi, Ouidah and Ouorghi, the mean total height of the trees was respectively 13.98 m; 13.50m; 12.79 m and 13.96 m. There is a highly significant dif- ference between the African fan palm trees populations as far as the trees total height is concerned (P < 0.001). When considering the ecological zones, the mean total height of trees varies from 12.38 m (populations of Gui- nean zone to 13.16 m (Populations of Sudanian zone) and 13.97 m (Population of Sudano-Guinean Transition zone). 3.4. Fru i t s P r od u ction of B. aethiopum Populations The fruit number increased with size as shown on Figure 4. The minimum fruiting diameter was 25 cm with a maximum of 85 cm. The maximum average number of fruit per tree (120 - 140 cm) was obtained for the dia- meter class centre 55 cm while in the younger diameter class (25 - 45 cm) this average number ranged from 60 to 80. Table 3 also showed fruit production feature of African fan palm populations. The average number of fruit per tree started to decrease from diameter class cen- tre 85 cm. The average number of fruits bunch varied from 3 (Population of Grand-Popo) to 10 (Population of Loumbou-loumbou). For the populations of Niaro, Ouidah, Ouorghi, Kongourou, the average number of fruit bunch was respectively: 7.4; 7.83; 7.47 and 8.2. There is a significant difference between the African fan palm po- pulations as far as the number of fruit bunch per tree is concerned. When considering the ecological zones, the average number of fruit bunch was 8.65 for the po- pulations of the Sudanian zone; 6.12 in the Sudano- Guinean Transition zone and 7.21 for the populations of Guinean zone. The average number of fruit per tree varied from 14.5 (Population of Grand Popo) to 103.4 (Population of Pahou). When considering the populations of Loumbou- loumbou, Niaro, Ouidah, Ouorghi and Kongourou, the average number of fruit per tree was respectively: 50.77; 86; 91.17; 89.74: 89.20. When considering the different ecological zones, the average number of fruit per tree was 62.58 for Sudanian zone populations; 77.48 for po- pulations of Sudano-Guinean Transition zone and 85.41 Copyright © 2011 SciRes. AJPS
Ecological Structure and Fruit Production of African Fan Palm (Borassus a ethiop um) Populations739 Table 4. Structural characterist ics and fruit production of African fan pal m trees. Means with the same letter are not statis- tically different according to Turkey homogeneity test. B. aeth iopum populations Mean DBH (cm) Mean total height (m) Average number of bunch of fruit per treeAverage number of fruit per tree Average number of seeds per fruit Fruit length Fruit width Akomiah 33.92ab 13.98c 6.12ab 77.48ab 2.50a 12.95a 14.98a Goroubi 34.43ab 13.50bc 7ab 35.67ab 2.75a 13.23a 12.87a Granp Popo 33.88ab 9.92a 3a 14.50a - - - Loumbou-lombou 36.32bc 17.52d 10b 50.77ab 1.69a 13.29a 14.26a Niaro 62.63d 11.50ab 7.40ab 86.00ab 2.11a 12.42a 13.56a Ouidah 42.99c 12.79bc 7.83ab 91.17ab - - - Ouorghi 31.55ab 13.96c 7.47ab 89.74ab 2.54a 13.39a 14.16a Pahou 39.1bc 13.38bc 7.7ab 103.40b - - - Sè 25.56a 13.77c 5.8ab 64.6ab 2.67a 12.47a 14.53a Kongourou 62.05d 9.77a 8.20ab 89.20ab 2.00a 14.30a 14.70a Probability (%) 0.000 0.000 0.030 0.020 0.155 0.939 0.557 Ecological zones Sudanian 52.20b 13.16ab 8.65b 62.58a 2a 13.03a 13.84a Sudano-Guinean Transition 32.73a 13.97b 6.12a 77.48a 2.67a 12.47a 14.53a Guinean 35.34a 12.38a 7.21ab 85.41a 2.52a 13.20a 14.52a Probability (%) 0.000 0.003 0.023 0.092 0.089 0.851 0.415 Figure 4. Diameter class distribution of number of fruits per African fan palm tree. for populations of Guinean zone. The mean length of the fruits varied from 12 cm (Populations of Niaro, Se) to 14.3 (Population of Kongourou) while the mean width of the fruit varied from 12 cm (Population of Goroubi) to 14.98 cm (Population of Akomiah). 3.5. Correlations between Fruit Production Parameters Table 5 showed the correlation patterns between trees dbh, the fruit number and the number of fruit bunch per tree. The number of fruit bunch and the number of fruits per tree were not correlated with the diameter at breast height. However, the number of fruit per tree was strongly correlated with the number of fruit bunch it bears. There was strong correlation between fruit size (length and width) and the number of seeds per fruit (Table 6). Table 7 showed the number of seed per fruit according to popu- lations and ecological zones. When considering the eco- logical zones, fruits in populations of the Sudanian zone (Goroubi, Loumbou-loumbou, Niaro) and in Sudano- Guinean transition zone (Akomiah, Ouorghi) have three seeds while those of Guinean zone have one or two seeds. The average fruit number per female tree varies between the populations within a given ecological zone but not from one ecological zone to the other. African fan palm trees in the Sudanian populations were larger (high mean dbh, high total height and high number of fruit cluster) but did not have the higher number of fruits per tree. Trees in the Ouidah population produced the highest number of fruit (91.17 fruit per tree in average). Tees in the Guinean zone produced the highest number of fruits with an average of 85.4 fruits per tree. Most of the palm trees in the Sudano-Guinean transition zone were female while those in the Sudanian zone were almost equal in sex (Table 8). 4. Discussion 4.1. Structural Characteristics of B. aethiopum Populations The mean number of palm trees per hectare recorded in Copyright © 2011 SciRes. AJPS
Ecological Structure and Fruit Production of African Fan Palm (Borassus a ethiop um) Populations 740 Table 5. Correlation between trees dbh, fruit number and number of fruit bunch per trees. Number of fruit bunch Fruit numberDbh Number of fruit bunch 1 Fruit number 0.707*** 1 Dbh 0.157 0.159 1 Table 6. Correlation between fr uit l ength and width and the number of seeds per fruit. Fruit length Fruit width Seeds number Fruit length 1 Fruit width 0.825*** 1 Seed number 0.677*** 0.568*** 1 *** Significant Correlation at the level of 1%. Table 7. Number of seed per fruit according to B. aethio- pum populati ons and ecological zones. Fruits number with: B. aethiopum populations 1 kernel 2 seeds 3 seeds4 seeds Goroubi 1 0 2 1 Loumbou-loumbou 5 4 3 0 Niaro 3 2 4 0 Sudanian Kongourou 0 1 0 0 Akomiah 2 1 7 0 Sudano -Guinean Transition Ouorghi 1 4 8 0 Ouidah - - - - Se 0 1 2 0 Pahou - - - - Guinean Grand popo - - - - Total 12 13 27 1 Table 8. Sex ratio of trees of B. aethiopum populations. Means with the same letter are not statistically significant according to Student t test at the level of 5%. Ecological zones B. aethiopum populations Male trees number Female trees number Ratio male/female Goroubi 0 3 - Loumbou- loumbou 8 13 0.61a Niaro 11 10 1.1a Sudanian Kongourou 12 5 2.4a Akomiah 5 25 0.2a Sudano-Guinean Transition Ouorghi 8 22 0.36a Ouidah 9 12 0.75a Se 9 11 0.81a Pahou 8 10 0.8a Guinean Grand popo 12 10 1.2a Total 82 121 - the different ecological zones is similar to those obtained by [24] in Wolokonto village in Burkina-Faso. The dia- meter at breast high for the palm trees in the Sudanian zone was higher than those of trees in Guinean and Sudano-Guinean transition zone. According to [25], Suda- nian climate was suitable for the optimal development of the African fan palm trees. However, there is generally a relationship between species temperament and their stem diameter class distribution. The shade tolerant species have an all-age population structure which gives the cha- racteristic inverse J-shaped curve or negative exponential stems distribution. These species are stable species as far as the equilibrium stand distribution is concerned [26-31]. The bell-shaped distribution species are generally light demanding species. They are generally gap depending for their regeneration. Mortality is higher in earlier stage under closed forest canopy. The number of large trees is higher. More caution is required as far as species tem- perament and stem class distribution are concerned. The stem diameter distribution of species greatly varies with observed plot size. Some species having a bell-shaped distribution within small plot could show inverse J- sha- ped when plots are larger (regular distribution to aggre- gative, [32]. In Ghana, for instance, [33] showed that on a national level, the majority of the forest species have sufficient natural regeneration and most species show inverse J-shaped curves. Furthermore, finding patterns among life histories is a challenge for modern ecology [34]. It requires linking evolution and demography and has led to many theore- tical models and field studies [35]. According to [23], the use of Weibull distribution probability density function is becoming increasingly popular for modeling the dia- meter distributions of both even and uneven-aged forest stands. The popularity of Weibull is derived from its fle- xibility to take on a number of different shapes corres- ponding to many different observed unimodal tree- diameter distributions. In addition, the cumulative distri- bution function of Weibull exists in closed form and thus allows for quick and easy estimation of the number of trees by diameter class without integration of the proba- bility density function once the parameters have been fitted. The bell shaped function obtained with the dia- meter or height classes distribution of the palm trees with a left dissymmetry corroborate the results of [36] with the African fan palm population of Wolokonto in Burkina- Faso. Similar results were also obtained by [37] with the Shea butter trees of Donfelgou in Togo. Also [23] ob- tained the same distribution as far as Afzelia africana trees populations are concerned in Benin. However this structure might not be derived only from the species temperament but also from human pressure. Copyright © 2011 SciRes. AJPS
Ecological Structure and Fruit Production of African Fan Palm (Borassus a ethiop um) Populations741 4.2. Fru i t and H y pocotyls Pro duct io n an d Sustainable Management of the B. aethiopum Populations Even though, the average number of fruit per tree did not vary significantly from one ecological zone to the other, trees in the Guinean zone and mainly the population of Pahou were more productive in term of number of fruit (103.4 fruits per tree). The palm tree with the higher number of fruits was from Pahou population (270 fruits). According to [16], [38] in [39], this number is higher than those obtained with the populations of African fan palm in Niger (250 fruits per tree) and in Côte d’Ivoire (50 - 100 fruits per tree). When we considered the number of seeds per fruit, trees of populations of the Sudano-Guinean transition zone have in majority (15 out of 23) three seeds. The number of fruit with three seeds is higher in the popula- tions of Sudano-Guinean Transition zone than other po- pulations indicating that seeds from Sudano-Guinean Transition zone are suitable for hypocotyls production. B. aethiopum tree is nowadays subjected to high pres- sure and this situation was emphasized by [13] who clas- sified the species as an endangered species on which many conservation actions should be focused. This pres- sure on the species is related to severe fruits and hypo- cotyls harvesting that hypothesised natural regeneration of the species. As each organ of the tree is of great importance, long term management of B. aethiopum populations is needed to improve local population livelihood. Therefore, sus- tainable management of African fan palm populations for providing woods for housing, fencing and fuel woods should mainly rely on the male trees with a sex ratio of three females for one male and an optimal number of 100 trees per hectare. On this basis, we suggested that the number of trees to be harvested should be: - 3 males from the population of Loumbou-loumbou, 7 from Niaro and 9 from Kongourou population as far as the Sudanian zone is concerned; - 10 and 2 female trees from Akomiah and Ouorghi po- pulations respectively as far as the Sudano-Guinean transition zone is concerned; - 5 male trees from each Ouidah, Sè and Pahou popu- lations in the Guinean zone. Natural regeneration of the palm trees should be pro- moted as well as hypocotyls production from nursery seed germination. 5. Conclusions Ecological structure of the African fan palm populations of Benin, adjusted to Weibull distribution showed a bell shaped curve with a left dissymmetry proving the pre- dominance of young trees within these populations. Stru- ctural characteristics of the stands greatly varied from one population to the other and from one ecological zone to the other. The trees mean dbh varied from 25.56 cm to 62.63 cm while the total height of the trees from 9.77 m to 17.52 m. The average number of fruit clusters varied from 3 (Population of Grand Popo) to 10 (Population of Loumbou-loumbou). The average number of fruits per tree varied from 14.5 (Population of Grand Popo) to 103.4 (Population of Pahou). The average number of seeds per fruit varied from 1 to 3. Most of fruits from populations of Sudanean and Sudano-Guinean Transi- tion zones had 3 seeds per fruit. The mean length of the fruits varied from 12.42 cm (Population of Niaro) to 14.3 cm (Population of Kongourou). The fruit mean width varied from 12.87 cm (Population of Goroubi) to 14.98 cm (Population of Akomiah). Environmental factors (Rainfall and temperature) seemed to discriminate the African fan palm populations of Benin and were key factors for the fruit production. When considering the ecological zones, trees of Guinean zone were higher fruit producers with an average fruits per tree of 85.4. Most of the fan palm trees of the Sudano-Guinean transition zone were female while those of the Sudanean zone were al- most equal in sex. Therefore sustainable management of African fan palm populations for providing woods for housing, fencing, and fuel woods should mainly rely on the male trees with a sex ratio of 3 females for 1 male and an optimal number per hectare of 100. However, as the species has particularly important socio-economic value for local people, there is a priority need to address the impact of its organs’ exploitation and eventually human-caused habitat fragmentation on its ge- netic structure for conservation purposes.” REFERENCES [1] B. C. Pamard, “Contraintes Ecologiques et Réalités Socio- Economiques. Exemple de Tafissou, Communauté Rurale du Contact Forêt—Savane sur la Branche Ouest du ‘V Baoulé’ (Côte D’Ivoire),” Cahiers ORSTOM, Série Scien- ces Humaines, Vol. 15, No. 2, 1978, pp. 51-79. [2] O. Bassir, “Some Nigerian Wines,” West Africa Journal of Biological and Applied Chemestry, Vol. 10, 1968, pp. 42-45. [3] N. Sokpon, E. Houankoun, C. Ouinsavi and J. Yabi, “Im- portance Socio-Economique du Rônier (Borassus ae- thiopum Mart): Différent Usages et Commercialisation de Quelques Sous Produit au Bénin,” Communication Présen- Tée à L’atelier Scientifique Sud-Centre, Abomey-Calavi, 2004. [4] D. S. E. Houankoun, “Importance Socio-Economique du Rônier (Borassus aethiopum): Différents Usages et Com- mercialisation de Quelques Sous-Produits au Bénin,” Mé- moire de DEA.FLASH/UAC, 2004, p. 90. Copyright © 2011 SciRes. AJPS
Ecological Structure and Fruit Production of African Fan Palm (Borassus a ethiop um) Populations 742 [5] J. Sakandé, O. G. Nacoulma, J. B. Nikiema, M. Compo, E. Bassene and I. P. Guissou, “Effect of Male Inflorescence of Borassusaethiopum,” Médécine d’Afrique Noire, Vol. 51, No. 5, 2004, pp. 280-282. [6] L. Price and B. O. Gaoh, “Local Communities and Mana- gement of the Palmyra Plantations of Dallol Naonis and the Niger River: The Example of a Dynamic of Sustainable Development in Niger,” Rapport, 1998. [7] S. Atta, “The Palmyra (Borassusa ethiopu m). Flowering, Biomass and Contribution of Nutrients,” Rapport, UICN/ PAIGLR/MH/E, Niger, 1997, p. 24. [8] E. Adjanohoun, V. Adjakidjè, M. A. Ahyi, et al., “Con- tribution Aux Etudes Ethnobotaniques et Floristiques en République Du Bénin,” Agence de Coopâeration Cultur- elle et Technique, Niamey, 1989, p. 895. [9] N. Sokpon, “Tenure Foncière et Propriété des Ligneux Dans Les Systèmes Agroforestiers Traditionnels au Bénin,” Annales de la Faculté des Sciences de Kisangani, Numéro Spécial, 1994, pp. 115-122. [10] N. Sokpon and J. Lejoly, “Les Plantes à Fruits Come- stibles D’une Forêt Semi—Caducifoliée de Pobè au Sud- est du Bénin,” In: C. M. Hladik, A. Hladik, H. Pagezy, O. F. Linares and A. Froment, Eds., L’alimentation en Forêt Tropicale: Interactions Bioculturelles et Perspectives de Développement, Vol. 1, UNESCO, Paris, 1996, pp. 115- 124. [11] C. Ouinsavi and N. Sokpon, “Traditional Agroforestry Systems as Tools for Conservation of Genetic Ressources of Milicia excelsa Welw. C.C. Berg. in Benin,” Agro- forestry Systems, Vol. 74, 2008, pp. 17-26. doi:10.1007/s10457-008-9109-6 [12] G. Agbahungba, N. Sokpon and O. et Gaoué, “State of Forest Genetic Resources of Benin,” Sub-Regional Work- shop FAO/IPGRI/ICRAF on the Conservation, Manage- ment, Use and Enhancement of Forest Genetic Resources in the Sahelian Zone, Ouagadougou, 22-24 September 1998, p. 30. [13] N. Sokpon and V. Agbo, “Sacred Groves as Tools for Indigenous Forests Management in Benin,” Annales des Sciences Agronomiques du Bénin, Vol. 2, 1999, pp. 161- 175. [14] C. Ouinsavi, N. Sokpon and S. O. Bada, “Utilization and Traditional Strategies of in Situ Conservation of Iroko (Milicia excelsa Welw. C.C. Berg.) in Benin,” Forest Ecology and Management, Vol. 207, No. 3, 2005, pp. 341-350. doi:10.1016/j.foreco.2004.10.069 [15] R. Vuattoux, “Le Peuplement de Palmier Rônier (Borassus aethiopum) D’une Savane de Côte D’Ivoire. Annales de l’Université D’Abidjan,” Série E. Université d’Abidjan, Abidjan, 1968. [16] N. W. Uhl and J. Dransfield, “Genera Palmarum,” Allen Press, Lawrence, 1987. [17] F. White, “The Vegetation of Africa,” UNESCO, Paris, 1983, p. 356. [18] F. Herzog, Z. Farah, and R. Amado, “Chemical Com- position and Nutritional Significance of Wines from the Palms Elaeis guineensis and Borassus aethiopum in the V-Baoulé, Côte d’Ivoire,” Tropical Scie nc e , Vol. 35, No. 1, 1995, pp. 30-39. [19] R. Portères, “Le Palmier Rônier (Borassus aethiopum Mart.) Dans la Province du Baoulé (Côte D’Ivoire),” Jour - nal D’Agriculture Tropicale et de Botanique Appliquée, Vol. 11, 1964, pp. 499-514. [20] P. B. Tomlinson and E. C. Jeffrey, “The Structural Bio- logy of Palms,” Clarendon Press, Oxford, 1990. [21] R. L. Bailey and T. R. Dell, “Quantifying Diameter Dis- tributions with the Weibull Function,” Journal of Forest Science, Vol. 19, No. 2, 1973, pp. 97-104. [22] J. Rondeux, “Les Mesures des Arbres et Des Peuplements Forestiers,” Presses Agronomiques de Gembloux, Gembloux, 1993, p. 521. [23] W. Bonou, R. Glèlè-Kakaï, A. E. Assogbadjo, H. N. Fonton and B. Sinsin, “Characterization of Afzelia africana Sm. Habitat in the Lama Forest Reserve of Benin,” Forest Ecology and Management, Vol. 258, No. 7, 2009, pp. 1084-1092. doi:10.1016/j.foreco.2009.05.032 [24] J. M. Boffa, “Agroforestry Parklands in Sub-Saharian Africa,” Food and Agriculture Organization of the United Nations, Rome, 2000, p. 259. [25] P. L. Giffard, “Le Palmier Rônier Borassus aethiopium,” Bois et forêt des tropiques, No. 116, 1967, p. 14. [26] B. Rollet, “L’architecture des Forêts Denses Humides Sempervirentes de Plaines,” Centre Technique Forestier Tropical, Paris, 1974, p. 298. [27] T. T. Veblen, F. M. Schlegel and R. B. Escobar, “Struc- ture and Dynamics of Old-Growth Nothofagus Forests in the Valdivian Andes,” Journal of Ecology, Vol. 68, No. 1, 1980, pp. 1-31. doi:10.2307/2259240 [28] C. J. Geldenhuys, “The Use of Diameter Distributions in Sustained-Use Management of Forests: Exemples from Southern Africa,” In: G. D. Piearce and D. J. Gumbo, Eds., Proceedings of an International Symposium on the Ecology and Management of Indigenous Forests in Sou- thern Africa, Victoria Falls, 27-29 July 1992, Zimbabwe Forestry Commission and SAREC, Harare, 1992, pp. 154- 167. [29] N. Sokpon, “Recherche Ecologique sur la Forêt dense Semi-Décidue de Pobè au Sud-Est du Bénin: Groupe- ments Végétaux, Structure, Régénération Naturelle et Chu- te de Litière,” Ph.D. Thesis, Universite Libre de Bruxelles, Bruxelles, 1995, p. 350. [30] N. Sokpon, S. H. Biaou, C. Ouinsavi and O. Hunhyet, “Bases Techniques Pour une Gestion Durable des Forêts Claires du Nord-Bénin: Rotation, Diamètre Minimal D’ex- ploitabilité et Régénération,” Bois et Forêts des Tropiques, Vol. 287, No. 1, 2006, pp. 45-57. [31] N. Sokpon and S. H. Biaou, “The Use of Diameter Dis- tribution in Sustained-Use Management of Remnant For- ests in Benin: Case of Bassila Forest Reserve in North Benin,” Ecology and Management, Vol. 161, No. 1-3, 2002, pp. 13-25. doi:10.1016/S0378-1127(01)00488-1 [32] F. Koubouana, “Les Forêts de la Vallée du Niari (Congo): Copyright © 2011 SciRes. AJPS
Ecological Structure and Fruit Production of African Fan Palm (Borassus a ethiop um) Populations Copyright © 2011 SciRes. AJPS 743 Etudes Floristiques et Structurales,” Ph.D. Thesis, Uni- versity of Paris, Paris, 1993, p. 122. [33] W. D. Hawthorne, “Ecological Profiles of Ghanaian Forest Trees,” Tropical Forest Paper 29, Oxford Forestry Institute, Oxford, 1995, p. 345. [34] M. Begon, J. L. Harper and C. R. Townsend, “Life-His- tory Variation,” Blackwell Scientific Publications, Oxford, 1990, pp. 473-509. [35] S. C. Stearns, “The Evolution of Life Histories,” Oxford University Press, Oxford, 1992. [36] J. Cassou and D. Depomier, “In Annonce: Réunion Tri- partie Sur L’agroforesterie,” Sikasso, Burkina, 1997. [37] W. Kperkouma, B. Sinsin, K. Guelly, K. Kokou and K. Akpagana, “Typologie et Structure des Parcs Agrofor- estiers Dans la Prefecture de Donfelgou,” Sécheresse, Vol. 1, No. 3, 2005, p. 8. [38] S. Barot and J. Gignoux, “Population Structure and Life Cycle of Borassus Aethiopum Mart.: Evidence of Early Senescence in Palm Tree,” Biotropica, Vol. 31, No. 3, 1999, pp. 439-448. doi:10.1111/j.1744-7429.1999.tb00386.x [39] Fao, “International Network of Tropical Trees, Partici- patory Management of Forest Resources, Case Study: Com- munity Management of Ecosystems and Conservation of Biodiversity (the Case of Gaya’ Palmyra Plantations),” Technical Paper, 1998, p. 72.
|