IfeJournalofScience vol.14,no.1(2012) 109 IMPACTOFRE-FORESTATIONOFARE-GROWTHSECONDARYFORESTWITH TECTONAGRANDIS (L.)TENYEARSAFTERONUNDERSTORYSPECIES COMPOSITIONANDDISTRIBUTIONINILE-IFE,SOUTHWESTERNNIGERIA

Odiwe,A.I.*,Ogunsanwo,O.andAgboola,O.O. Departmentof Botany,Facultyof Science,ObafemiAwolowoUniversity,Ile-Ife22005,OsunState,Nigeria *Correspondingauthor,e-mail:[email protected] :+2348034394721 (Received:March,2012;Accepted:June,2012)

ABSTRACT Understory species composition, density, diversity, and tree population structure were studied in aTectona grandis plantation 10 years after establishment and compared with a re-growth secondary forest of tropical forest zone in Southwestern Nigeria. The total number of understorey species in the re-growth secondary forest was 28 while 23 species were recorded in theTectona grandis plantation. The numbers of shrub and climber species in the re-growth secondary forest were higher (nine) than the plantation stand (five); however, the number of herbaceous species remained the same in both the plantation and the re-growth secondary forest, The dominant understorey species in the re-growth secondary forest were Asystasia gangetica which got reduced from 70 (26.5%) to 13 (12.3%) in the plantation;Combretum racemosum from 29 (12.2%) to 6 (5.7%); Cyathula achyranthiodes.from 25 (10.5%) to 5 (4.7%); barbata from 17 (7.1%) to 6 (5.7%). The families most abundantly represented in the plantation were Acanthaceae, Combretaceae and Papilionaceae while the family most abundantly represented in the re-growth secondary forest were Acanthaceae, Combretaceae and Amaranthaceae. In the re-growth secondary forest, lower dbh (0 - 20 cm) classes showed higher density while intermediate girth dbh class (41-60 cm) had the higher density in the plantation. In general, the forest stand had more density of trees in each dbh class with a peak in the intermediate girth class. The study reveals that the establishment ofTectona grandis ten years after, in a re-growth secondary forest, has altered the understory composition and structure.

Keywords: SpeciesDiversity;SpeciesComposition;Understorey;GirthSize-Class;Plantation;Re-growthSecondaryForest

INTRODUCTION 1995). Over the years, large areas of tropical A total of 7,895 species from 338 families lowland rainforest in Africa have been cleared for and 2,215 genera have been identified in Nigeria agriculture or converted to plantations of fast (FGN, 2006). Of these, 205 are endemic; the ninth growing exotics (e.g. oil palm, cocoa and rubber). highest number among the African countries. It Land-use change has been projected to have one has also been reported that there are 560 native of the largest global impacts on biodiversity tree species in Nigeria, 16 of which are critically (Chapinet al., 2000; Sala et al., 2000). The number endangered as listed in the IUCN Red list, 18 are of species occurring in different habitats is greatly endangered while 138 are vulnerable. In addition, determined by the type and intensity of land use. out of 4,715 species numbered in This applies particularly to arable land, forest and Nigeria, 205 are endemic while 170 are threatened plantations on which humans have an impact. (IUCN, 2004). The great diversity of plant species Therefore, the need to restore and conserve the found in Nigeria is related to the diversity of species and the ecosystems cannot be over- ecosystems and habitats as well as the tropical emphasized. Many studies have pointed this out climate in the country. However, despite the (Herdberg and Herdberg, 1968; Richardson, 1970; biological richness of Nigeria, these species are UNESCO, 1973; UNEP/FAO, 1975), and faced with threats and extinction. The following particularly in Nigeria (Charter, 1968; Okali, 1975; factors; agricultural activities, bush burning, fuel Ola-Adams and Iyamabo, 1977; Roche, 1973). wood collection, logging, grazing and gathering have been reported as threats and pressure on The conservation of lowland tropical rainforests biological diversity in Nigeria. The massive rate of is critical for the preservation of their biological deforestation is a direct cause of biodiversity loss. diversity and also for their role in climate Nigeria has been reported to have the highest rate stabilization as major carbon stores, for erosion of deforestation of primary forests between 2000 prevention and for global and local water balance. and 2005 (FAO,2005). As part of the effort to restore the lost forest vegetation and address some of the consequences Widespread deforestation and degradation of of deforestation especially on loss of biodiversity, natural habitats have contributed heavily to the wide-scale planting of tree crops like Tectona loss of species (Brookset al. 2002; Pimm et al. grandisand Elaesis guinensis was embarked upon by 110 Odiweetal.: Impactof Re-forestationof aRe-growthSecondaryForest the Authority of the Obafemi Awolowo MATERIALS AND METHODS University, Ile-Ife at various sites of the University Study Area estate 10 years ago (August, 2001) through the The study was conducted at the Obafemi Reforestation Project. Also because of the Awolowo University Estate, Ile-Ife, Osun state, ongoing process of species loss through Nigeria (7° 29' N, 4° 34' E). Ile-Ife lies between deforestation and change in land use, maintaining Latitude 70 32' N and Longitude 40 31' E. The biodiversity has become an important challenge. elevation of Ife ranges from 215 m to 457 m above sea level (Hall, 1969). The site for this study is Plantation and its replacement on the re-growth located on Latitude 070 31.311' N and Longitude secondary forest could have many effects on 040 30.983' E in Obafemi Awolowo University, Ile- natural fauna and flora composition, soil changes Ife. The elevation of the area ranges from 243m to and changes in plant species composition and 274m above the sea level. Figure 1A and 1B shows entrance or biological extinction of some native theTectona grandis plantation and re-growth species (Moraghebiet al., 2005; Pilehvar, 2007; secondary forest sites respectively. Roostami and Pourbabaei, 2007; Vatani, 2006). Studies have shown that plantations can speed The climate of the area is a tropical type with two recovery of biodiversity and promote woody prominent seasons, the rainy and dry seasons. species regeneration on re-growth secondary The dry season is short, usually lasting 4 months forest lands in tropical regions by speeding up from November to March and the longer rainy forest succession processes, increase of soil season prevails during the remaining months. The fertility and improving site conditions (Bernhard- annual rainfall averages 1413 mm yr-1 in a 5-year Reversat, 2001; Cusack and Montagnini, 2004). survey (Duncan, 1974) and it showed two peaks, The proximity to natural forest, seed dispersal one in July and the other in September. The mean characteristics and site qualities influence has been annual temperature ranges from 22.50 C to 31.4 0 C reported to affect regeneration (Koonkhunthod et (Duncan, 1974). The relative humidity in the early al., 2007). morning is generally high, usually over 90% throughout the year. At mid-day, it is rather lower, High species diversity of the forest depended on around 80% in the wet season as low as 50 -60 % in small trees in lower layers. The understory the dry season (Hall, 1969). have been reported to be a major component of The soil of the area is derived from material of forest ecosystems and play an important role in old basement complex which is made up of many ecological functions and processes (Nagaike granitic metamorphosised sedimentary rock (Hall, et al., 2006; Roberts, 2002; Yirdaw, 2001). 1969). The soils are moderately to strongly leached Therefore, conservation of small trees in lower and have low to medium humus content, weakly layers, especially bottom layer, is indispensable for acid to neutral surface layers and moderately to sound maintenance of forests (Ferozet al., 2006; strongly acid sub-soils (Smyth and Montgomery, Hagiharaet al., 2008). The potentials of this tree 1962). The soils which are usually acidic contain crop plantation in maintaining biological diversity less than 10% clay which is mainly kaolinite and are yet to be fully assessed in this part of the world. hence are characterized by low cation exchange capacity and low water holding capacity (Ayodele, The objectives of this paper were to (i) quantify 1986). The soil has been classified as lixisols, and document the understorey species diversity (ii) FAO/UNESCO (1974) and utisols (USDA, examine diameter distributions in this plantation 1975). and compare with a re-growth secondary forest The original vegetation of Ile-Ife is lowland in the vicinity in order to increase our rainforest as climax vegetation (Keay, 1959). The understanding of the recruitment, establishment forest sub-type is dry deciduous forest (Onochie, and succession of woody species following re- 1979). The vegetation of the areas has been forestation. The following key questions were described as the Guineo-Congolian drier forest addressed: (1) How does the understorey species type (White, 1983). Most of the original lowland richness and composition differ in the plantation rain forests are however being massively destroyed and the re-growth secondary forestland? (2) To leaving remnant of re-growth secondary forest what extent has reforestation been able to address scattered around. Tree crop plantations like the issue of biodiversity loss? Theobroma cacao, Cola nitida, Tectona grandis, Elaeis Odiweetal.: Impactof Re-forestationof aRe-growthSecondaryForest 111 guineensis are now common around the area. D =Ó ni (n i -1) N(N-1)

SamplingProcedure Where ni is the total number of individuals Four sample plots 25 m × 25 m, two plots in of species i which are counted/ encountered, and Tectona grandis and two plots in a nearby re-growth N is the total number of all individuals counted/ secondary forest were used for comparison. The encountered. plots were laid out with a measuring tape and demarcated by narrow cut lines. The degree of similarity in species composition between plantations and a nearby re-growth Data Collection and Analysis secondary forest was compared using Jaccard's The measurement was done in March (dry season Index of similarity: month). For each site, all the woody species were Jaccard'sIndex(J)=J/rx100 identified, counted, and their density per plot were Where j is the number of species found in both the determined in each plot. The diameters at breast plantation and the re-growth forest land. height (dbh) = 10 cm of all the identified woody r is the number of species found in only one or the species were measured. Five line transects were other. systematically laid at every 5 m mark in each plot and a quadrant of 50 cm x 50 cm placed at every 5 Data Analysis m point to identify and quantify all the understorey T-test analysis was used to test for significant plant species present in each plot. All the plants different treatment effects (plantation and re- encountered in the quadrants were identified with growth forest) on species diversity, Simpson index the help of IFE Herbarium for proper of similarity and evenness across the plots. identification and also using Flora of West Tropical Descriptive analysis was also employed to explain Africa (Hutchinson and Dalziel, 1954-72). some of the parameters determined. The Floristic composition,densities, diversity, statistical analyses were performed using SPSS distribution and population structure of the trees 13.0 model. and the understorey plants were determined using the following: species richness, diversity indices; RESULTS Shannon-Wiener index, Simpson's Index and Floristic Composition and Richness species evenness (E) (Pieolus index). Results revealed a total of 18 families distributed into 23 species and 23 genera in the plantation and The Diversity index H' of each sampled plot was 21 families distributed into 21 species and 28 calculated using the method prescribed by genera in the re-growth forest. Basal area was Shannon and Wiener index (1963) as: 2 -1 2 -1 17.78 m ha and 10.05 m ha in the plantation and re-growth secondary forest respectively and H= -Ó Pi ln Pi they are significantly different (p = 0.0025). The stem density in the re-growth secondary forest Where Pi is the relative abundance proportion of i -1 species and ln = Natural logarithm was found to be higher (3,808 ha ) than the recorded density in the plantation (1,696 ha-1 ) Eveness index (Pieolus, 1966) also calculated thus: (Table 1). The number of shrub species was higher in the re-growth secondary forest (nine) H E = than in the plantation stand (five), while the In S number of herbaceous species remained similar Where S is the total species number in each (seven) in the two study sites. Two grass species plantation, H is the diversity index and ln= natural (higher in occurrence), five climbers and four tree logarithm. saplings were encountered in the Tectona grandis plantation; while one grass species, nine climbers Simpson Index of diversity (1-D) which gives the and two tree saplings were found in the re-growth probability that two randomly selected individuals secondary forest (Figure 2, Table 2). in a zone belong to different subspecies and D is calculated thus: 112 Odiweetal.: Impactof Re-forestationof aRe-growthSecondaryForest

Figure1:Photographof theStudySites(A)Tectonagrandis Plantationand (B)Re-growthSecondaryForest Odiweetal.: Impactof Re-forestationof aRe-growthSecondaryForest 113 Table1: VegetationParametersMeasuredinTectonaGrandis PlantationandtheRe-growth SecondaryForestatObafemiAwolowoUniversity,Ile-Ife.

VegetationParametersStudySites Tectonagrandis plantation Re-growth secondaryforest Understoryspecies Numbersof Species 23 28 Numberof Families 18 21 Numberof Genus 23 28 Woodyspecies Basalaream2 ha-1 17.78 10.05 Stemdensityha-1 1696 3808

Figure 2: Growth Form of Understory Species Identified in the Re-growth and the Tectona Grandis Plantation at Obafemi Awolowo University, Ile-Ife. 114 Odiweetal.: Impactof Re-forestationof aRe-growthSecondaryForest Table 2: Comparison of Species Composition and Families inTectona Grandis Plantation and Re- growth Secondary Forest at Obafemi Awolowo University, Ile-Ife.

Re-growth Tectonagrandis Speciescomposition Family Forest Plantation Herb Asystaciagangetica Acanthaceae 70(29.41) 13(12.26) Caricapapaya Caricaceae - 2(1.89) Chromolaenaodorata Asteraceae 6(2.52) 30(28.30) Commelinabinnese Commelinaceae - 8(7.55) Commelinabenghalenses Commelinaceae 5(2.10) - Cyathulaachyranthiodes Amaranthaceae 25(10.50) 5(4.72) Nelsonia canescens Acanthaceae 1(0.42) - Phaulopsisbarteri Acanthaceae 1(0.42) - Phaulopsisfacilata Acanthaceae - 2(1.89) Talinumtriangulare Portulaceae 8(3.36) 1(0.94)

Shrubs Alchorneacordifolia Euphorbiaceae - 1(0.94) Allophylus africana Sapindaceae 1(0.42) 1(0.94) Capolobialutea Polygalaceae 1(0.42) - Combretumracemosa Combretaceae 29(12.18) 6(5.60) Euphorbiaspp Euphorbiaceae - 2(1.89) Mallotusoppositifolius Euphorbiaceae 3(1.26) - Microdesmuspuberula Pandaceae 1 (0.42) - Mimosapudica Mimosaceae - 1(0.94) Rauvolfiavomitoria Apocynaceae 1(0.42) - Securinegavirosa Euphorbiaceae 4(1.68) - Sphenocentrumjollyanum Menispermaceae 1(0.42) - Tithoniadiversifolia Compositaceae 1(0.42) - Climbers Baissea subsessilis Apocynaceae 6(2.52) - Cissus arguta Vitaceae 6(2.52) 3(2.83) Discoreabulbifera Dioscoreaceae 6(2.52) 1(0.94) Ipomoeainvolucrata Convolvulaceae 6(2.52) - Jateorhizamicrantha Menispermaceae 6(2.52) - Mondiawhitei Periplocaceae 8 (3.36) 5(4.72) Paulinapinnata Sapindaceae 10(4.20) 2(1.89) Triclisiasubcordata Menispermaceae 1(0.42) - Vignaspp Papilionaceae 8(3.36) 12(11.32)

Treesamplings Blighiaunijugata Sapindaceae 8(3.36) Manihotglaziovii Euphorbiaceae - 1 (0.94) Markhamiatomentosa Bigmonaceae - 1(0.94) Psidiumguajava Myrtaceae 1(0.42) - Tremaorientalis Ulmaceae - 1(0.94) Voacangaafricana Apocynaceae - 1(0.94) Grasses Panicummaximum - 1(0.94) Setariabarbata Poaceae 17(7.14) 6 (5.66) ValuesinBracketStandforPercentageFrequency Odiweetal.: Impactof Re-forestationof aRe-growthSecondaryForest 115 The dominant understorey species in the re- which is more dominant in climbers and shrubs growth secondary forest were AsystasiaGangetica , component as evidenced in Table 2. Results which got reduced from 70 (26.5%) to 13 (12.3%) showed that there were no significant differences in the plantation;Combretum racemosa from 29 between species diversity indices, Shannon wiener (12.2%) to 6 (5.7%);Cyathula Achyranthiodes from 25 (P= 0.2424), Simpson (P=0.4180) while evenness (10.5%) to 5 (4.7 %); Setaria barbata (grass species) index was found to be significantly different at 10 from 17 (7.1%) to 6 (5.7%) (Table 2). The families % (P = 0.0908) level in the plantation areas most abundantly represented in the plantation compared with the re-growth secondary forest. were Acanthaceae, Combretaceae, Papilionaceae while the families most abundantly represented in In the re-growth secondary forest lower dbh class re-growth secondary forest were Acanthaceae, (0-20 cm) showed higher density (Figure 4) than Combretaceae and Amaranthaceae. that of all other dbh girth classes in the plantation (41-100 cm). A trend of straight line relationship Results of diversity measurements in the (R2 = 0.75) between density and diameter understorey species related to Shannon Wiener, (negative linear curve) was observed in the re- Simpson's index and the Evenness measures in growth secondary forest while the plantation plantation and re-growth secondary forest are stand showed a normal distribution relationship shown in Figure 3. Shannon's Species Diversity (R2 = 0.72). In the plantation, trees with Index showed that the re-growth secondary forest intermediate dbh class (41-60 cm) showed highest had higher diversity (H = 2.59) as compared to that density per ha than all other dbh classes. of the plantation stand (H = 2.50). Pielou's Individuals of the lowest dbh class (0-20 cm) Evenness Index revealed that the plantation stand showed higher tree density per hectare in the re- had more consistency in species distribution. Re- growth secondary forest followed by other dbh growth secondary forest had lower evenness index classes in the order of 21-40 cm > 41- 60 cm > 61- (E = 0.47) compared to the plantation stand which 80 cm. In general, the plantation stand showed had higher evenness (E = 0.54; Figure 3). Jaccard's more density of trees than forest in each dbh similarity index between the two stands revealed classes, particularly the intermediate girth class that these stands had similarities to some extent (41-60 cm), except for individuals with lowest (12, 44.4%) in understorey species composition girth class (0-20 cm; Figure 4).

Figure 3: Comparison of Species Diversity Indices Measured across theTectona Grandis Plantation and Re-growth Secondary Forest Study Sites at Obafemi Awolowo University, Ile-Ife. 116 Odiweetal.: Impactof Re-forestationof aRe-growthSecondaryForest

Figure 4: Tree Density-diameter Distribution of Trees in theTectona Grandis Plantation Stand and the Re-growth Secondary Forest Ten Years after Establishment.

DISCUSSION forest, plantation and re-growth secondary forest Species Composition and Richness in Nigeria, have also reported higher undergrowth The aim of this study was to quantify understorey species in the re-growth secondary forest. This species diversity inTectona grandis plantation 10 however, is in contrast to the other studies in years after establishment to restore a re-growth plantation where higher understorey species secondary forest. The higher number of richness has been reported (Areset al., 2010; undergrowth species (richness) recorded in the re- Dodsonet al., 2008; Odiwe et al., 2012; Metlen and growth secondary forest might be related to the Fiedler, 2006; Thysell and Carey, 2001; Wienk et al., level of disturbance and canopy cover at the sites. 2004; Zenneret al., 2006). The higher understorey The re-growth secondary forest is less disturbed species in the plantation has been attributed to compared to plantation (land preparation and the various forms of disturbance like thinning and planting process at the establishment stage), the grazing. re-growth secondary forest is already undergoing successional process and this might have Higher number of climbers and shrub species accounted for the higher understorey species recorded in the re-growth secondary forest recorded at the re-growth secondary forest sites. compared toTectona grandis plantation follows the The canopy cover of the plantation is higher and findings of Ewel and Bigelow (1996) who this might not be favourable to the growth of reported that during the first 5 years of post slash understorey species because of low light and burn succession, there was decrease in availability that can inhibit seedling emergence. herbaceous vines, increase in shrubs and trees and This is consistent with a number of other studies a dramatic increase in epiphytes as the plants aged. which have reported that the overall cover and The result of higher number of shrubs and some biomass of forest understorey vegetation often herbaceous species that were recorded in this dramatically increases with canopy opening study is similar to the findings of Lüet al ., (2011) (Ehrenreich and Crosby, 1960; Stone and Wolfe, and Tchoutoet al . (2006), who found that the 1996). Ogunleyeet al., (2004) in their study on shrub layer was the most species-rich in all the Odiweetal.: Impactof Re-forestationof aRe-growthSecondaryForest 117 strata of the Campo-Ma'an rain forest in similarity of the stands (in terms of resource Cameroon tropical forest. The predominance of availability, similar environmental condition and shrubs and climbers in the re-growth secondary similar mode of propagation and dispersal), forest could be as a result of higher proportion of thereby making it possible for the species to light in the re-growth secondary forest which invade and survive (Davieset al., 2005; Levine, favours them compared with the herbaceous 2000; Stohlgrenet al , 1999; Lalfakawma et al ., plants. From our result, re-growth secondary 2009). forest has more grasses than the plantation, the presence of higher number (frequency) of grasses, Lower common species between plantations and especiallySetaria barbata is an indication of more secondary forest had earlier been reported by canopy openings since grasses can only thrive in an Odiweet al . (2012). Occurrence of more tree open area where more light is available. This species (especiallyBlighia unijugata ) in the observation is consistent with the results of Holl et plantations than the re-growth secondary forest al., (2000) and Hoper et al., (2004) where it was might indicate a potential of using plantations for reported that vines, ferns and persistent grasses restoring biodiversity of tree species in the studied can impede establishment and growth of woody sites. This position must however be treated with shrubs and trees in abandoned pastures. The caution since is not really clear cut and there is dominance of Acanthaceae family in herbaceous need to carry out more studies to ascertain this species is consistent with the findings of Richards position. The higher density of lower dbh class (0- (1996) where Acanthaceae in addition to other 20 cm) in the re-growth secondary forest might families has been reported to be dominant families indicate that the site is highly disturbed and high of ground herbs in tropical rain forests removal of higher girth sizes (Golf and West, 1975; Lorimeret al ., 2001) while the higher Species Diversity intermediate (41-50) dbh classes in the plantation The higher Shannon-wiener index diversity in the could be as a result of the fact that the trees were re-growth secondary forest might be connected planted in the plantation and that they have not with the higher value of species richness. This been harvested since establishment. This is similar result is in agreement with the findings of to the findings of Lalfakawmaet al., (2009), Saxena Ogunleyeet al ., (2004) where higher species et al., (1984), and Souza (1979), where mortality diversity was recorded in the re-growth secondary and lower removal rate across the intermediate forest compared with plantation. Lower species classes has been reported. diversity have also been reported in the plantations in some other studies (Munoz-Reinoso, 2004; Straight line relationship (negative linear curve) Pourbabaei and Roostami, 2007; Roberts, 2002; between the understory species density (plant ha-1 ) Yirdaw, 2001). However, this is in contrast to the and diameter size classes in the re-growth findings of Odiweet al ., (2012), Koonkhunthod et secondary forest is in agreement with the findings al., (2007), Nagaike (2002), Nagaike et al ., (2006); of Lalfakawmaet al. (2009), Rao (1990), Schmeiz Poorbabaei and Poorrrahmati (2009), who have all and Lindsey (1965), while the plantation did not reported higher species diversity in the plantations conform with this relationship (normal compared with the forest. The higher evenness distribution curve). The reason for this has been distribution in the plantation understorey might be explained above. Higher basal area recorded in the linked to the fact that trees are planted at a defined plantation compared to the re-growth secondary spacing and this created a micro-climate forest can be attributed to the trees of bigger sizes environment that might have influenced the that are of uniform girth sizes in the plantation. growth and regeneration of the understorey This contradicts the findings of Poorbabaei and species. Studies where evenness values are higher Poorrrahmati (2009) where a lower basal area in in plantations as obtained in this study are available theTectona grandis plantation was reported. The in literature (Vatani, 2006; Memarianet al., 2007, higher stem density (ha-1 ) recorded in the re- Koonkhunthodet al., 2007). growth forest can be contributed to the higher number of species recorded at the sites. The The presence of certain common species (12) and general pattern of girth size distribution in both higher value of similarity (44.4%) in the plantation Tectona grandis and the re-growth secondary forest and the re-growth secondary forest may be due to is an indication that the sites are disturbed though 118 Odiweetal.: Impactof Re-forestationof aRe-growthSecondaryForest the level of disturbance differs. One can therefore savanna: with special reference to conclude that the study sites are still growing; they eucalypts. Center for International Forestry are not yet matured. Research, Bogor, Indonesia, p. 71. Brooks, T. M., Mittermeier, R. A., Mittermeier, C. CONCLUSION G., Dafonseca , G.A B., Rylands, A. B., It can be inferred from the present study that Konstant, W. R., Flick, P., Pilgrim, J., diversity of understorey species is closely affected Oldfield, S., Magin, G. and Hilton-Taylor, by the land use change, species diversity increases C. 2002. Habitat loss and extinction in the in the re-growth secondary forest compared with hotspots of biodiversity.Conserv. Biol . 16, theTectona grandis plantation. It is obvious that re- 909-923. forestation withTectona grandis plantation has an Chapin F. S, Zavelata E. S, Eviner V. T, Naylor, R. appreciable effect on species richness and L. and Vitousek, P. M. 2000. composition, with less number of shrubs, and Consequences of changing biotic climbers being present. The purpose of carrying diversity.Nature 405, 234-242. out this study is to provide information on the Charter, J. R. 1968. The Conservation of species richness and composition, compared with vegetation and its consistuent species in re-growth secondary forest with the aim of Nigeria. AETFAT Symposium. providing information on the composition and Cusack, D. and Montagnini, F. 2004. The role of distribution of understorey species because little native species plantations in recovery of attention has been paid to the understorey understory woody diversity in degraded vegetation in the process of biodiversity pasturelands of Costa Rica. Forest Ecol. conservation in the tropical forests. As observed, Manage. 188, 1-15. some of the understory species identified in the re- Davies, K. F., Chessons, P., Harrison, S. B Inouye, growth secondary forest have decreased in the D., Melbourne, B. A. and Rice, K. J. 2005. plantation, especially tree species. Spatial heterogeneity explains the scale dependence of the native exotic diversity There is still need for more research to be carried relationship.Journal of Ecology 86, 1602- out, especially to the impact of seasonal changes 1610. on the distribution and composition of the Dodson, E. K., Peterson, D. W. and Harrod, R. J. understorey species and the role of different 2008. Understorey vegetation response to understorey species (shrubs, herbs, grasses, climbers) in nutrient cycling in the systems. thinning and burning restoration treatment in dry coniferous forests of the ACKNOWLEDGEMENT eastern Cascades, USA.For. Ecol. Manage . The authors wish to thank the Curator of IFE 255, 3130-3140 Herbarium, Mr. G. Ibhanesbor and and Mr. B Duncan, E. R. 1974. Weather information from Omomoh of the Department of Botany, OAU, the University of Ife. University Press, Ile- Ife for their assistance in identifying the species in Ife Nigeria. the field and for cross-checking others that cannot Ehrenreich, J.H. and Crosby, J.S. 1960. Herbage be identified in the field in the IFE herbarium. production is related to hardwood crown cover. Journal of Forestry 58, 564-565. REFERENCES Ewel, J. J. and Bigelow, S. W.1996. Plant life-forms Ares, A. Neill A. R. and Puettmann, K. J. 2010. and tropical ecosystem functioning. In Understorey abundance, species diversity Orians, G.H., Dirzo, R. and J. H. Cushman and functional attributes response to (eds.), Biodiversity and Ecosystem Processes in thinning in coniferous stands. For-Ecol Tropical Forests, Springer, New York, pp. Manage. 260, 1104-1113. 101126 Ayodele, O. J. 1986. Phosphorus availability in FAO/UNESCO. 1974. World soil classification. savanna soils of Western Nigeria. Tropical In: Legend to soil Map of the world. Agriculture (Trinodad) 63, 297-300. Volume 1. UNESCO,Paris. Bernhard-Reversat, F. 2001. Effect of exotic tree Food and Agriculture Organization (FAO). 2005. plantations on plant diversity and State of the World's Forests 2005. FAO, biological soil fertility in the Congo Rome, Italy. Odiweetal.: Impactof Re-forestationof aRe-growthSecondaryForest 119 Federal Government of Nigeria (FGN) 2006. Lalfakawma, U. K., Sahoo, F, Roy, S, Draft Report. National Capacity Needs Self- Vanlalhriatpuia, K.. and Vanalalhluna, P. Assessment for Environmental Management. C. 2009. Community composition and Federal Ministry of Environment. tree population structure in undisturbed Feroz, S. M, Hagihara, A. and Yokota, M. 2006. and disturbed tropical semi-evergreen Stand structure and woody species forest stands of North-East India. Applied diversity in relation to stand stratification Ecology and Environment Research 7(4),303- in a subtropical evergreen broadleaf 318 forest, Okinawa Island.J. Plant Res . 119 (4), Levine, J. 2000. Species diversity and biological 293-301. invasions relating local process to Goff, F. G. and West, D. C. 1975. Canopy- community pattern.Science 288, 852-854. understory interaction effects on forest Lorimer, C. G, Dahir, S. E. and Nordhiem, E. V. population structure.Forest Science 21, 98- 2001. Tree mortality rates and longetivity 108. in mature and old growth hemlock- Hagihara A, Feroz, S. M. and Yokota, M. 2008. hardwood forests. Journal of Ecology 89, Canopy Multilayering and Woody Species 960-971. Diversity of a Subtropical Evergreen Lü, X. T, Yin, J. X. and Tang, J. W..2011. Diversity Broad leaf Forest, Okinawa Island. Pacific and composition of understory Sci. 62(3), 363- 376. vegetation in the tropical seasonal rain Hall, J. B. 1969.The vegetation of Ile-Ife . University of forest of Xishuangbanna, SW China. Rev. Ife Herbarium Bulletin 1. Biol. Trop. 5(1). Herdberg, I. and Herdberg, O. 1968. Conservation Memarian, F, Tabari, M, Hoseini, M. and Banej of the vegetation in Africa south of the shafiei, A. 2007. Comparison of woody Sahara.Acta Phytogeogr. Suec. 54, 179-185. species diversity coniferous standard Holl, K. D, Loik, M. E, Lin, E. H. V,and Samuels, I. broad-leaved copoice stand in Kelardasht A. 2000. Tropical montane forest region.Iran J. Environ. 42, 103-108. restoration in Costa Rica: overcoming Metlen, K. L. and Fiedler, C. E. 2006. Restoration barriers to dispersal and establishment. treatment effect on the understoryof Restoration Ecology 8, 339-349. ponderosa pine/Douglas-fir forest in Hooper, E. R., Legendre, P. and Condit, R. 2004. western Montana, USA. For. Ecol. Manage. Factors affecting community composition 222, 355-369. of forest regeneration in deforested, Moraghebi, F, Aliahmadkorori, S, Khanjanishiraz, abandoned land in Panama.Ecology 85, B, Teymori, M. and Hemmeti, A. 2005. 3313-3326. Effects of Eucalyptus plantation on Hutchinson J, Dalziel, J. M, 1954-1972. (Flora of under-story vegetation and some West Tropical Africa Vol. I-III. The characteristics of soil comparing with Whitefriars Press, London, Tonbridge, adjacent forest region in Astara, Iran. J. England. Res. J. S. Iran 13(1(19), 93-109. International Union for Conservation of Nature Munoz-Reinoso, J. C. 2004. Diversity of maritime and Natural Resources (IUCN) (World juniper woodlands.For.Ecol .Manage . Conservation Union). 2004. Red list of 192:267-276. threatened species. IUCN, Gland, Nagaike, T. 2002. Differences in plant species Switzerland. Available from diversity between conifer (Larix kaempferi ) http://www.iucn.org/themes/ssc/redlist. plantations and broad-leaved (Quercus htm crispula) secondary forests in central Japan. Keay, R. W. J. 1959. An outline of the Nigerian For. Ecol. Manage. 168, 111-123. Vegetation (3r d edition) Federal Nagaike, T. Hayashi, A. Kubo, M,.Abe, M, and Government Printer, Lagos, Nigeria. Arai, N. 2006. Plant species diversity in a Koonkhunthod, N, Sakurai, K. and Tanaka, S. managed forest composed of Larix 2007. Composition and diversity of woody kaempferi plantations and abandoned regeneration in a 37-year- old teak (Tectona coppice forests in central Japan. For. Sci. grandis L.) plantation in Northern 52(3), 324-332. Thailand.For. Ecol. Manage . 247,246-254. 120 Odiweetal.: Impactof Re-forestationof aRe-growthSecondaryForest Odiwe, A. I, Olowoyo, J. O. and Ajiboye, O. 2012. Roberts, M. R. 2002. Effects of forest plantation Effects of Land-Use Change on Under management on herbaceous-layer Storey Species Composition and composition and diversity.Can. J. Bot . 80, Distribution in a Tropical Rainforest. Not. 378-389. Sci. Biol. .4(1), 150-156. Roche, L. 1973. The Conservation of the Ogunleye, A. J, Adeola, A. O, Ojo, L. O. and environment in Nigeria; an ecological Aduradola, A. M. 2004. Impact of farming perspective. Paper presented at a activities on vegetation in Olokemeji forest Symposium on the conservation of the reserve, Nigeria.Global Nest: the Int. J . 6 (2), environment in Nigeria 131-140. Roostami, S. T. and Pourbabaei, H. 2007. Study of Okali, D. U. U. 1975. The Case of conservation of plant species diversity in loblolly pine our forest wealth.Obeche 11: 8 -11. (Pinus taeda L.) plantations in the Azizkian Ola-Adams, B. A. and, Iyamabo, D. E. 1977. and Lakanareas, Rasht. Environ. J. Tehran Conservation of natural vegetation in Univ. 33(41), 85-96. Nigeria.Environ. Conserv . 4 (3), 217 - 226. Sala, O.E, Chapin, F. S, Armesto, J. J, Berlow, E, Onochie, C. F. A. 1979. The Nigerian Rainforest Bloomfield, J. and Dirzo, R. 2000. Global Ecosystem: In: Okali; D.U.U (ed.). The biodiversity scenerios for the year 2100. Nigerian Rainforest Ecosystem. Proceedings Science 287, 1770-1774. of the Man and Biosphere Workshop on the Saxena, A. K., Singh, S. P. and Singh, J. S. 1984. Nigerian Rainforest Ecosystem. University of Population structure of forests of Ibadan, 24-26 January, 1979. Pp 1-37 KumaunHimalaya: implications for Pielou, E. C. 1966: The measurement of diversity management. Journal of Environment in different types of biological collections. Management 19, 307-324. J. of Theoretical Biology 13, 131-144. Schmeiz, D. A. and Lindsey, A. A. 1965. Size-class Pilehvar, B. 2007. Concepts of Conservation in structure of old growth forests in Indiana. forest management and biodiversity. J. For. Forest Science 11, 258-264. Range, 75, 92-101. Shannon, C. E. and Wienner, W. 1963. The Pimm, S. L, Russell, G. J, Gittleman, J..L. and Mathematical Theory of Communication. Brooks, T. M. 1995. The future of University of Illinois Press, Urbans, USA. biodiversity.Science , 269, 347-350. pp.125. Poorbabaei, H. andPoorrahmati, G. 2009 Plant Smyth, A. J. and Montgomery, R. F. 1962. Soil and species diversity in loblolly pine (Pinus taeda Land use of Central Western Nigeria, L.) and sugi (Cryptomeria japonica D. Don.) Government Western Nigeria Press, plantation in the western Guilan, Iran. Ibadan Nigeria Internat. J. Biodivers. Conserv. 1(2), 38-44. Stohlgren, T. J, Binley, D, Chong, G. W, Kalkhan, Pourbabaei, H. and Roostami, S. T. 2007. Study of M. A, Shell, L. D, Bull, K. A, Otsuki, Y, plant species diversity in loblolly pine Newman, G, Bashking, M. and Son, Y. (Pinus taeda L.) plantations in the Azizkian 1999. Exotic species invade hot spots of and Lakan areas, Rasht. Environ J Tehran native plant diversity. Ecological Monographs Univ.33(41), 85-96. 69, 25-46. Rao, P, Barik, S. K., Pandey, H. N. and Tripathi, Stone, W. E. and Wolfe, M. L. 1996. Response of R..S. 1990. Community composition and understory vegetation to variable tree tree population structure in a sub-tropical mortality following a mountain pine beetle broad-leaved forest along a disturbance epidemic in lodgepole pine stands in gradient.Vegetatio 88, 151-162. northern Utah.Vegetatio 122, 1-12. Richards, P. W. 1996. The Tropical Rain Forest. Sousa, W. 1979. Disturbance in marine intertidal Cambridge University, Cambridge. boulder fields: the monequillibrium Richardson, S. D. 1970. Gene pools in Forestry. In: maintenance of species diversity. Ecology Genetic Resources in Plants-their 60, 1225-1239. Exploration and Conservation (eds.) Thysell, D. R. and Carey, A. B. 2001. Manipulation Frankel,O. H. and Bennett, E. Blackwell of density of Pseudotsuga menziesii Scientific Publications, Oxford. Pp: 353- canopies: preliminary effects on 365. understorey vegetation.Can. J. For. Res . 31, Odiweetal.: Impactof Re-forestationof aRe-growthSecondaryForest 121

1513-1525. descriptive Memoir to a vegetation Map Tchouto, M. G. P, De Boer, W. F, De Wilde, J. and Africa. UNESCO,Paris. Van der Maesen, L. J. G. 2006. Diversity Wienk, C.L, Sieg, C. H. and McPherson, G. R. patterns in the flora of Campo-Ma'an rain 2004. Evaluating the role of cutting forest, Cameroon: do tree species tell it all? treatments, fire and soil seed banks in an Biodivers. Conserv.15,1353-1374. experimental framework in ponderosa USDA, 1975.Soil . Agricultural pine forests of the Black Hills, South Handbook 436. US Department of Dakota.For. Ecol. Manage . 192, 375-393. Agriculture, Washington. Yirdaw, E. 2001. Diversity of naturally- UNEP/FAO, 1975. The methodology of regenerated native woody species in forest conservation of forest genetic resources - plantations in the Ethiopian highlands. Report on a pilot study. FAO,Rome. New For., 22:159-177.implications for UNESCO, 1973. Final report of export panel on vegetation management.For. Ecol. Manage . Project 8: Conservationof natural areas 79, 29-46. and of the genetic material they contain. Zenner, E. K., Kabrick, J. M, Jensen, R. G., Peck, J. MAB Series No. 12, UNESCO,Paris. E. and Grabner, J. K. 2006. Responses of Vatani, L. 2006. Study of woody species diversity ground flora to a gradient of harvest in planted area with Cupressus sempervirens intensity in the Missouri Ozarks. For. Ecol. L. var horizontalis (Mill) Gord. (Case Manage. 222, 326-334. study:Mazandaran wood and paper low forests). Symposium of natural resources and sustainable development in southern coastal zones of the Caspian sea (Islamic Azad University, Nour Branch, I.R. of Iran). pp. 790-791. White, F. 1983. The vegetation of Africa- a