Biodiv. Res. Conserv. 29: 63-80, 2013 BR C www.brc.amu.edu.pl DOI 10.2478/biorc-2013-0003 Submitted 27.01.2012, Accepted 31.03.2013 Plant community analysis and effect of environmental factors on the diversity of woody species in the moist Afromontane forest of Wondo Genet, South Central Ethiopia Mamo Kebede1, 2*, Eshetu Yirdaw1, Olavi Luukkanen1 & Mulugeta Lemenih2 Viikki Tropical Resources Institute (VITRI), Department of Forest Sciences, University of Helsinki, Latokartanonkarri 7, P.O. Box 27, 00014 Helsinki, Finland 2 Hawassa University, Wondo Genet College of Forestry and Natural Resources, P.O. Box 128, Shashamene, Ethiopia * corresponding author (e-mail: mamo.kebede@helsinki.fi) 1 Abstract: Floristic diversity and the composition of vascular plants are described for the moist Afromontane forest (MAF) of Wondo Genet, south-central Ethiopia. A total of 75 (20 x 20 m) quadrats were sampled and data on species identity, abundance, elevation, slope and aspect were recorded. Different diversity indices and ordination techniques were used to analyze the data. A total of 240 plant species including seven endemic plant species were found representing 94 families and 210 genera, of which trees constitute 23.8%, shrubs 25%, herbs 35%, lianas 11.3% and ferns 5%. Cluster and indicator species analyses revealed five plant communities described as: Teclea nobilis-Calpurnia aurea, Erythrococca trichogyne-Millettia ferruginea, Croton macrostachyus-Vernonia hochstetteri, Protea gaguedi-Rhus retinorrhoea and Dodonaea angustifolia-Hypericum quartinianum. Elevation (R2=0.48, P<0.001), slope (R2=0.14, P<0.001) and aspect (R2=0.04, P<0.01) correlated significantly and negatively with species richness, whereas only elevation (R2=0.30, P<0.001) and slope (R2=0.13, P<0.001) related significantly and negatively with abundance. Sørensen’s similarity coefficient indicates that the forest of Wondo Genet is similar to moist montane forests of southwestern and southeastern Ethiopia. Given the high diversity, coupled with the existence of endemic species, ecosystem conservation and restoration strategies with further research are warranted. 1. Introduction Ethiopia is endowed with diverse vegetation types ranging from high altitude Afroalpine vegetation in the central highlands to arid lowlands in the East, and rainforests in the West. The altitude of Ethiopia ranges from 125 m b.s.l. to 4533 m a.s.l. and it possesses more land above 2000 m than any other country in Africa (Friis et al. 2010). The highlands that host most of the afromontane vegetation are divided into the Western and Eastern highlands by the East African Rift Valley (Friis et al. 2010). The country has the fifth largest flora in Africa and tremendous floristic diversity, with an estimated 6,500-7,000 species of higher plants of which about 12% are endemic (Gebre-Egziabher 1991; Vivero et al. 2006). According to Friis et al. (2010), there are twelve major vegetation types in Ethiopia, some of these divided into subtypes: (1) Desert and semi-desert scrubland (DSS); (2) Acacia-Commiphora woodland and bushland (ACB); (3) Wooded grassland of the Western Gambella Region (WG); (4) CombretumTerminalia woodland and wooded grassland (CTW); (5) Dry evergreen Afromontane Forest and grassland complex (DAF); (6) Moist evergreen Afromontane Forest (MAF); (7) Transitional Rain Forest (TRF); (8) Ericaceous Belt (EB); (9) Afro-Alpine belt (AA); (10) Riverine Vegetation (RV); (11) Freshwater Lakes, lakeshores, swamps and floodplains Vegetation (FLV); and (12) Salt-water Lakes, lake shores, salt marshes and pan Vegetation (SLV). Topographic and altitudinal variation in Ethiopian landscapes has influenced the existence of varied vegetation types and floristic diversity. Several authors have reported that there is a correlation between floristic ©Adam Mickiewicz University in Poznań (Poland), Brought Department of Plant Taxonomy. All rights reserved. to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM ECOLOGY Key words: Ethiopia, plant community, species richness, Afromontane forest 64 Mamo Kebede et al. Plant community analysis and effect of environmental factors on the diversity of woody... composition and diversity and environmental gradients, such as elevation, slope and aspect (Smith & Huston 1989; Bale et al. 1998; Senbeta & Manfred 2006; Yimer et al. 2006; Fontaine et al. 2007; Woldemariam Gole et al. 2008; Sharman et al. 2009). An altitudinal gradient has an effect on diversity of plant species, which creates variation in climatic pattern and soil differentiation (Lomolino 2001). Studies show that elevation, slope, and aspect are determinants for the spatial and temporal distribution of factors such as radiation, precipitation, and temperature that influence species composition (Albert & Christian 2007). Geographic and climatic conditions change sharply with altitude (Kharkwal et al. 2005) and vegetation in mountain regions responds to small-scale altitude variation (Bale et al. 1998). Similarly, Ovales and Collins (1986) evaluated soil variability across landscapes in two contrasting climatic environments and concluded that topographic position and variation in soil properties were significantly related. In the South-eastern Ethiopian highlands, it has also been studied that in addition to topographic aspect, plant community types are influenced by the physical and chemical properties of the soil (Yimer et al. 2006). Topographic aspect has long been known as a potentially significant factor in generating differences in ecosystem characteristics (Bale & Charley 1994; Bale et al. 1998). Its impacts are various due to its compound character, potentially encompassing external variables such as solar radiation (Holland & Steyne 1975) and cloud cover (Smith 1977). Primary impacts of aspects are expressed through regulating energy budgets and site moisture relationships. Wondo Genet forest, where this study was conducted, is classified as Moist evergreen Afromontane Forest (Friis et al. 2010). In the 1970s, one of the major areas with remnant high forests was the South-central Rift Valley of Ethiopia, including Shashemene, Wondo Genet, and parts of Sidama (Chaffey 1979). The Wondo Genet forest is an upstream forest within the Hawassa watershed, in which over half a million people live (Dessie & Kleman 2007). Like most other forests in the country, Wondo Genet forest is experiencing a large scale deforestation. This forest is severely threatened by heavy anthropogenic disturbance and has declined from 16% of the catchment land area to 2.8%, within the past three decades alone, mainly driven by the expansion of small-scale agriculture, commercial farms and logging. In the decline of the Wondo Genet forest area, two major modes of change were observed: 1) internal, that is, openings created by small farm plots, grazing lands, and villages; and 2) external, that is, expansion of agriculture frontier from the exterior into the forests (Dessie & Kleman 2007). The valleys and mountain slopes of the Wondo Genet escarpment are very heterogeneous and rendered spatially variable plant communities. Analysis and evaluation of the spatial gradients is, therefore, essential to understand the factors affecting species richness, species abundance and plant community types and their distribution (Tesfaye et al. 2008). Several authors have published the results of Ethiopian plant community investigations (Bekele 1994; Woldemariam Gole et al. 2008; Soromessa et al. 2004; Aerts et al. 2006; Senbeta & Manfred 2006; Lulekal et al. 2008; Didita et al. 2010) that have shown a connection of plant community distribution with variation in environmental gradients. Understanding of vegetation composition, diversity of species and their habitats, and comparison with similar other habitats (Afromontane forests), may become a tool to estimate the level of adaptation to the environment. Information on floristic composition, diversity and their relationship with their environment is essential in understanding the forest dynamics. Conservation and management of the forest resources also requires data on plant species diversity and the forest communities in order to check out necessary actions for restoring and rehabilitating this forest. In the face of fast rate of deforestation and degradation, there is an urgency to generate information and assist national and regional action towards maintenance of this forest. The effect of topographic factors and altitude on species richness, abundance and plant community distribution has not been studied in this forest so far. The objectives of the present study are to: (i) describe species richness, abundance and plant community types of Wondo Genet Afromontane forest (ii) ascertain the pattern of species richness and abundance with elevation, slope and aspect, and (iii) carry out floristic similarity (Sørensen’s) comparison of the Wondo Genet forest with other Afromontane forests of Ethiopia and Afromontane forests of Mafi in Tanzania and Mt. Elgon in Kenya. We hypothesize that for the range of altitude and slope studied, floristic diversity increases with altitude, while it decreases with slope steepness, and this is dictated mainly by the moisture gradient. 2. Materials and methods 2.1. Study site Wondo Genet is situated in the southeastern central highlands of Ethiopia, about 263 km from Addis Ababa, at 7o5’30’’N to 7o7’40’’N latitude and 38o36’55’’E to 38 o39’00’’E longitude on the eastern slope of the Rift Valley escarpment (Fig. 1). The large-scale physiographic setting is defined by a tectonic depression bounded by steep escarpments. The floor of the depression is covered by lakes, wetlands and alluvial Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM Biodiv. Res. Conserv. 29: 63-80, 2013 Fig. 1. Location map of the study site and sample quadrats (P) and transects (first transect goes from quadrat P1 to P17, second from P18 to P36, third from P37 to P56 and fourth from P57 to P75) plains, which together cover half of the watershed. The remaining half consists of uplands and escarpments with slopes varying between 8 and 85 degrees. The altitude ranges from 1800-2500 m a.s.l. The mean annual rainfall is about 1200 mm and it is bimodal. Rain can be expected from March to April and June to August. The period from November to February is relatively dry. The mean monthly temperature ranges from 19oC in August to 25oC in March, April, May and September. The soils are young and of volcanic origin, characterized by well-drained loam or sandy loam, and they are shallow at steep convex slopes, but deeper at lower altitudes (Eriksson & Stern 1987). The valley floor is partly covered by lake deposits rich in plant nutrients. On higher ground the texture is sandy or silty, while clay dominates around the wetlands. The current land use is predominantly smallholder agriculture with an average landholding size of less than one hectare per household. The major crops include enset, khat, sugarcane, maize and potatoes. Wondo Genet is agriculturally fertile, with irrigation farming dominating in the flat and undulating sites. The Wondo Genet forest is the partly fragmented remnant of a formerly larger and more coherent forest covering the eastern rift flank (Dessie & Kleman 2007). It harbors important and rare fauna and flora, and provides watershed, ecosystem, economic, research, and educational services. The population of the Wondo Genet is composed of six main ethnic groups consisting of about half a million people (Dessie & Kleman 2007). 2.2. Data collection Systematic plot sampling was conducted in May 2010 in four transects and 75 quadrats of 20x20 m. Many researchers have used similar sample sizes and shapes in the different Afromontane forests in Ethiopia (Bekele 1994; Tadesse & Nigatu 1996; Teketay 1997; Senbeta & Teketay 2003; Senbeta & Manfred 2006; Woldemariam Gole et al. 2008). The first quadrat was located randomly and, afterwards, the quadrats were established at 100 m intervals along transects. Transects were spaced 350 m apart. In each quadrat, all species with DBH (Diameter at Breast Height) ≥2 cm and height ≥2 m were identified and counted. Diameter was measured using a caliper, and a diameter tape was used, when a tree was larger than what a caliper could measure. A Suunto clinometer was used to measure tree height. The presence of epiphytes, herbs, grasses, sedges and ferns was recorded for a floristic compilation. Vernacular names of species mainly in Oromiffa and Sidama local languages in the Wondo Genet area were provided by the key informants. Plant identification was done in the National Herbarium, Addis Ababa University. Nomenclature followed published guidelines of the Flora of Ethiopia and Eritrea (Hedberg & Edwards 1989, Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM 65 66 Plant community analysis and effect of environmental factors on the diversity of woody... Mamo Kebede et al. 1995; Edwards et al. 1995, 1997, 2000; Hedberg et al. 2003). Environmental parameters including slope (using clinometers), elevation (Garmin GPS-72 cross-checked with altimeter), exposition (using Silva compass), and coordinates (using GPS-72) were measured on a plot basis. 2.3. Data analysis Species diversity, cluster analysis, ordinations and phytogeographic comparison methods have been employed to analyse the data. Each of these methods is described in detail below. 2.3.1. Diversity and cluster analysis Species diversity was measured using Shannon diversity (H’), H’max, and Shannon evenness (J’), Simpson index (D) and Simpson evenness indices (Magurran 2004). Hierarchical cluster analysis of the data was done using PC-ORD for Windows version 5 created by McCune and Mefford (2006). Species abundance data was used as input. The Relative Euclidean Distance developed by McCune and Grace (2002) with Ward’s method (hierarchical grouping = minimum variance grouping) was used in order to minimize increases in the error sum of squares. The identified groups were tested for the hypothesis of no difference between two or more groups of entities using Multiple Response Permutation Procedure (MRPP) technique. Moreover, species indicator values were calculated following Dufrene and Legendre (1997). Indicator species analysis was used to contrast performance of individual species across two or more groups of samples. Indicator values are measures of faithfulness (closeness) of occurrence of a species in a particular group and ranges from zero (no indication) to 100 (perfect indication). The statistical significance of the indicator values were tested using Monte Carlo technique. The P-value is based on the proportion of randomised trials with indicator value equal to or exceeding the observed indicator value. In the present analysis, a species with a significant indicator value of P<0.05 is considered to be an indicator species of a community (group). The community types were named after two of the species that had indicator values of P<0.05. coolest slope (northwest) and one – the warmest slope (southeast). Data analysis was based on Nonmetric Multidimensional Scaling (NMDS) technique using Sørensen distance measure. The main advantages of NMDS are following: (1) it avoids the assumption of linear relationship among variables; (2) its use of ranked distances tends to linearize the relationship between distances measured in species space and distances in environmental space; (3) it allows the use of any distance measure or relativization. Gradient analyses were done by employing PCORD version 5.0 (McCune & Mefford 2006). NMDS was run on the log-transformed abundance data using “Autopilot” mode, relative Euclidean distance measure, six starting dimensions, 50 iterations and instability criterion of 10-5. To test for concordance between environmental variables and the NMDS dimension, Spearman rank correlation coefficients were calculated. Monte Carlo test was performed to evaluate whether NMDS extracts stronger axes than expected by chance. To check for the influence of linear regression of environmental gradient on species richness and abundance, “Tree Diversity Analysis software” R-Software (Kindt & Coe 2005) was used, whilst Pearson’s critical value was considered to check the significance level. 2.3.3. Phytogeographic comparison A similarity analysis was carried out to evaluate the relationship between the forests based on the presence of trees and shrubs. Evaluation was conducted using the Sørensen index (Sørensen’s similarity coefficient): Ss = 2a/2a+b+c where: a – number of species common to both forests, b – number of species found only in one forest (here, in Wondo Genet) but absent in the forest under comparison and c – number of species present in the other forest but not in Wondo Genet Forest, and N – number of species entered for comparison. Species data where retrieved from the publications: Ethiopia; Jibat, Chilimo, Menagesha and Wofwasha (Bekele 1994), Mena Angetu (Lulekal et al. 2008), Yayu (Woldemariam Gole et al. 2008); Mafi in Tanzania (Lyaruu et al. 2000) and Mt. Elgon in Kenya (Hitimana et al. 2004). 3. Results 2.3.2. Ordinations Ordination was also computed using a Nonmetric Multidimensional Scaling (NMDS). The species abundance and environmental matrices containing elevation (meters a.s.l.), slope (%), and aspect were used for the ordination. Aspect was measured in degrees and converted to scales from zero to one, following the formula (1-cos (Ø-45)/2, where Ø is aspect in degrees, East of true North, with zero value indicating the 3.1. Floristic composition A total of 240 plant species (herbs, lianas, shrubs, and trees) were identified from the studied quadrats in the Afromontane natural forest of Wondo Genet (Appendix 1). The identified species belong to 94 families and 210 genera, including 8 pteridophyte families, one gymnosperm family, 10 monocotyledon and 75 dicotyledon families. The family with the highest spe- Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM Biodiv. Res. Conserv. 29: 63-80, 2013 cies richness was Asteraceae (17 genera, 22 species), followed by Poaceae (15 genera, 16 species), Fabaceae (12 genera, 13 species), Lamiaceae (10 genera, 10 species), Rubiaceae (9 genera, 9 species), Orchidaceae (7 genera, 7 species) and Euphorbiaceae (6 genera, 6 species). The ten families with the highest species richness contributed 41.25% of the total species and 41.9% of the total genera. In comparison, the 20 families with the highest species richness contributed 55.41% of the total species and 55.71% of the total genera. With respect to plant life forms, trees account for 23.75%, shrubs 25%, herbs 35%, lianas 11.25% and ferns 5% of the species recorded. Species like: Aeollanthus abyssinicus Hochst. ex Benth, Droguetia iners subsp. pedunculata Schweinf, Millettia ferruginea Hochst, Phragmanthera macrosolen (Steud. ex A. Rich.) M.G. Gilbert, Solanecio gigas (Vatke) C. Jeffrey, Tiliacora troupinii Cufod., and Vepris dainellii (Pic. Serm.) Kokwaro, which are endemic to Ethiopia, were also recorded in this forest and constitute 2.9% of the total species recorded. The IUCN Vulnerable species Prunus africana (Hook. f.) Kalkman also inhabits this forest 3.2. Plant community types and the indicator species Five plant communities (clusters) (Fig. 2) with their indicator species (Table 1) were identified for the forest. The communities identified were: Ackokanthera schimperi – Calpurnia aurea community, Erythrococca trichogyne – Millettia ferruginea, Croton macrostachyus – Vernonia hochstetteri, Protea gaguedi – Rhus retinorrhoea and Dodonaea angustifolia Hypericum quartinianum. Ten combinations of pair-wise T-test comparisons were conducted and showed a significant difference (P<0.001). From the analyses, the Fig. 2. Dendrogram of the cluster analysis of species abundance of 72 tree and shrub species found in 75 quadrats (plots). The level of grouping was based on 50 to 30% of information remaining (similarity). The quadrat codes and their arrangment along the dendrogram from top to bottom are as follows: C1: 1, 2, 12, 15, 45, 3, 14, 4, 24, 36, 28, 32, 8, 10, 9, 29, 5, 25, 11, 37, 30; C2: 13, 19, 23, 44, 18, 43, 17, 33, 39, 38, 42, 64, 65, 22, 61, 62; C3: 16, 70, 71, 41, 72, 56, 58, 57, 6, 7; C5: 53, 52, 21, 34, 35, 26, 40, 59,60; C4: 20, 31, 47, 27, 51, 46, 48, 49, 50, 55, 54, 66, 63, 67, 68, 69, 73, 74, 75 Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM 67 68 Plant community analysis and effect of environmental factors on the diversity of woody... Mamo Kebede et al. Table 1. Results of the indicator species analysis Species Probability 1 2 3 4 5 Community I Teclea nobilis Calpurnia aurea Acokanthera schimperi Flacourtia indica Diospyros mespiliformis Allophylus macrobotrys Pittosporum viridiflorum Celtis africana Cassipourea malosana Chionanthus mildbraedii Maytenus arbutifolia Coffea arabica Diospyros abyssinica Afrocarpus falcatus Grewia ferruginea Canthium oligocarpum Acanthus eminens Prunus africana Clutia lanceolata Psydrax schimperiana 0.0002 0.0002 0.0002 0.0002 0.0002 0.0002 0.0008 0.0012 0.0024 0.0036 0.0114 0.0134 0.0212 0.0262 0.0336 0.0630 0.2152 0.2523 0.2581 0.2831 Community II 83 82 68 45 55 53 40 50 42 41 40 33 24 28 22 26 10 8 10 13 3 2 0 0 3 2 2 19 22 2 0 9 2 1 0 1 0 0 0 0 0 0 1 0 2 1 0 7 0 0 17 1 2 0 1 1 0 3 2 7 0 1 0 0 1 0 0 2 1 0 8 0 0 2 0 0 0 0 1 3 0 0 0 0 3 0 0 0 0 0 6 0 0 20 0 10 0 0 0 0 Erythrococca trichogyne Millettia ferruginea Vepris dainellii Pouteria adolfi-friedericii Dracaena afromontana Olea capensis subsp. welwitschii Lepidotrichilia volkensii Cordia africana Dracaena steudneri Fagaropsis angolensis Ehretia cymosa Allophylus abyssinicus Albizia schimperiana Ficus sur Oxyanthus speciosus Ficus vasta 0.0002 0.0002 0.0004 0.0010 0.0026 0.0026 0.0122 0.0366 0.0428 0.0604 0.0830 0.1120 0.1254 0.3873 0.4325 0.4827 Community III 11 1 6 2 0 18 0 0 1 15 0 0 17 0 1 0 58 54 61 46 33 41 27 18 23 24 12 15 23 7 12 6 0 1 0 0 0 1 1 3 9 0 0 1 10 3 7 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 Croton macrostachyus Vernonia hochstetteri Vernonia auriculifera Bersama abyssinica Clerodendrum myricoides Maesa lanceolata Carissa spinarum Maytenus undata Entada abyssinica Acacia abyssinica Oncoba spinosa Hypericum revolutum Ekebergia capensis Polyscias fulva 0.0002 0.0002 0.0008 0.0056 0.0764 0.1036 0.1362 0.2468 0.2627 0.4395 0.5073 0.6255 0.7057 0.9654 Community IV 1 0 0 1 4 0 4 0 0 0 0 5 6 3 6 2 1 12 0 0 0 0 0 0 4 0 0 3 59 43 48 38 19 19 13 10 10 6 8 9 7 4 0 0 0 0 1 12 0 0 0 0 0 2 0 2 1 0 0 0 8 3 3 0 0 4 0 7 1 0 0.0014 0.0030 0.0176 0.0402 0.1140 0.1364 1 0 2 0 0 0 0 0 0 0 0 0 3 3 8 18 6 19 37 35 34 29 16 21 1 5 23 27 0 10 Protea gaguedi Rhus retinorrhoea Myrsine africana Buddleja polystachya Abutilon bidentatum Osyris quadripartita Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM Biodiv. Res. Conserv. 29: 63-80, 2013 Species Premna schimperi Steganotaenia araliacea Probability 1 2 3 4 5 0.7634 0.9678 Community V 7 0 0 0 3 3 8 4 4 0 0.0004 0.0008 0.0046 0.0064 0.0066 0.0122 0.0174 0.0226 0.0834 0.1206 0.2895 0.3439 0.3507 0.5041 0 0 0 0 1 9 0 2 0 0 0 0 6 1 0 0 0 0 0 0 0 0 0 0 1 6 0 0 7 0 5 10 5 2 0 4 5 0 0 0 7 3 1 3 10 29 3 1 16 2 0 0 0 0 7 0 61 35 36 36 29 31 25 24 16 11 9 9 15 6 Dodonaea angustifolia Hypericum quartinianum Syzygium guineense subsp. guineense Rhus vulgaris Olea europaea subsp. cuspidata Schrebera alata Erica arborea Nuxia congesta Olinia rochetiana Syzygium guineense subsp. macrocarpum Ficus thonningii Phoenix reclinata Combretum molle Apodytes dimidiata T-value statistic for the five groups is -27.07 (P<0.001), while the A statistic (chance corrected for within-group agreement) is 0.554. The T-test statistic describes the separation between groups. The A statistics describes within-group homogeneity and falls between 0 and 1. When the items are identical, A=1. In community ecology, values of A are commonly below 0.1. The groups vary in size, ranging from 8 to 21 plots per group. The results of the indicator species analyses determine the degree to which species are associated with the different groups (i.e., communities). Each group has 3-4 indicator species with significant indicator values. In this analysis, a plant species with a significant indicator value at P<0.05 is considered an indicator species of the community. Pair wise comparison of the communities analysed using MRPP indicated T-values ranging from -7 to -20 (P<0.001). Community I (Ackokanthera schimperi – Calpurnia aurea) has twenty species and was found in 21 quadrats (28%). This forest community is the largest, mainly found between the altitudinal ranges of 1880-1990 m a.s.l., mostly, along the natural forest boundary with other land use areas. This community is mainly found along the north-western facing slopes. The slope gradient varies from 18-45%. The community has ten indicator species with significant indicator values (P<0.001); namely, Ackokanthera schimperi Oliv, Calpurnia aurea Benth, Flacourtia indica (Burm.f) Merr., Diospyros mespiliformis Hochst. ex A.DC, Teclea nobilis Delile and Allophylus macrobotrys Gilg. Community II (Erythrococca trichogyne – Millettia ferruginea) comprised sixteen species and sixteen quadrats. This forest community is mainly found between the altitudinal ranges of 1990-2050 m a.s.l. and is mostly distributed along the north-western facing slopes. The slope gradient varies from 15-50%. The community has 9 indicator species with significant indicator values. These are: Erythrococca trichogyne Prain., Millettia ferruginea, Vepris dainellii, Pouteria adolfi-friedericii (Engl.) Baehni, Dracaena afromontana Mildbr., Olea welwitschii Gilg & Schellenb., Lepidotrichilia volkensii (Güerke) J.-F. Leroy, Cordia africana Lam., and Dracaena steudneri Engl. The tree species that are illegally and selectively logged for their timber, namely, Cordia africana and Pouteria adolfi-friedericii, and the endemic tree species Millettia ferruginea and Vepris dainellii are found in this community. Community III (Croton macrostachyus – Vernonia hochstetteri) is represented by fourteen species and ten quadrats. This forest community is mainly found between altitudinal ranges of 1990-2280 m a.s.l., and is mostly distributed along the south-eastern facing slopes. The slope gradient varies from 20-50%. The community has 4 indicator species with significant indicator values, namely, Croton macrostachyus Hochst. ex Delile, Vernonia hochstetteri Sch. Bip. ex Hochst, Vernonia auriculifera Hiern and Bersama abyssinica Fresen. Community IV (Protea gaguedi – Rhus retinorrhoea) is represented by eight species and was found in nine quadrats. The community has 4 indicator species with significant indicator values; namely, Protea gaguedi J. F. Geml., Rhus retinorrhoea Steud. ex Oliv., Myrsine africana L. and Buddleja polystachya Fresen. This forest community is mainly found between altitudinal ranges of 2200-2400 m a.s.l. and along the south-eastern facing slopes. The slope gradient varies from 50-70%. Community V (Dodonaea angustifolia – Hypericum quartinianum) is the second largest community spread Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM 69 70 Plant community analysis and effect of environmental factors on the diversity of woody... Mamo Kebede et al. over nineteen quadrats and represented by fourteen species. This community has eight indicator species with significant indicator values. The indicator species are: Dodonea angustifolia L. f., Hypericum quartinianum A. Rich., Syzygium guineense subsp. (Willd.) DC macrocarpum Engl. macrocarpum, Rhus vulgaris Meikle, Olea europea subsp. cuspidata (Wall. ex G. Don) Cif., Schrebera alata Welw., Erica arborea L. and Nuxia congesta R. Br.. This forest community is mainly found between the altitudinal ranges of 2050-2150 m a.s.l. and along the south-eastern facing slopes. The slope gradient varies from 35-60%. 3.3. Species diversity, richness and evenness of the woody plant communities The overall Shannon-Wiener diversity and evenness of the woody species in the studied forest were 3.63± 0.438 and 0.84±0.10, respectively. The overall values indicate that the diversity and evenness of the woody species is relatively high (Table 2). Apparently, communities II, III, and IV had the highest species diversity, while communities I and V – the lowest. 3.4. Ordination In the NMDS ordination, the greatest reduction in ‘stress’ achieved was 14.71 with a three-dimensional solution and final instability of 1x10-5. The proportions of variance represented by the three axes were 0.288, 0.238 and 0.084, respectively (cumulative r=0.61). Species richness had a significant correlation with AXIS 1 (r=0.229, P<0.05) and AXIS 3 (r=0.567, P<0.01). Species abundance had a significant correlation with AXIS 3 (r=0.400, P<0.01). Elevation had the strongest significant correlation with AXIS 3 (r=0.768, P < 0.01), AXIS 2 (r=0.332, P<0.01) and AXIS 1 (r=0.281, P<0.05). Slope had significant correlation with AXIS 2 (r=0.354, P<0.01), AXIS 1 (r=0.278, Table 2. Species richness, diversity indices and evenness of plant community types Community Elevation Species richness I 1800-2000 51 (6.51) Shannon diversity (H') H' Max Evenness (J) Simpson 2.890 (0.44) 3.95 0.730 (0.11) 0.900 (0.12) II 1850-2000 44 (4.19) 3.026 (0.44) 3.78 0.800 (0.10) 0.932 (0.12) III 1900-2200 61 (7.67) 3.200 (0.42) 4.11 0.778 (0.08) 0.931 (0.07) IV 2200-2400 45 (3.64) 3.017 (0.39) 3.61 0.793 (0.07) 0.926 (0.09) V 2050-2150 37 (4.35) 2.836 (0.43) 3.80 0.785 (0.11) 0.897 (0.09) 1800-2445 72 (6.04) 3.630 (0.43) 4.32 0.840 (0.10) 0.960 (0.10) Over all Explanation: values in bracket are standard deviations Fig. 3. Ordination based on a Nonmetric Multidimensional Analysis (NMDS) with the abundance of 72 woody species from 75 quadrats Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM Biodiv. Res. Conserv. 29: 63-80, 2013 negative correlation with elevation (Fig. 4a-b). In a similar way, slope was correlated with species richness (r2=0.14, P<0.001) and abundance (r2=0.13, P<0.001) significantly negative (Fig. 4c-d). The correlation of aspect with species richness was negatively significant (r2=0.04, P<0.01), but its effect on abundance was not significant (Fig. 4e-f). P<0.01), and AXIS 3 (r=0.245, P<0.01). The correlation of aspect with the axes was not significant. Elevation explained 48.16% of the variation in species richness and 30.14% of species abundance. Slope explained 13.69% of the variations in species richness and 12.46% of species abundance. The percent of variation explained by aspect was quite low, 4.33% and 0.79% of the species richness and species abundance, respectively (Fig. 3). 3.6. Phytogeographical comparison 3.5. Linear relationships of environmental variables with species richness and abundance An attempt was made to compare some Afromontane forests in Ethiopia and Eastern Africa (Tanzania and Kenya) on the basis of similarities in their species composition. Sørensen’s similarity coefficient indicates Both species richness (r2=0.48, P<0.001) and abundance (r2=0.30, P<0.001) have shown significantly a b c d e f Fig. 4. Relationship between elevation (a-b), slope (c-d) and aspect (e-f) and species richness and abundance in the Wondo Genet Forest, south-central Ethiopia Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM 71 72 Plant community analysis and effect of environmental factors on the diversity of woody... Mamo Kebede et al. Table 3. Comparison of the Wondo Genet Forest with five other Ethiopian and two Eastern African montane forests Forest Jibat Chilimo Menagesha Wof-washa Yayu Mena Angetu Mafi (Tanzania) Mt. Elgon (Kenya) N a b c Ss 51 31 31 30 87 117 61 51 35 19 16 19 33 34 13 23 37 53 56 53 39 38 59 49 16 12 15 11 39 35 48 28 0.56 0.36 0.31 0.37 0.45 0.47 0.19 0.37 that the studied forest is similar to moist montane forests of southwestern and southeastern Ethiopia (Table 3). 4. Discussion The inventory presented in this study shows that the Wondo Genet forest is one of the most diverse forests in Ethiopia with respect to plant species richness. Afromontane forests of Ethiopia has been studied by several authors (Bekele 1994; Teketay 1995; Teketay & Bekele 1995; Woldemariam Gole et al. 2000; Friis et al. 1982; Friis 1992; Senbeta & Manfred 2006; Woldemariam Gole et al. 2008). Comparisons of the recorded species richness of the Wondo Genet forest in this study with other published studies of species richness of Afromontane forests, reveal that the level of the diversity in the studied forest is high. With about 240 species, the Wondo Genet Afromontane forest has higher species richness, than, for instance, Jibat forest with 58 species (Bekele 1994), Chilimo forest with 90 (Woldemariam Gole et al. 2000), Dakata forest with 202 (Teketay 1995), Bonga forest with 154 (Friis et al. 1982), the Harrena forest with 128 (Tadesse & Nigatu 1996) and Yayu with 220 (Woldemariam Gole et al 2008). However, Wofwasha forest was reported to have more than 250 species (Teketay & Bekele 1995), so presents an exceptional species richness. With a record of seven endemic species to Ethiopia, Wondo Genet has a relatively high number compared to, for example, Yayu forest with three endemic species (Woldemariam Gole et al. 2008), but lower than Wofwasha, which has 29 species (Teketay & Bekele 1995). Sidamo floristic region, where this forest is located, is among the richest centers of endemism in Ethiopia (Friis et al. 2001). It was noted that endemic plant species account for 11-25% of Afromontane species composition (Woldemariam Gole et al. 2008). In general, the diverse evergreen Afromontane forests have a lower number of unique species (Friis et al. 2010). In this study, five new records were made for the Sidamo floristic region of the Flora of Ethiopia and Eritrea. These are Rhus Source Bekele 1994 Bekele 1994 Bekele 1994 Bekele 1994 Wodemariam Gole et al. 2008 Lulekal et al. 2008 Lyaruu et al. 2000 Hitimana et al. 2004 retinorrhoea, Dracaena afromontana, Erythrococca trichogyne, Sida tenuicarpa and Rubus volkensii. The high diversity of the Wondo Genet forest is probably the result of diverse physiographic nature of this area, with its mountain slopes, valleys and fluvial landforms, in addition to springs/river flowing from the foot of the mountains, made the vegetation unusually diverse. Several vegetation types and associated tree species occur in this forest. From, The Undifferentiated Afromontane forest (DAF/U) (Friis et al. 2010), which is a subtype of the Dry evergreen Afromontane forest and grassland complex (DAF), is clearly represented at Wondo Genet by characteristic species, like Afrocarpus falcatus, Olea europea subsp. cuspidata, Croton macrostachyus, Allophylus abyssinicus, Apodytes dimidiata, Cassipourea malosana, Celtis africana, Millettia ferruginea, Ekebergia capensis, Lepidotrichilia volkensii, Olinia rochetiana, Prunus africana, Vepris dainellii, Teclea nobilis, Pittosporum viridiflorum, Ritchiea albersii and Solanecio gigas. The higher elevation slopes of Wondo Genet harbour some elements of vegetation of the Ericaceous Belt (EB), consisting of scattered trees of Erica arborea and Hypericum revolutum. According to Friis et al. (2010), this is the lower limit of Ericaceous belt (EB) adjoining the Dry Afromontnae forest and grassland complex (DAF). The moist evergreen Afromontane forest (MAF), predominantly broadleaved characteristic species like Pouteria adolfi-fridericii, Albizia schimperiana, Ficus sur, Ficus thonningii, Ekebergia capensis, Cassipourea malosana were identified in the Wondo Genet forest. This is the vegetation type found in southwestern Ethiopia and Harrena forest in the southeast Ethiopia, in Bale Mountains (Friis et al. 2010). Edges of the moist evergreen Afromontane forest, bushland, woodland and wooded grassland (MAF/BW) are represented at Wondo Genet by Acacia abyssinica, Cordia africana, Calpurnia aurea, Maesa lanceolata and Carissa spinarum. The Wondo Genet forest also has the riverine vegetation (RV) type along the outlets of springs into the lower valley. In this type, such characteristic species (Friis et al. 2010) as Syzygium guineense, Oncoba spinosa, Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM Biodiv. Res. Conserv. 29: 63-80, 2013 Diospyros mespiliformis, Salix subserrata and Phoenix reclinata were found in this study. It has been suggested that different elevation and slopes influence species richness, and dispersion behaviour of tree species (Ellu & Obusa 2005). In the Yayu forest, in southwestern Ethiopia, the plant species distribution, and hence the patterns in forest vegetation are mainly influenced by the terrain gradient variables, such as altitude, slope and distance from the river banks (Woldemariam Gole et al. 2008). It was found that altitude was significantly and negatively correlated with density and species richness (Sharma et al. 2009). In the Wondo Genet forest, both species richness and abundance, hence diversity, were significantly and negatively correlated with altitude. A similar pattern for species richness and diversity was reported for the vegetation around Dello Menna in southeastern Ethiopia (Didita et al. 2010). Slope was also significantly and negatively correlated with density and species richness (Sharman et al. 2009). Slope influences drainage, impacting soil formation processes, and chemical properties, since the soils on steeper slopes are influenced by bed rock and tend to be less moist and less acidic (Tewolde 1986, as cited in Woldemariam Gole et al. 2008). At Wondo Genet, slope is significantly and negatively correlated with both species richness and abundance. Aspect-induced regime of fundamental niche characteristics, such as frost, light compensation level and permanent wilting point, enforces some sorting of species (Austin et al. 1990). The magnitudes of inter-aspect differences in the mean monthly temperatures were sufficient to contribute to a sorting of canopy species (Bale et al. 1998). At Wondo Genet, aspect has significantly influenced species richness; however, the influence of aspect on species abundance was not significant. 5. Conclusions The Wondo Genet remnant forest contains a substantial amount of Afromontane plant species composition and diversity. The high diversity, coupled with the presence of endemic species, calls for immediate conservation strategies with the involvement of government and local communities that would lead to the restoration and rehabilitation of this remnant forest. Conserving this forest will allow to preserve refugia for many species and also retain the dispersal pool for the restoration and rehabilitation of the forest itself and the nearby degraded areas under agricultural systems. The further detailed ecological studies concerning the species composition, diversity and distribution of the possible plant community types in relation to other environmental factors such as soil properties, moisture regime, temperature fluctuation, frost occurrence, and the like, which were not the subject of this study, will be of vital importance. Ethnobotanical studies should be conducted to harness the indigenous knowledge on the uses of plant resources contained in the forest. Acknowledgments. We are grateful to the Swedish International Development Agency (SIDA), Center for International Mobility (CIMO, Finland), and International Foundation for Science (IFS Grt. No. D/5053-1) for financial support to the first author. The Wondo Genet College of Forestry and Natural Resources and the University of Helsinki are also acknowledged for offering a postgraduate study opportunity for the first author. We also thank a GIS specialist, Tigneh Eshete, for his assistance in mapping the study site; Gemechu Koroso, our key informant on species local names, and Dr Jennifer Rowland for her assistance in editing this manuscript. We are especially grateful to the anonymous reviewers for their many useful comments and suggestions on improving our manuscript. References AerTs r., VAn oVerTVeLd K., hAiLe m, hermy m. decKers j. & muys B. 2006. Species composition and diversity of small Afromontane forest fragments in northern Ethiopia. Plant Ecol. 187: 127-142. ALBerT r. s. & chrisTiAn s. 2007. Interactions of Elevation, Aspect, and Slope in models of Forest Species Composition and Productivity. Forest Sci. 53: 486-492. AusTin m. p., nichoLLs A. o. & mArguLes c. r. 1990. Measurement of the realized qualitative niche: environmental niches of Eucalyptus species. Ecological Mono. 60: 161-177. BALe c. L. & chArLey j. L. 1994. The impact of aspect on forest floor characteristics in some eastern Australian sites. Forest Ecol. Manage. 67: 305-317. BALe c. L., WiLLiAms B. j. & chArLey j. L. 1998. The impact of aspect on forest structure and floristics in some eastern Australian sites. Forest Ecol. Manage. 110: 363-377. BeKeLe T. 1994. Phytosociology and Ecology of a Humid Afromontane Forest on the Central Plateau of Ethiopia. J. Veg. Sci. 5: 87-98 chAFFey d. r. 1979. Southwest Ethiopia Forest Inventory Project. An Inventory of Forest at Munessa and Shashemene. Project Report 29. Land Resources Division, Ministry of Overseas Development, London. dessie g. & KLemAn j. 2007. Pattern and Magnitude of Deforestation in the South Central Rift Valley Region of Ethiopia. Mountain research and development 27: 162-168. Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM 73 74 Mamo Kebede et al. Plant community analysis and effect of environmental factors on the diversity of woody... didiTA m., nemomissA s. & WoLdemAriAm goLe T. 2010. Floristic and structural analysis of the woodland vegetation around Dello Menna, Southeast Ethiopia. J. For. Res. 21: 395-408. duFrene m. & Legendre p. 1997. Species assemblages and indicator species: the need for a flexible asymmetrical approach. Ecological Mono. 67: 345-366. edWArds s., demisseW s. & hedVerg i. (eds.). 1997. Flora of Ethiopia and Eritrea, vol. 6. National Herbarium, Addis Ababa and Uppsala University, Uppsala. edWArds s., TAdesse m., demisseW s. & hedVerg i. (eds.). 2000. Flora of Ethiopia and Eritrea, part 1, vol. 2. National Herbarium, Addis Ababa and Uppsala University, Uppsala. edWArds s., WoLdemAriAm goLe T. & hedVerg i. (eds.). 1995. Flora of Ethiopia and Eritrea. vol. 2, part 2. Canellaceae to Euphorbiaceae. Addis Ababa and Uppsala University, Uppsala. eLLu g. & oBusA j. 2005. Tree conditions and natural regeneration in disturbed sites of Bwindi Impenetrable forest national park, South-western Uganda. Tropical Ecol. 46: 99-111. eriKsson h. & sTern m. 1987. A soil study at Wono Genet forestry resources institute, Ethiopia. Swedish University of Agricultural Sciences, International Rural Development Centre. Working paper 48, 65 pp., Uppsala. FonTAine m., AerTs r., özAKAn K, merT A., guLsoy s., sueL h., WAeLKens m. & muys B. 2007 Elevation and exposition rather than soil types determine communities and their suitability in Mediterranean mountain forest. Forest Ecol. Manage. 247: 18-25. Friis i. 1992. Forests and forest trees of northeast tropical Africa – their natural habitats and distribution patterns in Ethiopia, Djibouti and Somalia. Kew Bull. Add. Ser. 15: 1-396. Friis i., rAsmussen F. n. & VoLLesen K. 1982. Studies in the flora and vegetation of SW Ethiopia. Opera Botanica 63: 1-70. Friis i., edWArds s., ensermu K. & seBseBe d. 2001. Diversity and endemism in the flora of Ethiopia and Eritrea-what do the published Flora volumes tell us? In: i. Friis & o. ryding (eds.). Proceedings of the 3rd International Symposium on the Flora of Ethiopia & Eritrea. Biol. Skr. 54: 173-193. Friis i., demisseW s. & BreugeL p. V. 2010. Atlas of the potential vegetation of Ethiopia. The Royal Danish Academy of Sciences and Letters, Copenhagen, Denmark. geBre-egziABher T. 1991. Diversity of Ethiopian flora. In: j. m. m. engeLs, j. g. hAWAKes & W. meLAKu (eds.). Plant Genetic Resources of Ethiopia, pp 75-81. Cambridge University Press, Cambridge. hedBerg i. & edWArds s. (eds.). 1989. Flora of Ethiopia, vol. 3. Pittosporaceae to Araliaceae, 659 pp. The National Herbarium, Addis Ababa and Uppsala University, Uppsala. hedBerg i. & edWArds s. (eds.). 1995. Flora of Ethiopia, vol. 7. Poaceae (Gramineae), 420 pp. The National Herbarium, Addis Ababa and Uppsala University, Uppsala. hedBerg i., edWArds s. & nemomissA S. (eds.). 2003. Flora of Ethiopia and Eritrea, vol. 4 (2), Part 1: Apiaceae to Dipsacaceae, 352 pp. The National Herbarium, Addis Ababa and Uppsala University, Uppsala. hiTimAnA j., LegiLisho K. j. & njunge j. T. 2004. Forest structure characteristics in disturbed sites of Mt. Elgon Moist Lower Montane Forest, western Kenya. Forest Ecology Management 194: 269-291 hoLLAnd p. g. & sTeyne d. g. 1975. Vegetation response to latitudinal variations in slope angle and aspect. J. Biogeo. 2: 179-183. KhArKWAL g., mehroTA p., rAWAT y. s. & rico-grAy V. 2005. Distribution of plant life forms along an altitudinal gradient in the Central Himalayan region of India. Current Sci. 89: 873-878. KindT r. & coe r. 2005. Tree diversity analysis. A manual and software for common statistical methods for ecological and biodiversity studies, 207 pp. World Agroforestry Centre (ICRAF) Nairobi. LomoLino m. V. 2001. Elevation gradients of species-density: historical and prospective views. Global Ecol. Biogeo. 10: 3-13. LuLeKAL L., KeLBessA e., BeKeLe T. & yineger h. 2008. Plant species composition and structure of the Mana Angetu moist montane forest, south-eastern Ethiopia. J. East Afri. Nat. Hist. 97: 165-185. LyAruu h. V., eLiApendA s. & BAcKeus i. 2000. Floristic, structural and seed bank diversity of a dry Afromontane forest at Mafai, central Tanzania. Biodivers Conserv 9: 241-263. mAgurrAn A. E. 2004. Measuring biological diversity. 256 pp. Black-well Sciences, Oxford, UK. mccune B. & grAce j. B. 2002. Analysis of Ecological Communities. MjM Software Design, Gleneden Beach. mccune B. & meFFord m. j. 2006. PC-ORD. Multivariate analysis of ecological data, Version 5. MjM Software design, Gleneden Beach. oVALes F. A. & coLLins m. e. 1986. Soil-landscape relationships and soil variability in North Central Florida. Soil Sci. Soc. Ame. J. 50: 401-408. senBeTA F. & TeKeTAy d. 2003. Diversity, community types and population structure of woody plants in Kimphee Forest: a Unique Nature Reserve in Southern Ethiopia. Ethiopian Journal of Biological Society 2: 169-187. senBeTA F. & mAnFred d. 2006. Effects of wild coffee management on species diversity in the Afromontane rainforests of Ethiopia. Forest Ecology and Management 232: 68-74 shArmAn c. m., suyAL s., gAiroLA s. & ghiLdiyAL s. K. 2009. Species richness and diversity along an altitudinal gradient in moist temperate forest of Garhwal Himalaya. Journal of American Science 5: 119-128. smiTh j. m. B. 1977. Vegetation and microclimate of east- and west-facing slopes in the grasslands of Mt. Wihelm, Papua New Guinea. J. Ecol. 65: 39-53. smiTh T. & husTon m. 1989. A theory of spatial and temporal dynamics of plant communities. Vegetatio 83: 49-69. soromessA s., TeKeTAy d. & demisseW s. 2004. Ecological study of the vegetation in Gamo Gofa zone, southern Ethiopia. J. Trop. Ecol. 45: 209-221. Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM Biodiv. Res. Conserv. 29: 63-80, 2013 TAdesse m. & nigATu L. 1996. An ecological and ethnobotanical study of wild or spontaneous coffee, Coffea arabica in Ethiopia. In: L. j. g. VAn der mAesen, x. m. VAn der BurgT & j. m. VAn eedenBAch de rooy (eds.). The Biodiversity of African Plants, pp. 227-294. Kluwer Academic Publishers, Dordrecht. TeKeTAy d. 1995. Floristic composition of Dakata Valley, south-eastern Ethiopia, an implication for the conservation of biodiversity. Mt. Res. Dev. 15: 183-186. TeKeTAy d. 1997. Seedling population and regeneration of woody species in Dry Afromontane Forests of Ethiopia. For. Ecol. Manage. 98: 149-165. TeKeTAy d. & BeKeLe T. 1995. Floristic composition of WofWasha natural forest, Central Ethiopia: implications for the conservation of biodiversity. Feddes Repertorium 106: 127-147. TesFAye g., BeKeLe T. & demisseW s. 2008. Dryland woody vegetation along an altitudinal gradient on the eastern escarpment of Welo, Ethiopia. Eth. J. Sci. 31: 43-54. TeWoLde B. g. e. 1986. Preliminary studies on the ecology of a natural coffee (Coffea arabica) forest with emphasis on coffee. Symb. Bot. Ups. 26: 146-156. ViVero j. L., KeLBessA e. & demisseW s. 2006. Progress on the Red List of plants of Ethiopia and Eritrea: conservation and biogeography of endemic flowering taxa. In: s. A. ghAzAnFAr & h. j. BeenTje (eds.). Taxonomy and ecology of African plants, their conservation and sustainable use, pp. 761-778. Royal Botanic Gardens, Kew. WoLdemAriAm goLe T., TeKeTAy d., edWArds s. & oLsson m. 2000. Woody plants and avian species diversity in a dry Afromontane forest on the central plateau of Ethiopia: biological indicators for conservation. Eth. J. Nat. Res. 2: 255-293. WoLdemAriAm goLe T., Borsch T., denich m. & TeKeTAy d. 2008. Floristic composition and environmental factors characterizing coffee forests in south-west Ethiopia. Forest Ecol. Manage 255: 2138-2150. yimer F., ABdeLKAdir A. & Ledin s. 2006. Soil property variations in relation to topographic aspect and vegetation community in the South-eastern highlands of Ethiopia. Forest Ecol. Manage. 232: 90-99. Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM 75 76 Mamo Kebede et al. Plant community analysis and effect of environmental factors on the diversity of woody... Appendix 1. List of plant species recorded in the indigenous Afromontane forest of Wondo Genet Local name Species Family (Oromiffa & sidama) Habit Distribution type* A. PTERIDOPHYTA Adianthaceae Aspleniaceae Aspleniaceae Aspleniaceae Dennstaedtiaceae Dryopteridaceae Pteridaceae Selaginellaceae Sinoperidaceae Thelypteridaceae Doryopteris concolor (Langsd. & Fisch.) Kuhn. Asplenium bugoiense Hieron. Asplenium mannii Hook. Asplenium theciferum (Kunth) Mett. Pteridium aquilinum (L.) Kuhn. Tectaria gemmifera (Fée) Alston Pteris catoptera Kunze Selaginella abyssinica Spring. Pellaea viridis (Forssk.) Prantl. Christella chaseana (Schelpe) Holttum H H H H H H H H H H B. GYMNOSPERMAE Afrocarpus falcatus (Thunb.) Mirb. Podocarpaceae bibirsa C. ANGOISPERMAE – MONOCTYLEDONAE Chlorophytum Ker Gawl. Arisaema Mart. Phoenix reclinata Jacq. Anthericaceae Araceae Arecaceae Asparagus africanus Lam. Commelina africana L. Commelina erecta L. Carex chlorosaccus C. B. Clarke Cyperus rigidifolius Steud. Scleria bulbifera Hochst. ex A. Rich. Dioscorea bulbifera L. Dracaena afromontana Mildbr. Dracaena steudneri Engl. Asparagaceae Commelinaceae Commelinaceae Cyperaceae Cyperaceae Cyperaceae Discoreaceae Dracaenaceae Dracaenaceae Aerangis brachycarpa (A. Rich.) Durand. & Schinz Disperis anthoceros Rchb. f. Eulophia guineensis Lindl. Habenaria cultriformis Kraenzl. ex Engl. Nervilia crociformis Seidenf. Polystachya Hook. Pteroglossaspis eustachya Rchb.f. Andropogon distachyos L. Aristida adoensis Hochst. Brachiaria brizantha (A. Rich.) Stapf Brachiaria ovalis Stapf Cynodon dactylon (L.) Pers. Digitaria abyssinica (Hochst.) Stapf Eragrostis schweinfurthii Chiov. Exotheca abyssinica Andersson. Hyparrhenia hirta (L.) Stapf Ischaemum afrum (J. F.Gmel.) Dandy Melinis repens (Willd.) Zizka Oplismenus compositus (L.) P. Beauv. Pennisetum thunbergii Kunth Setaria megaphylla T. Durand & Schinz Sporobolus pyramidalis P. Beauv. Themeda triandra Forssk. Orchidaceae Orchidaceae Orchidaceae Orchidaceae Orchidaceae Orchidaceae Orchidaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae adila zenbaba seriti qortobe qortobe dhalladuu yetrara ser boroda lenticho serte digalo gasha serdo muriye T H H T L H H H H H L T T H H H H H H H H H H H H H H H H H H H H H H H DAF/U; MAF/P DAF/WG,MAF/ P,BW,RV,FLU ACB,DAF/WG, EB CTW, RV DAF/U, MAF/P DAF/U-WG, MAF/PBW D. ANGIOSPERMAE – DICOCTYLEDONAE Acanthus eminens C. B. Clarke Hypoestes Sol. ex R. Br. Justicia schimperiana T. Anderson. Acanthaceae Acanthaceae Acanthaceae anshokala dergu dhumuga S H S Cyathula cylindrica Moq. Amaranthaceae garbabo H Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM DAF/U,MAF/P-RV DAF/WG;MAF/BW; RV Biodiv. Res. Conserv. 29: 63-80, 2013 Local name (Oromiffa & sidama) Species Family Aerva lanata (L.) Schult. Celosia schweinfurthiana Schinz Rhus retinorrhoea Steud. ex Oliv. Rhus vulgaris Meikle Alepidea longifolia E. Mey. ex Steud. Heteromorpha arborescens Cham. & Schltdl. Foeniculum vulgare Mill. Pimpinella L. Steganotaenia araliacea Hochst. Acokanthera schimperi (A. DC.) Schweinf. Carissa spinarum L. Amaranthaceae Amaranthaceae Anacardiaceae Anacardiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apocynaceae Apocynaceae kararu agamsa Landolphia buchananii (Hallier f.) Stapf. Polyscias fulva (Hiern) Harms Ceropegia L. Gomphocarpus phillipsiae (N. E. Br.) Goyder Periploca linearifolia Quart.-Dill. & A. Rich. Sonchus L. Acmella caulirhiza Delile. Ageratum conyzoides L. Asplia L. M. A. A. Du Petit-Thouars Athrixia rosmarinifolia Oliv. & Hiern Berkheya spekeana Oliv. Bidens pilosa L. Bidens L. Carduus leptacanthus Fresen. Conyza pyrrhopappa Sch. Bip. ex A. Rich. Crassocephalum Moench Guizotia Cass. Helichrysum schimperi Moeser Plectocephalus varians (A. Rich.) C. Jeffrey in Cufod. Solanecio angulatus (Vahl) C. Jeffrey Solanecio gigas (Vatke) C. Jeffrey Apocynaceae Araliaceae Asclepiadaceae Asclepiadaceae Asclepiadaceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae hopi H yezingero wanbar T H L otisa L hato H yemidr barbere H abadebo H H S qoree H chogogit H H uticho S H H H S/L H J/S shokoko goman T/S Solanecio tuberosus (Sch. Bip. ex A. Rich.) C. Jeffrey Tagetes minuta L. Vernonia auriculifera Hiern Vernonia brachycalyx O. Hoffm. Vernonia hochstetteri Sch. Bip. ex Hochst. Vernonia inulaefolia Steud. Impatiens hochstetteri Warb Cordia africana Lam. Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Balsaminaceae Boraginaceae Cynoglossum coeruleum Hochst. ex DC. Ehretia cymosa Thonn. gagabsa tateesa insilal Habit H H T/S T/S H S H H T T S wadessa H H T/S S S H H T Boraginaceae Boraginaceae maxane ulaga H T Erucastrum arabicum Fisch. & C. A. Mey. Wahlenbergia abyssinica (Hochst. ex A. Rich.) Thulin Ritchiea albersii Gilg Drymaria cordata Willd. ex Schult Hippocratea goetzei Loes. Maytenus arbutifolia (Hochst. ex A. Rich) R. Wilczek Maytenus undata (Thunb.) Blakelock Combretum molle R. Br. ex. G. Don Dichondra repens J. R. Forst & G. Forst. Ipomoea kituiensis Vatke Kalanchoe lanceolata Pers. Momordica foetida Schumach. Zehneria scabra Sond. Diospyros abyssinica (Hiern) F. White Brassicaceae Campanulaceae Capparidaceae Caryophyllaceae Celastraceae Celastraceae Celastraceae Combretaceae Convolvulaceae Convolvulaceae Crassulaceae Cucurbitaceae Cucurbitaceae Ebenaceae siraro loko H S T L L T T/S T H L H H L T Diospyros mespiliformis Hochst. ex A. DC Erica arborea L. Ebenaceae Ericaceae babe asta T/S T/S gimal kital reeji homba kombolcha abalo yayit joro anano hanchura kire Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM Distribution type* ACB,DAF/WG ACB,CTW,DAF/WG DAF/U-SD-WG-TR ACB,CTW,DAF/WG ACB;DAF/SD-TR-WG ACB,DAF/U-SD,MAF/ P,RV MAF/P,TRF DAF/WG;RV DAF/U;DAF/WG; MAF/P-BW;EB DAF/WG;MAF/BW CTW;DAF/WG DAF/WG CTW,DAF/U, MAF/PBW,TRF DAF/U-SD-WG, MAF/P-BW, RV DAF/U, MAF/P MAF/P, TRF DAF/U-WG DAF/U-SD;MAF/P CTW, DAF/WG DAF/WG, ACB DAF/U, MAF/P, TRF, RV RV DAF/U-SD, EB, AA 77 78 Mamo Kebede et al. Plant community analysis and effect of environmental factors on the diversity of woody... Local name Species Family (Oromiffa & sidama) Habit Distribution type* Croton macrostachyus Hochst. ex Delile. Euphorbiaceae bakanisa T/S Clutia lanceolata Forssk. Erythrococca trichogyne Prain. Euphorbia schimperiana Hochst. ex A. Rich Phyllanthus L. Tragia L. Dalbergia lactea Vatke Acacia abyssinica Hochst. ex Benth Acacia brevispica Harms Albizia schimperiana Oliv. Euphorbiaceae Euphorbiaceae Euphorbiaceae Euphorbiaceae Euphorbiaceae Fabaceae Fabaceae Fabaceae Fabaceae muka foni muka kara bingile yagbero lafto ledi sasa S T/S H H H S/L T L/S T CTW, DAF/U, WG, MAF/P-BW DAF/U-SD-WG,EB MAF/P Calpurnia aurea Benth. Fabaceae cheqa S Crotalaria incana L. Desmodium repandum (Vahl.) DC. Entada abyssinica Steud. Eriosema (DC.) Desv. Indigofera L. Millettia ferruginea Hochst. Senna septemtrionalis (Viv.) H. S. Irwin & Barneby Zollernia paraensis Huber. Dovyalis verrucosa Warb. Flacourtia indica (Burm.f) Merr. Oncoba spinosa Forssk. Geranium L. Pelargonium L'Hér. Hypericum peplidifolium A. Rich. Hypericum quartinianum A. Rich. Hypericum revolutum Vahl Hypoxis villosa L.f. Apodytes dimidiata E. Mey. ex Arn. Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae Flacourtiaceae Flacourtiaceae Flacourtiaceae Geraniaceae Geraniaceae Hypericaceae Hypericaceae Hypericaceae Hypoxidaceae Icacinaceae titako dongicho H H T H H T S T T/S T T/S H H H S T H T Aristea abyssinica Pax ex Engl. Hesperantha petitiana Baker Clerodendrum myricoides R. Br. Iridaceae Iridaceae Lamiaceae marachisa H H S Aeollanthus abyssinicus Hochst. ex Benth. Fuerstia africana T. C. E. Fr. Leucas martinicensis (Jacq.) R. Br. Ocimum grandiflorum (Lam.) Pic. Serm. Ocimum lamiifolium Hochst. Plectranthus punctatus L'Hér. Salvia tiliifolia Vahl. Satureja punctata (Benth.) Briq. subsp. ovata (Benth.) Seybold Stachys L. Buddleja polystachya Fresen. Nuxia congesta R. Br. Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Loganiaceae Loganiaceae bulancho bitana H T/S T Phragmanthera macrosolen (Steud. ex A. Rich.) M. G. Gilbert Abutilon bidentatum (Hochst.) ex A. Rich. Sida tenuicarpa Vollesen. Dissotis Benth. Ekebergia capensis Sparrm. Lepidotrichilia volkensii (Güerke) J. -F. Leroy Bersama abyssinica Fresen Loranthaceae tekatila H Malvaceae Malvaceae Melastomataceae Meliaceae Meliaceae Melianthaceae danisa chifreg ononu alayo azamir T S H T T T Stephania abyssinica Walp. Tiliacora troupinii Cufod. Ficus sur Forssk. Menispermaceae Menispermaceae Moraceae kalala lukuta harbu L L T Ficus thonningii Blume. Moraceae dimbicho T/s ganchacha yemidr kolo birbira hagala akuku garamba garmaba chabicha Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM H H H S S H H S DAF/U-WG,MAF/P,RV DAF/WG,MAF/BW ACB,DAF, WG DAF/U-WG;MAF/PBW;TRF DAF/U-WG,MAF/PBW CTW; DAF/WG DAF/U,MAF/P DAF/U-SD,EB ACB;DAF/U-WG;RV CTW, DAF/WG,RV DAF/U-WG DAF/U;EB;AA DAF/U-WG, MAF/PBW, RV CTW,DAF/U-SD-TRWG ACB; DAF/WG ACB, DAF/WG DAF/SD-WG;EB DAF/U-SD-WG;MAF/ P-BW;EB ACB DAF/U-SD;MAF/P-BW DAF/U;MAF/P;RV DAF/U-SD-WG;MAF/ P-BW;RV DAF/U, MAF/P DAF/U, MAF/P, TRF, RV DAF/U, MAF/P, TRF, RV Biodiv. Res. Conserv. 29: 63-80, 2013 Local name Species Family (Oromiffa & sidama) Habit Distribution type* Ficus vasta Forssk Embelia schimperi Vatke Maesa lanceolata Forssk. Moraceae Myrsinaceae Myrsinaceae qilxu kanku abayi T T/L T/S Myrsine africana L. Syzygium guineense (Willd.) DC. subsp. guineense Syzygium guineense (Willd.) DC. subsp. macrocarpum (Engl.) F. White Ochna holstii Engl Chionanthus mildbraedii (Gilg & G. Schellenb.) Stearn Jasminum abyssinicum R.Br. Olea europaea L. subsp. cuspidata (Wall. ex G. Don) Cif. Olea welwitschii (Knobl.) Gilg. & Schellenb. Schrebera alata Welw. Olinia rochetiana A. Juss. Opilia amentacea Roxb. Oxalis obliquifolia Steud. ex A. Rich. Peperomia abyssinica Miq. Peperomia tetraphylla (G. Forst.) Hook. & Arn. Pittosporum viridiflorum Sims Plantago palmata Hook.f. Drynaria volkensii Hieron. Pleopeltis macrocarpa (Bory. ex Wild.) Kaulf. Rumex abyssinicus Desf. Rumex nepalensis Spreng. Lysimachia ruhmeriana Vatke Protea gaguedi J. F. Geml. Clematis hirsuta Guill. & Perr. Myrsinaceae Myrtaceae Myrtaceae qacama dokma dokma T/S T T DAF/WG, RV DAF/U;RV DAF/U-SD-WG;MAF/ BW;RV DAF/U;EB/AA RV, FLU/MFS CTW/DAF/WG Ochnaceae Oleaceae Oleaceae Oleaceae sigida dhala xorsicho waira T/S T/S L T DAF; MAF/P DAF/U;MAF/P DAF/U-WG DAF/U-SD-TR-WG Oleaceae Oleaceae Olinaceae Opiliaceae Oxalidaceae Piperaceae Piperaceae Pittosporaceae Plantaginaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Primulaceae Proteaceae Ranunculaceae T T/S T L H L H T/S H H H H S H T/S L DAF/U;MAF/P DAF/U-SD-TR-WG DAF/U, EB DAF/WG, RV Thalictrum rhynchocarpum Quart. -Dill. & A. Rich. Caylusea abyssinica Fisch. & C. A. Mey Gouania longispicata Engl. Ranunculaceae Resedaceae Rhamnaceae arencho H H L Helinus mystacinus E. Mey. ex Steud. Rhamnus prinoides L'Hér. Rhamnaceae Rhamnaceae galimo gesho L T/S Cassipourea malosana Alston Prunus africana (Hook.f.) Kalkman Rubus niveus Thunb. Rubus steudneri Schweinf. Rubus volkensii Engl. Canthium oligocarpum Hiern Rhizophoraceae Rosaceae Rosaceae Rosaceae Rosaceae Rubiaceae tilo tikurenchet gora gora injori gallo T/S T L S/L L T/S Coffea arabica L. Gardenia ternifolia Schumach. Kohautia platyphylla (K. Schum.) Bremek. Oxyanthus speciosus DC. Pavetta abyssinica Fresen. Pentas lanceolata (Forssk.) Deflers Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae buna T/S T H T/S T/S S Psydrax schimperiana (A. Rich.) Bridson Rubia cordifolia L. Fagaropsis angolensis (Engl.) H. M.Gardner Teclea nobilis Delile Teclea simplicifolia (Engl.) Engl. Toddalia asiatica Lam. Vepris dainellii (Pic. Serm.) Kokwaro Salix subserrata Willd. Osyris quadripartita Salzam. ex Decne. Allophylus abyssinicus Radlk. Rubiaceae Rubiaceae Rutaceae Rutaceae Rutaceae Rutaceae Rutaceae Salicaceae Santalaceae Sapindaceae seged Allophylus macrobotrys Gilg. Dodonaea angustifolia L. f. Pouteria adolfi-friedericii (Engl.) Baehni Sopubia ramosa Hochst Sapindaceae Sapindaceae Sapotaceae Scrophulariaceae walincho dhamaye nole yebrechew ara karkasho inbwacho shult bashanka danshe haso mukabuna sisa hadhessa gao lela karo hirkamo kitkita qeraro Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM T/S H T/S T/S T L T/S T/S T/S T T/S S T H DAF/U, MAF/P-BW DAF/WG DAF/U-SD, MAF, BW, EB DAF/U-WG, MAF/PBW, TRF,RV DAF/WG, RV DAF/U-SD-WG, MAF/P, EB, RV DAF/U, MAF/P, EB DAF/U;MAF/P DAFU-WG ACB/;DAF/TR-WG; MAF/P-BW MAF/P;TRF CTW; DAF/WG DAF/U;MAF/P;RV DAF/SD-WG;RV CTW;DAF/WG;MAF/ P-BW ACB; CTW; DAF/WG DAF/U;MAF/P;TRF DAF/U-SD;MAF/P;RV DAF/U-SD-WG DAF/U-WG;EB;RV DAF/U;MAF/P;TRF DAF/U-WG DAF/U-SU;MAF/PBW;RV MAF/P DAF/U-SD-WG-TR MAF/P 79 80 Mamo Kebede et al. Plant community analysis and effect of environmental factors on the diversity of woody... Local name Species Family (Oromiffa & sidama) Habit Verbascum L. Brucea antidysenterica J. F. Mill. Scrophulariaceae Simaroubaceae gurahare hatawicho H T/S Discopodium penninervium Hochst. Solanaceae meraro T Physalis peruviana L. Solanum giganteum Jacq. Solanaceae Solanaceae awut H T/S Solanum incanum L. Solanum nigrum L. Solanum villosum Mill. Gnidia chrysantha (Sch.) Gilg. Gnidia lamprantha Gilg. Grewia ferruginea Hochst. Triumfetta rhomboidea Jacq. Celtis africana Burm.f. Droguetia iners Schweinf. Girardinia bullosa (Hochst. ex Steud.) Wedd. Girardinia diversifolia (Link) Friis Premna schimperi Engl. Ampelocissus abyssinica Planch. Cayratia gracilis (Guill. & Perrott..) Suess. & Suess Cyphostemma niveum (Hochst ex Schweinf.) Desc. Rhoicissus tridentata (L.f.) Wild. & R. B. Drumm. Solanaceae Solanaceae Solanaceae Thymelaeaceae Thymelaeaceae Tiliaceae Tiliaceae Ulmaceae Urticaceae Urticaceae Urticaceae Verbanaceae Vitaceae Vitaceae Vitaceae Vitaceae tunaye hidi warabesa tunaye ya'aa H H H H S T/S H T H S H T L L/H H S/L dokonu daro qawut dobi sonicho urgessa sariti alqa Distribution type* DAF/U-WG;MAF/PBW DAF/U-SD-WG, EB, AA DAF/U, MAF/BW, TRF, RV CTW, DAF/WG DAF/U-SD-WG DAF/U, MAF/P, RV DAF/WG, RV ACB, CTW, DAF/SDWG Explanations: T – tree, S – shrub, H – herb, L – liana, T/S – tree shrub, T/L – tree liana, S/L – shrub liana; * – for the distribution types see the Introduction part Brought to you by | Kansalliskirjasto Authenticated Download Date | 5/17/16 1:17 PM