Effects of Environmental and Disturbance Factors on Plant Community Distribution in Tropical Moist Afromontane Forests, South-West Ethiopia

Semegnew Tadese; Teshome Soromessa; Getaneh Gebeyehu

doi:10.1155/2023/8521303

Effects of Environmental and Disturbance Factors on Plant Community Distribution in Tropical Moist Afromontane Forests, South-West Ethiopia

Tadese, Semegnew;Soromessa, Teshome;Gebeyehu, Getaneh 2023-04-06 00:00:00 Hindawi International Journal of Forestry Research Volume 2023, Article ID 8521303, 17 pages https://doi.org/10.1155/2023/8521303 Research Article Effects of Environmental and Disturbance Factors on Plant Community Distribution in Tropical Moist Afromontane Forests, South-West Ethiopia 1 1 2 Semegnew Tadese , Teshome Soromessa , and Getaneh Gebeyehu Addis Ababa University, College of Natural and Computational Sciences, Centre for Environmental Science, Addis Ababa, Ethiopia Injibara University, Department of Biology, Injibara, Ethiopia Correspondence should be addressed to Semegnew Tadese; semetade@gmail.com Received 1 November 2022; Revised 14 February 2023; Accepted 15 March 2023; Published 6 April 2023 Academic Editor: Anna Zro´bek-Sokolnik Copyright © 2023 Semegnew Tadese et al. Tis is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Tis study was carried out to investigate the efects of environmental and disturbance factors on plant community distribution in the Majang Forest Biosphere Reserve (MFBR) in south-west Ethiopia. A systematic sample design was conducted to collect vegetation and environmental factors in four study sites. In a nested plot design, the vegetation data were collected from 140 main 2 2 2 plots, i.e., 400 m (trees), 25 m subplots (shrubs, lianas, seedlings, and saplings), and 1 m (herbs), respectively. Te plant community classifcation was performed using agglomerative hierarchical cluster analysis (Ward’s Linkage method) in R software (version 4.0.1). Te distribution of plant communities along an environmental gradient was computed using canonical cor- respondence analysis (CCA). In this study, a total of 15 (9.5%) endemic plant species were recorded in MFBR. Four plant community types were identifed, and these were Celtis zenkeri-Blighia unijugata, Pouteria altissima-Lecaniodiscus fraxinifolius, Antiaris toxicaria-Celtis toka, and Dracaena afromontana-Cyathea manniana. Environmental and disturbance factors, such as elevations, slopes, harvesting indexes, soil pH, silt, and herbaceous cover, were the most important for determining plant community distribution in the area. Elevation and slope were found to have a signifcant (P < 0.05) negative and positive relationship with species diversity and richness, respectively. Terefore, the fnding of this study provides baseline information that could be necessary for making further conservation and management in MFBR. required for successful ecological restoration and bio- 1. Introduction diversity protection [7, 8]. Plant ecologists have been successful in defning the Environmental factors have been a cornerstone in the variations in species diversity of communities along evolution of ecological theory [9, 10]. A variety of factors infuence the spatial and temporal patterns of vegetation environmental gradients at various spatial scales [1]. Plant community responses are well related to climatic change, including the physical environment, land use his- factors at regional and global scales [2, 3], whereas at local tory, prior disturbance, and initial vegetation composition or plot scales, topographic and edaphic factors play an [11–14]. Te most important data required to understand important role in controlling community structure and vegetation patterns in forest landscapes are relationships species distribution [4–8]. As a result, environmental between species diversity and environmental factors [15–21]. factors are important not only in confrming plant As a result, the interaction between physical environmental community structure and variability of species distri- variables such as elevations, slopes, and anthropogenic bution at a spatial scale but also for providing insight factors may infuence species diversity and composition into the environmental requirements of the tree species [19, 22–29]. Elevations, slopes, and aspects are determinants 2 International Journal of Forestry Research of the spatial and temporal distribution of elements such as temperatures range between 13.9 and 31.8 C at the Tinishu radiation, precipitation, and temperature that determine Meti weather station. In Ermichi, the average annual rainfall species composition [30]. Soil heterogeneity, such as soil is 2053 mm and the mean monthly minimum and maximum texture, moisture content, electric conductivity (EC), and temperatures range between 11.8 and 29.7 C. Te maximum ° ° pH, creates niches with specifc conditions, which in turn average monthly temperature is between 29.8 C and 31.8 C afect the distribution pattern of plants [31–33]. in February, while the minimum is between 11.9 C and Ethiopia has greater geographic diversity with ten eco- 13.9 C in July for Ermichi and Tinishu Meti, respectively. systems, 18 major and 49 minor agroecological zones [34]. Te maximum rainfall is between April and October, and the Tis diverse physiographic feature is responsible for the minimum rainfall is from November to March in Tinishu existence of a wide range of habitats and is conducive to the Meti and Ermichi (Figure 2). Tis temperature and rainfall survival of various plant and animal species [35]. Accord- variation between two stations is due to variations in the ingly, Ethiopia has a diverse set of plant, animal, microbial, elevation gradient (NMSA 2018). and genetic resources [36]. Ethiopia is recognized as a major Based on the vegetation classifcation of Ethiopia, the centre of diversity and endemism for a number of plant main vegetation types of the Majang Biosphere is montane species in the Horn of Africa region [37, 38]. Te fora of evergreen forests. Besides, the vegetation of this area has Ethiopia’s is diverse, with an estimated 6000–7000 species of diferent categories in terms of life forms such as high higher plants, including 647 (10.74%) endemic taxa [36]. natural forests, bush lands, and grasslands [43]. Some en- Te study of plant community types and factors (soil, vironmental factors, such as elevations, slopes, soil pH, total topographic, and disturbance) that infuence the distribution nitrogen, and phosphorus, vary between each study site in patterns of plant communities are essential inputs for forest MFBR (Table 1). conservation planning and management. For instance, previous studies have reported an association between plant community distributions and variations in environmental 2.2. Vegetation and Soil Data Collection. A systematic gradients in protected forests [25], fragmented forests [39], sampling design was established to arrange quadrats and and community-managed forests of Ethiopia [40]. However, transects as well as to collect vegetation data [44]. Te study there are limited studies on the efects of environmental and area was stratifed into four sites using a digital elevation anthropogenic disturbance factors governing plant com- model (DEM) in Arc GIS software. Te study sites’ polygon munity distribution patterns in southwestern forests [41, 42] was digitized using Google Earth by elevation classes. Tese in general and the Majang Forest Biosphere Reserve (MFBR) were site I (<1200 m.a.s.l), site II (1200–1500 m.a.s.l), site III in particular. Tis problem leads to some research questions: (1500–1800 m.a.s.l), and site IV (>1800 m.a.s.l) (Table 2). A (a) What are the main community types in the Majang total of 140 quadrats were established for vegetation and Forest Biosphere (MFBR)? (b) How do environmental and forest soil data collection, i.e., site I (1–45), site II (46–85), disturbance factors infuence plant community distribution site III (86–115), and site IV (116–140). patterns in Majang Forest Biosphere Reserves? Terefore, to Te quadrats’ X-Y coordinates were generated in Arc answer these questions, we aimed to study to (1) identify the GIS software and loaded to a global positioning system type of plant community in MFBR and (2) determine plant (GPS) receiver for tracking quadrats. Later, a measuring 2 2 community distribution patterns in response to environ- tape was used to lay out 20 × 20 m (400 m ) quadrats in mental and disturbance factors (elevation, slope, distur- each site in the biosphere. Te sampling intervals between bance, and physical and chemical composition of soil) the transect line and the quadrats were 2 km apart [43, 52]. in MFBR. Te sizes of the quadrats were determined based on the 2 2 growth forms of plants [44], i.e., 400 m (tree), 25 m (shrubs and lianas), and 1 m (herbs), respectively, in 2. Materials and Methods a nested plot design (Figure 3). 2.1. Description of the Study Area. Te study was conducted Te tree diameters (DBH ≥5 cm) were determined, and in the Majang Forest Biosphere Reserve (MFBR), situated in their number of stems was counted and recorded in the main the Majang zone, Gambella People National Regional State of plots (400 m ) (Figure 3). Te diameter at breast height and Ethiopia. It has unique biogeography and shares a boundary the height of the individual’s tree species were measured with Sale Nono woreda of Oromia regional state and using a diameter tape, clinometer, and meter tape, re- Anderacha, Yeki, Sheka, and Guraferda woreda of South- spectively. When the branching of multistemmed in- West regional state (Figure 1). It covers a total area of dividuals occurred below DBH, the DBH of each stem was 233,254 ha of forest and agricultural land and rural settle- measured separately and developed into a common diameter ments and towns. MFBR is located between the latitudes of of all stems by summing up and then taking an average ° ° 07 08′00″N and 07 50′00″N and the longitudes of diameter. To determine the diversity and estimate the ° ° 34 50′00″E and 35 25′00″E, with elevations ranging from abundance of shrubs and lianas, subplots (area � 25 m each) 562 m to 2444 m. were established (Figure 3) [44]. Te climate in the area is generally hot and humid, which Seedlings with a height of less than 1.30 m and a DBH of is marked on most rainfall maps of Ethiopia as the wettest less than 2.5 centimeters, saplings with a height of more than part of the country. Te average annual rainfall is 1774 mm, 1.30 m and a DBH of 2.5 to 5 cm, and trees as plants with and the mean monthly minimum and maximum a DBH of more than 5 cm were selected [53]. Herbaceous International Journal of Forestry Research 3 34°50′0″E35°0′0″E35°10′0″E 35°20′0″E Ethio-Region Gambella Regions- Zone 0 5 10 20 km 34°50′0″E 35°0′0″E 35°10′0″E 35°20′0″E Elevation (m) <1200 1500-1800 1200-1500 >1800 Figure 1: Location of the study sites (sites I–IV). Source: https://earthexplorer.usgs.gov. 1277 m 22.1°C 1774 mm 1582 m 20.5°C 2053 mm °C mm °C mm 50 100 50 100 40 80 40 80 29.8 31.8 30 60 30 60 20 40 20 40 11.9 13.9 10 20 10 20 0 0 0 0 J F MA M J JA S ON D JF M A M J JA SO N D Months Months (a) (b) Figure 2: Mean monthly temperature and rainfall recorded. (a) Tinishu Meti (1987–2017) and (b) Ermichi (1987–2017) (NMSA 2018). Temprature 7°10′0″N 7°20′0″N 7°30′0″N 7°40′0″N Temprature 7°10′0″N 7°20′0″N 7°30′0″N 7°40′0″N 4 International Journal of Forestry Research Table 1: Topographic and soil characteristics of the study sites. Study sites Ele (m) Slo Area (ha) SP Site I 1042 ± 42 5.3 ± 0.4 22,826.1 40 Site II 1365 ± 24 5.4 ± 0.4 25,220.5 45 Site III 1635 ± 24 7.2 ± 0.5 14,053 30 Site IV 2011 ± 42 11.1 ± 1.2 11,783.5 25 Note. Ele � elevation, Slo � slope, and SP � sample plots. Table 2: List of equations used for calculation of vegetation parameters. Vegetation parameters Equation Equation no. Reference Density D � n/N 1 [45] √�� Species richness D � n/ N 2 [46] H H � 􏽐 (Pi) (Inpi) 3 [44, 47] i�1 ′ ′ Shannon evenness J � H /Hmax � H /In s 4 [44, 47] SSI S � (2a/(2a + b + c)) 5 [44, 48] Beta (β) diversity β � (b + c/(2a + b + c)) 6 [49, 50] Harvesting index HI � 􏽐 (SD∗ EF/LD 7 [51] i�1 “a” � number of tree species common to sites A and B; “b” � number of tree species recorded only in the site A; “c” � number of tree species recorded only in site B, “N”: total number of individuals of all the species; “n”: total number of individuals of the species; Shannon diversity index (H) � where pi is the th proportion of individuals found in the i species; SSI � Sorensen similarity index, H � Shannon diversity index; SD � stump density; EF � expansion factor (3.4 for tropical forest); LD � live tree density. University. Nomenclature of plants in this article follows 20 m those published in the fora of Ethiopia as well as fora of Ethiopia and Eritrea [57–62]. 5 m Soil samples were taken with a soil auger from the topsoil at a depth of 0–30 cm, which is recommended as the default 5 m sampling depth for soil [63]. Te soil samples were taken from 1 m fve diferent locations in each plot, four from the quadrat’s 20 m 1 m corners and one from the quadrat’s centre. A total of 220 soil samples, 140 from forestland, 40 from farmland, and 40 from grassland, were collected, composited separately, labelled, and transported to the laboratory. To determine soil bulk density, soil was collected on the centre of the quadrats using a stainless core sampler, then placed in plastic bags, and transported to the laboratory for dry weight determination. Fresh wet soil weights Figure 3: Sampling design used for data collection [44]. Note: were measured in the feld with a kitchen balance with 0.1 g subplots (1 × 1 m) are measured for herbs, grasses, and soil samples; precision. A composite sample of 200 g was taken from each subplots (5 × 5 m) are measured for shrubs and lianas, and the main quadrat to analyze its chemical composition [64] in the Water plot (20 × 20 m) is measured for trees, saplings, and seedlings. Works Design and Supervision Enterprise (WWDSE) labora- tory in Addis Ababa. cover abundance was assessed in fve subquadrats (1 m ) within the main plot (400 m ). 2.3. Environmental and Disturbance Data Collection. Te herbaceous cover abundance from each quadrat was Environmental factors such as elevations, slopes, and aspects changed into 9 cover abundance [54] scale classes: were measured and recorded for each of 140 quadrats using 1 � (<0.5%), 2 � (0.5–1.5%), 3 � (1.5–3%), 4 � (3–5%), clinometers and Garmin GPS, respectively. Te elevation 5 � (5–12.5%), 6 � (12.5–25%), 7 � (25–50%), 8 � 50–75%, was coded into four elevation ranges, namely, and 9 ≥ 75% [55]. Canopy openness was measured by using 1 ≤1200 m.a.s.l, 2 �1200–1500 m.a.s.l, 3 �1500–1800 m.a.s.l, a densiometer located at the centre of each plot. We classifed and 4 ≥1800 m.a.s.l [65]. Aspect values were collected and canopy openness by the following classifcation scheme: very arranged (N � 0, NE � 1, E � 2, SE � 3, S � 4, SW � 3.25, dense canopy >70%, moderately dense canopy 40–70%, W � 2.5, and NW � 1.25) in each quadrat [52, 66]. Te slope open canopy 10–40%, scrub <10%, and nonforest 0% [56]. range was classifed into three major slope classes [67]. As Plant species were identifed in the feld, and for those a result, the classes were as follows: (1) fat <10, (2) in- species that were difcult to identify in the feld, voucher termediate 10–20, and (3) steep >20. specimens were collected with the help of local guides, Human disturbance (which includes harvesting/cutting coded, pressed, and dried for subsequent identifcation and trees for fuel wood, charcoal, timber, and house construction verifcation at the National Herbarium (ETH), Addis Ababa illegally) was computed as the harvesting index. Te International Journal of Forestry Research 5 Shannon evenness index (J) were calculated using equations harvesting index was computed by counting individual stumps that were illegally logged trees within the quadrats. It 3 and 4 in Table 2. Te Sorensen similarity index was used to assess the degree of foristic similarity between plant com- is calculated from the density of individual tree stumps by following equation (7) in Table 2 [51]. Stumps are a small munities. Te Sorensen similarity index was calculated using portion of the trunk that remains after a tree with about 5 cm equation 5 in Table 2. Beta diversity was computed by the chopped down [68]. change in diversity of species from one community to an- other and calculated using equation 6 in Table 2 [49]. Pearson correlation coefcients were calculated to see the 2.4. Data Analysis relationship between diversity indices and environmental disturbance factors in MFBR. 2.4.1. Cluster Analysis. Te vegetation data, which were originally estimated in the feld as a percentage of cover abundance values, were converted into Domin cover 2.4.3. Soil Laboratory Analysis. Te soil samples were an- scales prior to analysis [54]. Te vegetation data were alyzed in the Water Works Design and Supervision En- examined using an agglomerative hierarchical cluster terprise (WWDSE) laboratory in Addis Ababa, Ethiopia. Te (AHC) analysis in package cluster to classify vegetation Bouyoucos hydrometer method was used to determine soil into plant community types. Te similarity ratio (SR) textures. Soil pH was determined using a pH meter a 1 : 2.5 with Ward’s group linkage methods (minimum variance soil to water suspension potentiometric method [72]. Te clustering) was used for cluster analysis. Te similarity micro-Kjeldahl [73] and Walkley and Black [74] methods ratio is one of the similarity indices that determine how were used to determine total nitrogen (N) and soil organic similar or dissimilar vegetation samples, quadrats, or carbon, respectively. Te Bray-I method was used to de- plots are. Ward’s group linkage method evaluates cluster termine available phosphorus, and the absorbance of the distances using the analysis of variance methodology Bray-I extract was measured in a spectrophotometer at [52]. Te goal of this strategy is to eliminate diferences in 882 nm [75]. Based on C and N concentrations, the carbon- total abundance between sample units (quadrats). In to-nitrogen ratio (C/N) was calculated. Te dry weight of addition, Ward’s method minimizes the total within- each soil sample (MS) was determined using oven-drying set group mean of squares or residual sum of squares [69]. to 105 C for 24 h to achieve a constant weight [76]. Te Te maximum number of clusters in a data set is volume of the core sampler (VC) was determined as VC � π a central issue in partitioning clustering, such as k-means r h, where r is the radius and h is the height of the core clustering, which requires the user to specify the number sampler (VC � 3.14 × (2.5 cm) 2 × 5 cm � 98.125 cm ) to of clusters k to be created. In this study, the optimum calculate the bulk density for each sample. number of clusters was identifed using the gap statistic method and determined by plotting the gap statistic against the number of clusters (k-means). Te point, 2.4.4. Ordination. Te canonical correspondence analysis where the maximum gap statistic in the plot is, de- (CCA) ordination method was used to fnd out plant termines the proper number of clusters. In this study, the community distribution along the environmental gradient maximum gap statistic value in the plot is shown at k- [77]. Te selection of CCA was based on the results of means � 4 (the maximum number of clusters or detrended correspondence analysis (DCA), which showed communities) [70]. that the longest axis of DCA for a data set was greater than 3 Characteristic species were determined using syn- (�4.88) (Table 3). Tis indicates the presence of higher optic table analysis. Plant community types were named β-diversity or heterogeneous vegetation data due to the after two distinctive species with high synoptic cover- unimodal relationship between species and environmental abundance ratings [44]. Indicator species analysis was variables. Permutational multivariate analysis of variance performed using the package labs in R software. Te (function adonis test) was used to choose environmental signifcance of the indicator value was tested using the factors that were relatively more relevant in explaining the Monte Carlo test (P < 0.05) [71]. Te indicator values species data and signifcance before CCA ordination [78]. were calculated from the product of relative abundance Pearson’s correlation coefcient was calculated to evaluate (specifcity) and relative frequency of species (fdelity) the relationship between environmental factors and plant within a community [71]. In this study, the species with diversity. the highest signifcant indicator value is considered an indicator species of a community. 3. Result 2.4.2. Community Diversity Analysis. Species richness, 3.1. Plant Diversity. In this study, a total of 15 (9.5%) en- evenness, Shannon diversity, and beta diversity indices were demic plant species were recorded from Ethiopia. Of the computed using R software (version 4.0.1). Species richness total endemic species, 9 are herbaceous, 3 are shrubs, 2 are is usually expressed as the number of species per sample unit trees, and 1 is a liana in life form (Table 4). Rinorea friisii and and calculated using equation 2 in Table 2 [49]. Shannon’s Crotalaria rosenii were listed as near threatened among the index was computed using abundance and evenness of the 15 endemic species, while eight others were listed as least species present. Te Shannon diversity index (H′) and the concerned (Table 4). 6 International Journal of Forestry Research Table 3: Detrended correspondence analysis result of MFBR vegetation. DCA axes DCA1 DCA2 DCA3 DCA4 Eigen values 0.8356 0.2267 0.1917 0.1600 Decorana values 0.8599 0.2441 0.1845 0.1449 Axis lengths 4.8857 2.2693 2.4418 2.2889 3.2. Plant Community Types. Four plant community types Table 4: Endemic plants, their habits, and IUCN status in the were identifed using agglomerative hierarchical cluster Majang Forest Biosphere Reserve. analysis of the entire vegetation data set of MFBR (Figure 4 IUCN and Table 5). Te identifed communities were Celtis Plant species Family name LF category zenkeri-Blighia unijugata community (C1), Pouteria Acanthopale aethiogermanica altissima-Lecaniodiscus fraxinifolius community (C2), Acanthaceae NA S Ensermu Antiaris toxicaria-Celtis toka community (C3), and Dra- Bothriocline schimperi Oliv. & caena afromontana-Cyathea manniana community (C4). Asteraceae LC H Hiern ex Benth Clematis longicauda Steud. ex C1: Celtis zenkeri-Blighia unijugata community Ranunculaceae LC L A. Rich Tis community was found in the elevation range of Crotalaria rosenii (Pax) Fabaceae NT H 830–1896 m.a.s.l and was represented by 77 species. Of Milne-Redh. ex Polhill these, 67 species are commonly shared with other Dorsetnia soerensenii Friis Moraceae NA H communities, while 12 are found only in this com- Euphorbia dumalis S. Carter Euphorbiaceae LC H munity type. Among the total species, four species were Impatiens rothii Hook.f Balsaminaceae NA H signifcant indicator species, with higher indicator Millettia ferruginea (Hochst.) Bak Fabaceae LC T Pycnostachys abyssinica Fresen Lamiaceae NA H values than those of total species (Table 6), namely, Rinorea friisii M.G.Gilbert Violaceae NT S Blighia unijugata, Pouteria alnifolia, Margaritaria Solanecio gigas (Vatke) C. Jefrey Asteraceae LC S discoidea, and Garcinia buchananii. Te most domi- Solanum marginatum L.f Solanaceae NA H nant shrub species included Argomuellera macrophylla, Vepris dainellii (Pic. Serm.) Whitfeldia elongata, Dracaena fragrans, Vernonia Rutaceae LC T Kokwaro amygdalina, and Phyllanthus reticulatus. Leptaspis Vernonia fligera Oliv. & Hiern Asteraceae LC H zeylanica, Aspilia mossambicensis, Asplenium Vernonia leopoldi (Sch. Bip. ex Asteraceae LC H bugoiense, Asplenium anisophyllum, and Pollia con- Walp.) Vake densata were the dominant herb layers in the com- Note. LF � life form, T �tree, S � shrub, H � herb, and L � liana; IUCN threat munity. Te common lianas of this community were categories (LC � least concern, NA � not available, and NT �near Embelia schimperi, Combretum paniculatum, Dregea threatened). schimperi, and Goupia glabra. C2: Pouteria altissima-Lecaniodiscus fraxinifolius community herb layer was dominated by Asplenium bugoiense and Aspilia mossambicensis. Tis community was distributed in the elevation range of 800 to 2310 m.a.s.l. and was represented by 77 C3: Antiaris toxicaria-Celtis toka community species. Of these, 71 species are commonly shared with Tis community was found in the elevation range of other communities, while 6 are found only in this 850–2300 m.a.s.l and was represented by 105 species. community type. Among the total species, four species Of these, 92 species are commonly shared with other were signifcant indicators with higher indicator values communities, while 13 are found only in this com- (Table 6), namely, Pouteria altissima, Lecaniodiscus munity type. Among the total species, four species are fraxinifolius, Ficus exasperata, and Ritchiea albersii. signifcant indicator species with higher indicator Moreover, the dominant tree species layers were values (Table 6), namely, Celtis zenkeri, Morus meso- Lannea welwitschii, Ritchiea albersii, Grewia molli, zygia, Vernonia hochstetteri, and Celtis toka. Moreover, Ficus sur, Ficus mucuso, Ficus ovata, Ficus exasperata, the dominant species in the tree layer were Diospyros Ficus umbellata, Allophylus macrobotrys, and Croton abyssinica, Morus mesozygia, Combretum molle, Apo- sylvaticus. Erythrococca trichogyne, Whitfeldia elon- dytes dimidiata, Olea capensis, Dombeya torrida, gata, Dracaena fragrans, Rinorea friisii, Vernonia Buddleja polystachya, and Lepisanthes senegalensis. Te amygdalina, Acalypha orbata, Pittosporum viridi- dominant species in the shrub layer were Vernonia forum, Phyllanthus reticulatus, and Oxyanthus spe- hochstetteri, Acanthopale pubescens, Erythrococca ciosus were prominent shrubs in this community. trichogyne, Plectranthus sp, and Capparis tomentosa. Jasminum dichotomum, Combretum paniculatum, Marantochloa leucantha, Solanum nigrum, Achyr- Phytolacca dodecandra, Dregea schimperi, Symphytum anthes aspera, and Asplenium aethopicum were the ofcinale, Pseuderanthemum tunicatum, Uncaria afri- dominant species in the herb layer. Te dominant li- cana, and Saba comorensis were common lianas. Te anas in this community were Peponium vogelii, International Journal of Forestry Research 7 Table 5: Synoptic cover value of plants in MFBR for species ≥1% in at least one community. Scientifc name C1 C2 C3 C4 Cluster size 36 25 44 35 Celtis zenkeri 3.450 2.120 2.659 0.000 Blighia unijugata 2.806 1.040 1.841 0.000 Apodytes dimidiata 2.444 2.000 1.364 0.000 Pouteria alnifolia 2.222 1.240 1.000 0.000 Trichilia prieuriana 1.694 1.200 0.864 0.000 Margaritaria discoidea 1.694 0.640 0.818 0.000 Argomuellera macrophylla 1.611 1.320 1.000 0.000 Diospyros abyssinica 1.556 0.800 1.955 0.000 Lannea welwitschii 1.500 1.600 1.409 0.000 Combretum molle 1.389 0.600 0.432 0.000 Pouteria altissima 2.472 3.400 2.864 0.000 Lecaniodiscus fraxinifolius 1.472 3.220 1.955 0.000 Erythrococca trichogyne 1.361 2.040 0.750 0.943 Apodytes dimidiata 2.444 2.000 1.364 0.000 Ritchiea albersii 0.944 1.840 1.227 0.000 Ficus exasperata 1.000 1.760 0.364 0.000 Antiaris toxicaria 1.778 1.640 3.532 0.000 Celtis toka 1.417 1.200 2.909 0.000 Morus mesozygia 0.556 0.480 1.477 0.000 Croton macrostachyus 0.472 0.800 1.045 1.171 Deinbollia kilimandschrica 0.250 1.040 1.045 0.000 Grewia mollis 1.222 1.400 1.159 0.000 Ficus sur 0.222 1.520 1.136 0.943 Dracaena afromontana 0.000 0.000 0.000 3.514 Cyathea manniana 0.000 0.000 0.000 3.414 Schefera abyssinica 0.000 0.000 0.000 2.800 Trilepisium madagascariense 0.000 0.000 0.000 2.686 Galiniera saxifraga 0.000 0.000 0.000 2.600 Schefera myriantha 0.000 0.000 0.000 1.914 Vernonia bipotini 0.000 0.000 0.000 1.886 Allophylus abyssinicus 0.000 0.000 0.000 1.857 Pouteria adolf-friedericii 0.000 0.000 0.000 1.829 Clausena anisata 0.000 0.000 0.000 1.829 Note. C1 � community one, C2 � community two, C3 � community three, and C4 � community four. 5.0 4.5 4.0 3.5 3.0 2.5 2.0 1.5 1.0 0.5 0.0 Plots Figure 4: Dendrogram of the cluster analysis results of species abundance found in 140 of MFBR. C1: 1, 17, 5, 21, 8, 24, 14, 30, 31, 38, 45, 61, 86, 90, 78, 98, 7, 23, 10, 26, 15, 50, 66, 43, 59, 33, 47, 63, 40, 56, 74, 94, 83, 103, 77, and 97; C2: 2, 18, 9, 25, 13, 29, 3, 19, 11, 27, 72, 92, 85, 105, 76, 96, 4, 20, 89, 12, 28, 75, 95, 82, and 102; C3: 6, 22, 35, 16, 54, 70, 36, 53, 69, 42, 58, 49, 65, 71, 91, 79, 99, 87, 80, 100, 84, 104, 88, 32, 39, 55, 46, 62, 48, 64, 52, 68, 73, 93, 81, 101, 34, 37, 44, 60, 41, 57, 51, and 67; C4: 106, 126, 110, 130, 119, 139, 112, 132, 116, 136, 123, 109, 129, 120, 140, 117, 137, 114, 134, 125, 107, 127, 118, 138, 111, 131, 121, 115, 135, 124, 108, 128, 113, 133, and 122. Dissimilarity 8 International Journal of Forestry Research Table 6: Higher indicator values with signifcance of species in the 3.2.1. Species Richness, Evenness, and Diversity among Plant clusters. Community Types. Te overall Shannon diversity index and evenness in MFBR were 3.64 and 0.94, respectively (Table 7). Te Species Community R. Ab R. Fr Indval highest Shannon diversity was shown in community 1 (3.8), value while the least was exhibited in community 4 (3.5). Te max- Blighia unijugata C1 0.493 0.778 0.384 0.001 imum Shannon evenness was indicated in community type four Pouteria alnifolia C1 0.498 0.667 0.332 0.001 (0.9), followed by community types three, two, and one (each Margaritaria C1 0.537 0.583 0.314 0.001 with 0.7 evenness) (Table 8). Te highest species richness was discoidea Garcinia buchananii C1 1.000 0.306 0.306 0.001 exhibited in community type one (102 species), whereas the Pouteria altissima C2 0.283 0.722 0.389 0.001 lowest plant species richness was revealed in community type Lecaniodiscus four (70) (Table 7). C2 0.225 0.278 0.343 0.001 fraxinifolius Similarity coefcients of all communities range from 0.61 Ficus exasperata C2 0.320 0.361 0.293 0.001 to 0.91. Te highest Sorensen similarity was exhibited be- Ritchiea albersii C2 0.235 0.306 0.275 0.001 tween communities 1 and 2 (0.91) followed by communities Celtis zenkeri C3 0.841 0.365 0.307 0.005 2 and 3 (0.90). Te lowest Sorensen similarity was revealed Morus mesozygia C3 0.523 0.588 0.307 0.001 between communities 1 and 4 (0.61) and communities 2 and Vernonia hochstetteri C3 0.341 0.799 0.272 0.001 4 (0.63) of the total species in MFBR (Table 8). Celtis toka C3 0.636 0.422 0.268 0.005 Sorenson similarity coefcients and beta diversity were Dracaena C4 1.000 1.000 1.000 0.001 afromontana inversely related to each other. Terefore, communities with Cyathea manniana C4 1.000 1.000 1.000 0.001 the lowest similarity coefcients had the highest beta di- Vernonia bipotini C4 1.000 0.914 0.914 0.001 versity (0.45) in communities 1 and 4, whereas communities Schefera abyssinica C4 1.000 0.886 0.886 0.001 with the highest similarity coefcients had the lowest beta Note. R.Ab � relative abundance, R.Fr � relative frequency, diversity (0.102) (Table 8). Indval � indicator values in the class, and indicator species in the respective community types at P < 0.05. 3.2.2. Species Richness, Evenness, and Diversity along Envi- ronmental and Disturbance Factor. Species diversity varied Ampelocissus bombycina, Sericostachys scandens, from 2.33 to 3.64; the higher value was recorded at the higher Tacazzea apiculata, Paullinia pinnata, Lagenaria elevation gradient (>1800 m.a.s.l), and the lower value was siceraria, and Cayratia gracilis. recorded at the lower elevation gradient (<1200 m.a.s.l). Species richness and Shannon diversity exhibited a signifcant positive C4: Dracaena afromontana-Cyathea manniana correlation with elevations (r� 0.302, p � 0.0002) and community (r� 0.493, p � 0.00005), respectively, while evenness showed Te community is distributed in the elevation range of a signifcant negative correlation with the elevation gradient 1820–2359 m.a.s.l and was represented by 70 species in (r� −0.385, p � 0.00003) (Table 9). Similarly, the slope showed 35 plots. Of these, 35 species are commonly shared with a signifcant positive correlation with species richness (r � 0.262, other communities, while 35 are found only in this p � 0.00002) and species diversity (r� 0.373, p � 0.00001), community type. Among the total species, four species while the slope showed a signifcant negative correlation with are signifcant indicator species with higher indicator species evenness (r� −0.181, p � 0.0005) (Table 9). values (Table 6), namely, Dracaena afromontana, Te highest species diversity was shown on the in- Cyathea manniana, Vernonia bipotini, Schefera termediate slope (10–20) gradient, while the lowest diversity abyssinica, and Pouteria adolf-friedericii. Te domi- recorded the lower slope (<10). Te herbaceous cover nant tree species in the community were Allophylus showed a signifcant positive correlation with species rich- abyssinicus, Pouteria adolf-friedericii, Triumfetta ness and Shannon diversity (r � 0.137, p � 0.01) and tomentosa, Pycnostachys eminii, Dracaena refexa, (r � 0.171, p � 0.004), respectively. Species richness and Ekebergia capensis, Ilex mitis, Elaeodendron buchana- Shannon diversity showed a signifcant negative correlation nii, Albizia schimperiana, Dracaena afromontana, with the harvesting index (r � −0.243, p � 0.003) and Cyathea manniana, Schefera abyssinica, Trilepisium (r � −0.376, p � 0.00004), respectively. Similarly, soil pH and madagascariense, Galiniera saxifrage, and Schefera silt showed a signifcant negative relationship with species myriantha. Justicia schimperiana, Acanthopale aethio- richness (r � −0.321, p � 0.0001) and (r � −0.504, germanica, Clausena anisata, Vernonia auriculifera, p � 0.0004) and Shannon diversity (r � −0.302, p � 0.0002) Maesa lanceolata, Maytenus gracilipes, Clematis long- and (r � -0.458, p � 0.00001), respectively. However, soil icauda, and Vernonia bipotini were the dominant pH and silt showed a signifcant positive relationship with species in the shrub layer. Te dominant species in herb species evenness (r � 0.365, p � 0.0009) and (r � 0.353, layer were Alchemilla fscheri, Bothriocline schimperi, p � 0.00001), respectively (Table 9). Justicia sp, Asplenium friesiorum, Setaria megaphylla, Asplenium sandersonii, and Physalis angulata. Te dominant lianas in this community were Clematis 3.3. Relationship between Community Types with Environ- mental and Disturbance Factors. Te length of the frst axis simensis, Culcasia falcifolia, Hippocratea pallens, and Jasminum abyssinicum. of DCA indicates species heterogeneity in vegetation as International Journal of Forestry Research 9 Table 7: Species richness, evenness, and diversity indices of plant (r � 0.0445, p � 0.001), elevation (r � 0.0906, p � 0.001), community types. slope (r � 0.0143, p � 0.014), pH (r � 0.0127, p � 0.032), and silt (r � 0.0114, p � 0.0047) exhibited a signifcant Community type Vegetation parameters diference among environmental and disturbance factors C1 C2 C3 C4 (Table 10). Tus, signifcant environmental and distur- Species richness 102 77 93 70 bance factors infuence the species distribution pattern in H′ 3.8 3.6 3.7 3.5 all community types. Shannon evenness 0.7 0.7 0.7 0.9 2 −1 CCA ordination was performed using data on cover Basal area (m )/ha 58.55 48.82 64.64 76.3 −1 abundance values of plant species in the plots and six sta- Density (ha ) 1532 1018 1478 1309 −1 tistically signifcant environmental disturbance factors. In Seedling density (ha ) 3349 3087 4431 3804 −1 the canonical correspondence analysis (CCA), the eigen- Sapling density (ha ) 1629 1457 1789 1509 values obtained in the frst and second axes were 54.4 and Note. H′ � Shannon diversity index, C1 � community 1, C2 � community 2, C3 � community 3, and C4 � community 4. 4.4, respectively. Te correlation results of environmental disturbance factors and the CCA axis showed both a positive and negative correlation from the frst to the last axis Table 8: Pairwise comparison of Sorensen’s similarity coefcient (Table 11). Te elevation and slope showed a positive cor- and beta diversity among four plant communities. relation (r � 0.868, p < 0.05) and (r � 0.638, p < 0.05), re- spectively, in the frst axis. Among signifcant environmental Communities C1 C2 C3 C4 disturbance factors, the elevation was an extremely im- C1 1 portant constraining factor followed by the slope and har- C2 0.91 (0.102) 1 C3 0.87 (0.13) 0.90 (0.104) 1 vesting index (HI) on plant species distribution patterns in C4 0.61 (0.45) 0.63 (0.43) 0.64 (0.38) 1 MFBR. On the other hand, pH and silt showed a strong Note. Values outside the bracket indicate Sorensen’s coefcient, while those negative correlation (r � −0.907, p < 0.05) and (r � 0.792, inside the bracket indicate the beta diversity index. C1 � community 1, p < 0.05), respectively, in the frst axis. CCA showed that the C2 � community 2, C3 � community 3, and C4 � community 4. cumulative proportion and proportion explained in the frst axis accounted 76.3%, respectively, which is explained as species and environmental disturbance factor variations (Table 11). Table 9: Pearson correlation coefcients between diversity indices Ordination of the study plots of MFBR formed four and environmental and disturbance factors in MFBR. communities based on the species composition. Tese Environmental disturbance Diversity indices four community types were separated by following the site SR H′ J arrows of the environmental variables. Red, green, blue, factors and purple represent sample plots of community types 1, ns ∗ ∗ Hca 0.137 0.171 −0.030 ns 2, 3, and 4, respectively (Figure 5). Community four ∗∗∗ ∗∗∗ HI −0.243 −0.376 −0.040 ∗∗∗ ∗∗∗ ∗∗∗ generally occurs at a higher elevation, while species in Ele 0.302 0.493 −0.385 ∗∗∗ ∗∗∗ ∗ communities one, two, and three are distributed at the Slope 0.262 0.373 −0.181 ∗∗∗ ∗∗∗ ∗∗∗ middle and lower elevations. Te distribution of species in Soil pH −0.321 −0.504 0.365 ∗∗∗ ∗∗∗ ∗∗∗ communities four was infuenced by elevation, slope, Silt −0.302 −0.458 0.353 harvesting index, and herbaceous cover factors, whereas Note. Hca � herbaceous cover, HI � harvesting index, Ele � elevation, SR � species richness, H′ � species diversity index, and J � species evenness. the distribution of species in communities one, two, and Positive signs indicate a positive correlation, and negative signs indicate three was strongly infuenced by soil pH and silt factors ∗ ∗∗ ∗∗∗ inverse relations. p < 0.05, p < 0.01, p < 0.001, and ns � no (Figure 5). Te length and direction of the vectors in the signifcance. plots and environment and disturbance biplot represent the degree of correlation between the plots and envi- a result of environmental variables (Table 3). Te frst axis of ronmental disturbance factors. Communities’ types one, DCA was> 3 (4.88), which indicates a wider length of the two, and three are more closely related to one another, gradient and the presence of higher β- diversity or het- whereas community type four is diferent from the other erogeneous vegetation data due to the unimodal relationship three community types (Figure 5). between species and environmental factors (Table 3). Te correlation results between environmental and Terefore, CCA was selected to show the efect of envi- disturbance factors exhibited both positive and negative ronmental and disturbance factors on the patterns of plant relationships in MFBR (Table 12). Te elevation showed community distribution. a signifcant positive correlation with slopes (r � 0.44), MC Furthermore, signifcant environmental and distur- (r � 0.58), sand (r � 0.33), and TN (r � 0.43), while it showed bance factors (P < 0.05) were selected using permuta- a signifcant negative relationship with pH (r � −0.76), silt tional multivariate analysis of variance (function adonis (r � −0.90), and clay (r � −0.91). Similarly, the slope showed test). Out of 14 environmental and disturbance factors, a signifcant negative relationship with MC (r � −0.38), six factors were signifcant in explaining patterns of plant pH (r � -0.46), silt (r � −0.42), TN (r � −0.47), and clay community distribution. More specifcally, herbaceous (r � −0.46), while it showed a signifcant positive correlation cover (r � 0.0409, p � 0.001), harvesting index with sand (r � 0.45). Soil pH showed a signifcant positive 10 International Journal of Forestry Research Table 10: Result of the function Adonis test of signifcant envi- ronmental and disturbance factors in MFBR. Sums of Mean Variables Df F model R Pr(>F) squares squares ∗∗∗ Hca 1 1.8900 1.8898 6.9218 0.0409 0.001 CaOp 1 0.1770 0.1766 0.6470 0.0038 0.767 ∗∗∗ HI` 1 2.0610 2.0608 7.5481 0.0445 0.001 ∗∗∗ Ele 1 4.1910 4.1912 15.3514 0.0906 0.001 Slope 1 0.6600 0.6600 2.4173 0.0143 0.014 Aspect 1 0.4640 0.4640 1.6994 0.0100 0.08 -3 MC 1 0.2000 0.1998 0.7316 0.0043 0.703 pH 1 0.5860 0.5863 2.1473 0.0127 0.032 Sand 1 0.2280 0.2284 0.8367 0.0049 0.559 -6 Silt 1 0.5290 0.5293 1.9388 0.0114 0.047 Clay 1 0.3360 0.3365 1.5142 0.0631 0.441 -1 012 TN 1 0.4140 0.4135 1.5146 0.0089 0.137 CCA1 OM 1 0.2900 0.2898 1.0614 0.0063 0.369 Communities P 1 0.1750 0.1748 0.6403 0.0038 0.814 a Community 1 a Community 3 Residuals 126 34.4000 0.2730 0.7436 Total 139 46.2640 1.0000 a Community 2 a Community 4 Note. Hca � herbaceous cover, CaOp � canopy openness, HI � harvesting Figure 5: Canonical correspondence analysis (CCA) ordination index, Ele � elevation, MC � moisture content of the soil, TN � total ni- diagram of 140 plots and 6 signifcant environmental variables and trogen, OM � organic matter, and P � available phosphorus. plot number distribution in four community types. forests of Ethiopia may be due to ecological and geographical Table 11: Scores of constraining variables and correlations between environmental and disturbance factors with CCA axes in MFBR. separations [84]. Based on the plant species endemic to Ethiopia, Rinorea friisii and Crotalaria rosenii were listed as Environmental variables CCA1 CCA2 CCA3 CCA4 near threatened, while other eight were registered as least Hca 0.429 −0.128 0.160 −0.541 concerned (Table 4) [85]. Tis suggests that special attention HI 0.522 0.028 0.207 0.335 needs to be given to the forest stand containing species Ele 0.868 0.066 0.100 −0.014 which are threatened and least concerned. Slope 0.638 −0.238 0.135 −0.280 Te overall diversity investigation of MFBR showed the pH −0.907 0.047 −0.029 −0.020 higher Shannon diversity index (3.64) and evenness (0.94). Silt −0.792 0.060 −0.187 −0.243 Te value of species diversity and evenness in MFBR is Eigenvalue 0.544 0.044 0.040 0.033 Proportion explained 0.763 0.062 0.057 0.046 greater than that of Gesha-Sayilem forest, Kibate forest, Cumulative proportion 0.763 0.825 0.882 0.928 Gerba Dima forest, Wurg forest, and Agama forest, while it Note. Hca � herbaceous cover, HI � harvesting index, Ele � elevation, and is lower than that of Belete forest (Table 13). Tis diference MFBR � Majang Forest Biosphere Reserve. may be due to links with geographical locations, climatic conditions, and elevation factors. relationship with silt (r � 0.91), TN (r � 0.93), and clay 4.2. Plant Community Types. In this study, a total of four (r � 0.91), while it showed a signifcant negative relationship plant communities were identifed, which are rich in species with sand (r � −0.93) (Table 12). composition. Te major distinguishing features of the identifed plant communities were the diference in domi- 4. Discussion nant and indicator plant species. However, communities 1–3 4.1. Endemic Plant Species and Diversity. Te Majang Forest share more species (>70 species) than community 4 (35 Biosphere Reserve had shown high endemism as proposed species), which could be directly related to environmental by the World Conservation and Monitoring Centre (com- and disturbance factors that make the plant communities have their own distinct or characteristic species [42]. Almost prises high or 9.5% endemic species) (Table 4) [79]. Tus, endemism is higher than reported in other moist Afro- all communities identifed more than four indicator species with a signifcant value. Indicator plant species are plants montane forests of southwestern Ethiopia. For instance, Bonga forest recorded 13 endemic species (5.35%) [80] and that are easily monitored and predict the condition of the Yayu forest with only 3 endemic species (1.36%) [81]. On the environment where they originated. Indicator species can other hand, the number of endemic species recorded in this also be a sign of a distinctive set of environmental qualities or study is lower than in some dry Afromontane forests of characteristics found in a specifc place [91]. Ethiopia. For example, Wof Washa forest has 29 species Blighia unijugata and Pouteria alnifolia were indicator (12%) [82]; Chilimo forest has 18 endemic species (8.45%) species in community 1. Blighia unijugata is the dominant [83]. Te lower number of endemics in the moist Afro- and indicator species and is among the indicator species of montane forests of the southwestern and dry Afromontane moist Afromontane forests in the middle canopy [92, 93]. CCA2 International Journal of Forestry Research 11 Table 12: Pearson’s correlation coefcient matrix for environmental and disturbance factors (N � 14) in MFBR. Clay Hca CaOp HI Elevation Slope Aspect MC pH Sand Silt TN OM P (ns) Hca CaOp 0.27 ns ns HI 0.13 0.05 ns ∗∗ ∗∗∗ Elevation 0.23 0.10 0.33 ns ns ∗ ∗∗∗ Slope 0.15 −0.07 −0.28 0.44 ns ns ns ns ns Aspect −0.08 0.08 −0.03 −0.09 −0.11 ns ns ∗ ∗ ∗∗∗ ∗∗∗ MC 0.18 −0.05 −0.14 0.58 −0.38 −0.09 ∗∗ ns ∗∗∗ ∗∗∗ ∗∗∗ ns ∗∗∗ pH −0.28 −0.08 −0.32 −0.76 −0.46 0.09 −0.57 ns ns ∗∗ ∗∗∗ ∗∗∗ ∗∗∗ ∗∗∗ ∗∗∗ Sand −0.27 0.08 0.31 0.33 0.45 −0.06 −0.47 −0.93 ∗∗ ns ∗∗ ∗∗∗ ∗∗∗ ns ∗∗∗ ∗∗∗ ∗∗∗ Silt 0.25 −0.09 −0.29 −0.90 −0.42 0.06 0.45 0.91 −0.98 ns ns ∗∗ ∗∗ ∗∗∗ ∗∗∗ ∗∗∗ ∗∗∗ ∗∗∗ ∗∗∗ TN −0.24 −0.11 −0.27 0.43 −0.47 0.02 −0.57 0.93 −0.87 0.85 ∗∗∗ ns ∗∗∗ ∗∗∗ ∗∗∗ ns ∗∗∗ ∗∗∗ ∗∗∗ ∗∗ ∗∗∗ Clay −0.28 −0.06 −0.32 −0.91 −0.46 0.06 0.67 0.91 −0.98 −0.52 0.85 ns ns ns ns ns ns ns ns ns ns OM 0.03 0.01 −0.07 0.32 −0.11 −0.03 0.21 −0.12 −0.13 0.12 0.41 0.16 ns ns ns ns ns ns ns ns ns ns ns P 0.03 0.10 −0.03 0.09 0.01 0.03 0.08 −0.09 0.11 0.10 0.01 0.01 0.18 ∗ ∗∗ ∗∗∗ Note. Te magnitude indicates the degree of correlation, positive signs indicate positive correlations, and negative signs indicate inverse relations. p < 0.05, p < 0.01, p < 0.001, and ns � no signifcance. N � number of variables, MFBR � Majang Forest Biosphere Reserve, Hca � herbaceous cover, CaOp � canopy openness, HI � harvesting index, MC � moisture content of the soil, TN � total nitrogen, OM � organic matter, and P � available phosphorus. 12 International Journal of Forestry Research Table 13: Comparisons of the Shannon diversity index and Te communities with the highest similarity coefcients evenness of MFBR with other moist Afromontane forests in had the least beta diversity, while communities with the least southwestern Ethiopia. similarity coefcients had the highest beta diversity. Te higher Sorensen similarity between communities 1 and 2 Forest area Elevation (m) H′ J Sources might be due to the narrow topographical distance between Gesha-Sayilem forest 1734–2803 3.56 0.84 [86] the two communities where most of the plots forming these MFBR 800–2400 3.64 0.94 Present study communities may have relatively similar environmental Gerba Dima forest 1677–2240 3.45 0.93 [87] factors [52, 102–105]. Wurg forest 900–2500 3.38 0.90 [88] Agama forest 1800–2370 3.25 0.78 [89] Belete forest 1800–2300 3.79 0.95 [90] 4.3. Species Richness, Evenness, and Diversity Indices along Note. H′ � Shannon diversity index, J � evenness, and MFBR � Majang Environmental Gradients. We found high species richness Forest Biosphere Reserve. and diversity at higher elevations than at lower elevations (Table 9). Higher species richness and diversity at higher elevations may be due to variations in climatic and edaphic Celtis zenkeri is the dominant species and signifcant in- factors [106]. Te maximum species diversity and richness dicator species in this community among fve species. Tis were found on intermediate sloping terrain, followed by on community comprises an important fodder tree for honey lower sloping terrain (Table 9). Te slope has been identifed production, which contributes to the means of livelihood for as one of the topographic factors that infuence species local communities in the area [94]. diversity and richness [95, 96, 107]. Tis result agrees with P. altissima was the dominant and signifcant in- the fndings of Wondie et al. [108], who stated that forests dicator species in community 2 (Table 6) and is among grow favourably on intermediate sloping terrain. Te in- the indicator species in the upper canopy of the moist crease in species richness and diversity at intermediate Afromontane forest [92, 93], while L. fraxinifolius was sloping terrain may be due to soil moisture availability and also the dominant species and signifcant indicator edaphic factors (soil nutrients and low soil erosion) species in the middle canopy. [109, 110]. A. toxicaria is the dominant species and the distinctive species of moist Afromontane forests in the middle canopy, whereas Celtis toka was the other dominant and signifcant 4.4. Relationship between Community Types with Environ- indicator species in community 3 and is among the indicator mental and Disturbance Factors. Studies focusing on the species of moist Afromontane forests in the middle canopy relationship between plant communities with environ- [92, 93]. Te community had the highest number of plant mental and disturbance factors have become increasingly species compared to other communities. Tis could be due important in understanding the ecology of forest commu- to their location away from human disturbances since they nities [111–115]. Plant species distribution patterns and plant community formation are highly infuenced by mi- are located in difcult terrain such as sloppy and deep gorges [95, 96]. croclimate, edaphic, and anthropogenic factors [41, 93, 116–119]. Plants form a community when a plant Dracaena afromontana and Cyathea manniana were the dominant and indicator species with a signifcant indicator species repeatedly appears in a similar environment. Simi- value in this community, and they are among the distinctive larly, it is the classifcation of plant communities, which species in the middle canopy of the moist Afromontane temporally and spatially determines the boundaries of forest [92, 93]. Te D. afromontana-C. manniana com- a plant community [120–122]. Each community has its own munity type comprises diferent species as compared to distribution area with a specifc combination or preference other plant communities, which are found at higher ele- of environmental variables. vations. Te species composition diference in the com- Relatively, the elevation was the most signifcant envi- munity may be due to an environmental and anthropogenic ronmental factor infuencing and describing the community type distribution [42, 81, 123–129]. Accordingly, the vari- factor [95, 96]. Te Shannon diversity index was extended from 3.5 to ation of plant communities was also closely related to other environmental, disturbance, and site factors, including 3.8 in four plant community types, which indicates the existence of high diversity in MFBR. Te plant communities herbaceous cover, harvesting index, slope, pH, and silt showed a minor variation in their species richness, diversity, (Table 10). and evenness. Somewhat, community types such as Disturbance (illegal logging) afects the distribution of B. unijugata-P. alnifolia and P. altissima-L. fraxinifolius were plant communities by hindering seedling establishment and higher in species richness, while community types regeneration in tropical forests [130–133]. It can also assist in A. toxicaria-C. toka and D. afromontana-C. manniana had the ground layer plant species growth by facilitating the availability of light [134, 135]. the lowest species richness. Tis variation in species richness among communities could be due to the variations of to- Te correlation among environmental, disturbance, and site factors governs the pattern of species distribution in the pography, anthropogenic impact, climate, and edaphic factors [97, 98]. Each community has its own distribution plant communities of MFBR (Table 12). Tere was a positive correlation between the elevation and organic matter. Te area with a specifc combination of environmental variables [27, 99–101]. high amount of organic matter at higher elevations can be International Journal of Forestry Research 13 edaphic gradients,” Journal of Ecology, vol. 100, no. 1, attributed to a low rate of decomposition due to relatively pp. 253–263, 2012. low temperatures [113]. High organic matter content in soils [3] S. Harrison, M. J. Spasojevic, and D. Li, “Climate and plant of higher elevations was also reported in other Afromontane community diversity in space and time,” Proceedings of the forests of Ethiopia [136]. National Academy of Sciences, vol. 117, no. 9, pp. 4464–4470, 5. Conclusion [4] C. A. Baldeck, K. E. Harms, J. B. 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Publisher: Hindawi Publishing Corporation
ISSN: 1687-9368
eISSN: 1687-9376
DOI: 10.1155/2023/8521303
Publisher site: See Article on Publisher Site

Abstract

Hindawi International Journal of Forestry Research Volume 2023, Article ID 8521303, 17 pages https://doi.org/10.1155/2023/8521303 Research Article Effects of Environmental and Disturbance Factors on Plant Community Distribution in Tropical Moist Afromontane Forests, South-West Ethiopia 1 1 2 Semegnew Tadese , Teshome Soromessa , and Getaneh Gebeyehu Addis Ababa University, College of Natural and Computational Sciences, Centre for Environmental Science, Addis Ababa, Ethiopia Injibara University, Department of Biology, Injibara, Ethiopia Correspondence should be addressed to Semegnew Tadese; semetade@gmail.com Received 1 November 2022; Revised 14 February 2023; Accepted 15 March 2023; Published 6 April 2023 Academic Editor: Anna Zro´bek-Sokolnik Copyright © 2023 Semegnew Tadese et al. Tis is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Tis study was carried out to investigate the efects of environmental and disturbance factors on plant community distribution in the Majang Forest Biosphere Reserve (MFBR) in south-west Ethiopia. A systematic sample design was conducted to collect vegetation and environmental factors in four study sites. In a nested plot design, the vegetation data were collected from 140 main 2 2 2 plots, i.e., 400 m (trees), 25 m subplots (shrubs, lianas, seedlings, and saplings), and 1 m (herbs), respectively. Te plant community classifcation was performed using agglomerative hierarchical cluster analysis (Ward’s Linkage method) in R software (version 4.0.1). Te distribution of plant communities along an environmental gradient was computed using canonical cor- respondence analysis (CCA). In this study, a total of 15 (9.5%) endemic plant species were recorded in MFBR. Four plant community types were identifed, and these were Celtis zenkeri-Blighia unijugata, Pouteria altissima-Lecaniodiscus fraxinifolius, Antiaris toxicaria-Celtis toka, and Dracaena afromontana-Cyathea manniana. Environmental and disturbance factors, such as elevations, slopes, harvesting indexes, soil pH, silt, and herbaceous cover, were the most important for determining plant community distribution in the area. Elevation and slope were found to have a signifcant (P < 0.05) negative and positive relationship with species diversity and richness, respectively. Terefore, the fnding of this study provides baseline information that could be necessary for making further conservation and management in MFBR. required for successful ecological restoration and bio- 1. Introduction diversity protection [7, 8]. Plant ecologists have been successful in defning the Environmental factors have been a cornerstone in the variations in species diversity of communities along evolution of ecological theory [9, 10]. A variety of factors infuence the spatial and temporal patterns of vegetation environmental gradients at various spatial scales [1]. Plant community responses are well related to climatic change, including the physical environment, land use his- factors at regional and global scales [2, 3], whereas at local tory, prior disturbance, and initial vegetation composition or plot scales, topographic and edaphic factors play an [11–14]. Te most important data required to understand important role in controlling community structure and vegetation patterns in forest landscapes are relationships species distribution [4–8]. As a result, environmental between species diversity and environmental factors [15–21]. factors are important not only in confrming plant As a result, the interaction between physical environmental community structure and variability of species distri- variables such as elevations, slopes, and anthropogenic bution at a spatial scale but also for providing insight factors may infuence species diversity and composition into the environmental requirements of the tree species [19, 22–29]. Elevations, slopes, and aspects are determinants 2 International Journal of Forestry Research of the spatial and temporal distribution of elements such as temperatures range between 13.9 and 31.8 C at the Tinishu radiation, precipitation, and temperature that determine Meti weather station. In Ermichi, the average annual rainfall species composition [30]. Soil heterogeneity, such as soil is 2053 mm and the mean monthly minimum and maximum texture, moisture content, electric conductivity (EC), and temperatures range between 11.8 and 29.7 C. Te maximum ° ° pH, creates niches with specifc conditions, which in turn average monthly temperature is between 29.8 C and 31.8 C afect the distribution pattern of plants [31–33]. in February, while the minimum is between 11.9 C and Ethiopia has greater geographic diversity with ten eco- 13.9 C in July for Ermichi and Tinishu Meti, respectively. systems, 18 major and 49 minor agroecological zones [34]. Te maximum rainfall is between April and October, and the Tis diverse physiographic feature is responsible for the minimum rainfall is from November to March in Tinishu existence of a wide range of habitats and is conducive to the Meti and Ermichi (Figure 2). Tis temperature and rainfall survival of various plant and animal species [35]. Accord- variation between two stations is due to variations in the ingly, Ethiopia has a diverse set of plant, animal, microbial, elevation gradient (NMSA 2018). and genetic resources [36]. Ethiopia is recognized as a major Based on the vegetation classifcation of Ethiopia, the centre of diversity and endemism for a number of plant main vegetation types of the Majang Biosphere is montane species in the Horn of Africa region [37, 38]. Te fora of evergreen forests. Besides, the vegetation of this area has Ethiopia’s is diverse, with an estimated 6000–7000 species of diferent categories in terms of life forms such as high higher plants, including 647 (10.74%) endemic taxa [36]. natural forests, bush lands, and grasslands [43]. Some en- Te study of plant community types and factors (soil, vironmental factors, such as elevations, slopes, soil pH, total topographic, and disturbance) that infuence the distribution nitrogen, and phosphorus, vary between each study site in patterns of plant communities are essential inputs for forest MFBR (Table 1). conservation planning and management. For instance, previous studies have reported an association between plant community distributions and variations in environmental 2.2. Vegetation and Soil Data Collection. A systematic gradients in protected forests [25], fragmented forests [39], sampling design was established to arrange quadrats and and community-managed forests of Ethiopia [40]. However, transects as well as to collect vegetation data [44]. Te study there are limited studies on the efects of environmental and area was stratifed into four sites using a digital elevation anthropogenic disturbance factors governing plant com- model (DEM) in Arc GIS software. Te study sites’ polygon munity distribution patterns in southwestern forests [41, 42] was digitized using Google Earth by elevation classes. Tese in general and the Majang Forest Biosphere Reserve (MFBR) were site I (<1200 m.a.s.l), site II (1200–1500 m.a.s.l), site III in particular. Tis problem leads to some research questions: (1500–1800 m.a.s.l), and site IV (>1800 m.a.s.l) (Table 2). A (a) What are the main community types in the Majang total of 140 quadrats were established for vegetation and Forest Biosphere (MFBR)? (b) How do environmental and forest soil data collection, i.e., site I (1–45), site II (46–85), disturbance factors infuence plant community distribution site III (86–115), and site IV (116–140). patterns in Majang Forest Biosphere Reserves? Terefore, to Te quadrats’ X-Y coordinates were generated in Arc answer these questions, we aimed to study to (1) identify the GIS software and loaded to a global positioning system type of plant community in MFBR and (2) determine plant (GPS) receiver for tracking quadrats. Later, a measuring 2 2 community distribution patterns in response to environ- tape was used to lay out 20 × 20 m (400 m ) quadrats in mental and disturbance factors (elevation, slope, distur- each site in the biosphere. Te sampling intervals between bance, and physical and chemical composition of soil) the transect line and the quadrats were 2 km apart [43, 52]. in MFBR. Te sizes of the quadrats were determined based on the 2 2 growth forms of plants [44], i.e., 400 m (tree), 25 m (shrubs and lianas), and 1 m (herbs), respectively, in 2. Materials and Methods a nested plot design (Figure 3). 2.1. Description of the Study Area. Te study was conducted Te tree diameters (DBH ≥5 cm) were determined, and in the Majang Forest Biosphere Reserve (MFBR), situated in their number of stems was counted and recorded in the main the Majang zone, Gambella People National Regional State of plots (400 m ) (Figure 3). Te diameter at breast height and Ethiopia. It has unique biogeography and shares a boundary the height of the individual’s tree species were measured with Sale Nono woreda of Oromia regional state and using a diameter tape, clinometer, and meter tape, re- Anderacha, Yeki, Sheka, and Guraferda woreda of South- spectively. When the branching of multistemmed in- West regional state (Figure 1). It covers a total area of dividuals occurred below DBH, the DBH of each stem was 233,254 ha of forest and agricultural land and rural settle- measured separately and developed into a common diameter ments and towns. MFBR is located between the latitudes of of all stems by summing up and then taking an average ° ° 07 08′00″N and 07 50′00″N and the longitudes of diameter. To determine the diversity and estimate the ° ° 34 50′00″E and 35 25′00″E, with elevations ranging from abundance of shrubs and lianas, subplots (area � 25 m each) 562 m to 2444 m. were established (Figure 3) [44]. Te climate in the area is generally hot and humid, which Seedlings with a height of less than 1.30 m and a DBH of is marked on most rainfall maps of Ethiopia as the wettest less than 2.5 centimeters, saplings with a height of more than part of the country. Te average annual rainfall is 1774 mm, 1.30 m and a DBH of 2.5 to 5 cm, and trees as plants with and the mean monthly minimum and maximum a DBH of more than 5 cm were selected [53]. Herbaceous International Journal of Forestry Research 3 34°50′0″E35°0′0″E35°10′0″E 35°20′0″E Ethio-Region Gambella Regions- Zone 0 5 10 20 km 34°50′0″E 35°0′0″E 35°10′0″E 35°20′0″E Elevation (m) <1200 1500-1800 1200-1500 >1800 Figure 1: Location of the study sites (sites I–IV). Source: https://earthexplorer.usgs.gov. 1277 m 22.1°C 1774 mm 1582 m 20.5°C 2053 mm °C mm °C mm 50 100 50 100 40 80 40 80 29.8 31.8 30 60 30 60 20 40 20 40 11.9 13.9 10 20 10 20 0 0 0 0 J F MA M J JA S ON D JF M A M J JA SO N D Months Months (a) (b) Figure 2: Mean monthly temperature and rainfall recorded. (a) Tinishu Meti (1987–2017) and (b) Ermichi (1987–2017) (NMSA 2018). Temprature 7°10′0″N 7°20′0″N 7°30′0″N 7°40′0″N Temprature 7°10′0″N 7°20′0″N 7°30′0″N 7°40′0″N 4 International Journal of Forestry Research Table 1: Topographic and soil characteristics of the study sites. Study sites Ele (m) Slo Area (ha) SP Site I 1042 ± 42 5.3 ± 0.4 22,826.1 40 Site II 1365 ± 24 5.4 ± 0.4 25,220.5 45 Site III 1635 ± 24 7.2 ± 0.5 14,053 30 Site IV 2011 ± 42 11.1 ± 1.2 11,783.5 25 Note. Ele � elevation, Slo � slope, and SP � sample plots. Table 2: List of equations used for calculation of vegetation parameters. Vegetation parameters Equation Equation no. Reference Density D � n/N 1 [45] √�� Species richness D � n/ N 2 [46] H H � 􏽐 (Pi) (Inpi) 3 [44, 47] i�1 ′ ′ Shannon evenness J � H /Hmax � H /In s 4 [44, 47] SSI S � (2a/(2a + b + c)) 5 [44, 48] Beta (β) diversity β � (b + c/(2a + b + c)) 6 [49, 50] Harvesting index HI � 􏽐 (SD∗ EF/LD 7 [51] i�1 “a” � number of tree species common to sites A and B; “b” � number of tree species recorded only in the site A; “c” � number of tree species recorded only in site B, “N”: total number of individuals of all the species; “n”: total number of individuals of the species; Shannon diversity index (H) � where pi is the th proportion of individuals found in the i species; SSI � Sorensen similarity index, H � Shannon diversity index; SD � stump density; EF � expansion factor (3.4 for tropical forest); LD � live tree density. University. Nomenclature of plants in this article follows 20 m those published in the fora of Ethiopia as well as fora of Ethiopia and Eritrea [57–62]. 5 m Soil samples were taken with a soil auger from the topsoil at a depth of 0–30 cm, which is recommended as the default 5 m sampling depth for soil [63]. Te soil samples were taken from 1 m fve diferent locations in each plot, four from the quadrat’s 20 m 1 m corners and one from the quadrat’s centre. A total of 220 soil samples, 140 from forestland, 40 from farmland, and 40 from grassland, were collected, composited separately, labelled, and transported to the laboratory. To determine soil bulk density, soil was collected on the centre of the quadrats using a stainless core sampler, then placed in plastic bags, and transported to the laboratory for dry weight determination. Fresh wet soil weights Figure 3: Sampling design used for data collection [44]. Note: were measured in the feld with a kitchen balance with 0.1 g subplots (1 × 1 m) are measured for herbs, grasses, and soil samples; precision. A composite sample of 200 g was taken from each subplots (5 × 5 m) are measured for shrubs and lianas, and the main quadrat to analyze its chemical composition [64] in the Water plot (20 × 20 m) is measured for trees, saplings, and seedlings. Works Design and Supervision Enterprise (WWDSE) labora- tory in Addis Ababa. cover abundance was assessed in fve subquadrats (1 m ) within the main plot (400 m ). 2.3. Environmental and Disturbance Data Collection. Te herbaceous cover abundance from each quadrat was Environmental factors such as elevations, slopes, and aspects changed into 9 cover abundance [54] scale classes: were measured and recorded for each of 140 quadrats using 1 � (<0.5%), 2 � (0.5–1.5%), 3 � (1.5–3%), 4 � (3–5%), clinometers and Garmin GPS, respectively. Te elevation 5 � (5–12.5%), 6 � (12.5–25%), 7 � (25–50%), 8 � 50–75%, was coded into four elevation ranges, namely, and 9 ≥ 75% [55]. Canopy openness was measured by using 1 ≤1200 m.a.s.l, 2 �1200–1500 m.a.s.l, 3 �1500–1800 m.a.s.l, a densiometer located at the centre of each plot. We classifed and 4 ≥1800 m.a.s.l [65]. Aspect values were collected and canopy openness by the following classifcation scheme: very arranged (N � 0, NE � 1, E � 2, SE � 3, S � 4, SW � 3.25, dense canopy >70%, moderately dense canopy 40–70%, W � 2.5, and NW � 1.25) in each quadrat [52, 66]. Te slope open canopy 10–40%, scrub <10%, and nonforest 0% [56]. range was classifed into three major slope classes [67]. As Plant species were identifed in the feld, and for those a result, the classes were as follows: (1) fat <10, (2) in- species that were difcult to identify in the feld, voucher termediate 10–20, and (3) steep >20. specimens were collected with the help of local guides, Human disturbance (which includes harvesting/cutting coded, pressed, and dried for subsequent identifcation and trees for fuel wood, charcoal, timber, and house construction verifcation at the National Herbarium (ETH), Addis Ababa illegally) was computed as the harvesting index. Te International Journal of Forestry Research 5 Shannon evenness index (J) were calculated using equations harvesting index was computed by counting individual stumps that were illegally logged trees within the quadrats. It 3 and 4 in Table 2. Te Sorensen similarity index was used to assess the degree of foristic similarity between plant com- is calculated from the density of individual tree stumps by following equation (7) in Table 2 [51]. Stumps are a small munities. Te Sorensen similarity index was calculated using portion of the trunk that remains after a tree with about 5 cm equation 5 in Table 2. Beta diversity was computed by the chopped down [68]. change in diversity of species from one community to an- other and calculated using equation 6 in Table 2 [49]. Pearson correlation coefcients were calculated to see the 2.4. Data Analysis relationship between diversity indices and environmental disturbance factors in MFBR. 2.4.1. Cluster Analysis. Te vegetation data, which were originally estimated in the feld as a percentage of cover abundance values, were converted into Domin cover 2.4.3. Soil Laboratory Analysis. Te soil samples were an- scales prior to analysis [54]. Te vegetation data were alyzed in the Water Works Design and Supervision En- examined using an agglomerative hierarchical cluster terprise (WWDSE) laboratory in Addis Ababa, Ethiopia. Te (AHC) analysis in package cluster to classify vegetation Bouyoucos hydrometer method was used to determine soil into plant community types. Te similarity ratio (SR) textures. Soil pH was determined using a pH meter a 1 : 2.5 with Ward’s group linkage methods (minimum variance soil to water suspension potentiometric method [72]. Te clustering) was used for cluster analysis. Te similarity micro-Kjeldahl [73] and Walkley and Black [74] methods ratio is one of the similarity indices that determine how were used to determine total nitrogen (N) and soil organic similar or dissimilar vegetation samples, quadrats, or carbon, respectively. Te Bray-I method was used to de- plots are. Ward’s group linkage method evaluates cluster termine available phosphorus, and the absorbance of the distances using the analysis of variance methodology Bray-I extract was measured in a spectrophotometer at [52]. Te goal of this strategy is to eliminate diferences in 882 nm [75]. Based on C and N concentrations, the carbon- total abundance between sample units (quadrats). In to-nitrogen ratio (C/N) was calculated. Te dry weight of addition, Ward’s method minimizes the total within- each soil sample (MS) was determined using oven-drying set group mean of squares or residual sum of squares [69]. to 105 C for 24 h to achieve a constant weight [76]. Te Te maximum number of clusters in a data set is volume of the core sampler (VC) was determined as VC � π a central issue in partitioning clustering, such as k-means r h, where r is the radius and h is the height of the core clustering, which requires the user to specify the number sampler (VC � 3.14 × (2.5 cm) 2 × 5 cm � 98.125 cm ) to of clusters k to be created. In this study, the optimum calculate the bulk density for each sample. number of clusters was identifed using the gap statistic method and determined by plotting the gap statistic against the number of clusters (k-means). Te point, 2.4.4. Ordination. Te canonical correspondence analysis where the maximum gap statistic in the plot is, de- (CCA) ordination method was used to fnd out plant termines the proper number of clusters. In this study, the community distribution along the environmental gradient maximum gap statistic value in the plot is shown at k- [77]. Te selection of CCA was based on the results of means � 4 (the maximum number of clusters or detrended correspondence analysis (DCA), which showed communities) [70]. that the longest axis of DCA for a data set was greater than 3 Characteristic species were determined using syn- (�4.88) (Table 3). Tis indicates the presence of higher optic table analysis. Plant community types were named β-diversity or heterogeneous vegetation data due to the after two distinctive species with high synoptic cover- unimodal relationship between species and environmental abundance ratings [44]. Indicator species analysis was variables. Permutational multivariate analysis of variance performed using the package labs in R software. Te (function adonis test) was used to choose environmental signifcance of the indicator value was tested using the factors that were relatively more relevant in explaining the Monte Carlo test (P < 0.05) [71]. Te indicator values species data and signifcance before CCA ordination [78]. were calculated from the product of relative abundance Pearson’s correlation coefcient was calculated to evaluate (specifcity) and relative frequency of species (fdelity) the relationship between environmental factors and plant within a community [71]. In this study, the species with diversity. the highest signifcant indicator value is considered an indicator species of a community. 3. Result 2.4.2. Community Diversity Analysis. Species richness, 3.1. Plant Diversity. In this study, a total of 15 (9.5%) en- evenness, Shannon diversity, and beta diversity indices were demic plant species were recorded from Ethiopia. Of the computed using R software (version 4.0.1). Species richness total endemic species, 9 are herbaceous, 3 are shrubs, 2 are is usually expressed as the number of species per sample unit trees, and 1 is a liana in life form (Table 4). Rinorea friisii and and calculated using equation 2 in Table 2 [49]. Shannon’s Crotalaria rosenii were listed as near threatened among the index was computed using abundance and evenness of the 15 endemic species, while eight others were listed as least species present. Te Shannon diversity index (H′) and the concerned (Table 4). 6 International Journal of Forestry Research Table 3: Detrended correspondence analysis result of MFBR vegetation. DCA axes DCA1 DCA2 DCA3 DCA4 Eigen values 0.8356 0.2267 0.1917 0.1600 Decorana values 0.8599 0.2441 0.1845 0.1449 Axis lengths 4.8857 2.2693 2.4418 2.2889 3.2. Plant Community Types. Four plant community types Table 4: Endemic plants, their habits, and IUCN status in the were identifed using agglomerative hierarchical cluster Majang Forest Biosphere Reserve. analysis of the entire vegetation data set of MFBR (Figure 4 IUCN and Table 5). Te identifed communities were Celtis Plant species Family name LF category zenkeri-Blighia unijugata community (C1), Pouteria Acanthopale aethiogermanica altissima-Lecaniodiscus fraxinifolius community (C2), Acanthaceae NA S Ensermu Antiaris toxicaria-Celtis toka community (C3), and Dra- Bothriocline schimperi Oliv. & caena afromontana-Cyathea manniana community (C4). Asteraceae LC H Hiern ex Benth Clematis longicauda Steud. ex C1: Celtis zenkeri-Blighia unijugata community Ranunculaceae LC L A. Rich Tis community was found in the elevation range of Crotalaria rosenii (Pax) Fabaceae NT H 830–1896 m.a.s.l and was represented by 77 species. Of Milne-Redh. ex Polhill these, 67 species are commonly shared with other Dorsetnia soerensenii Friis Moraceae NA H communities, while 12 are found only in this com- Euphorbia dumalis S. Carter Euphorbiaceae LC H munity type. Among the total species, four species were Impatiens rothii Hook.f Balsaminaceae NA H signifcant indicator species, with higher indicator Millettia ferruginea (Hochst.) Bak Fabaceae LC T Pycnostachys abyssinica Fresen Lamiaceae NA H values than those of total species (Table 6), namely, Rinorea friisii M.G.Gilbert Violaceae NT S Blighia unijugata, Pouteria alnifolia, Margaritaria Solanecio gigas (Vatke) C. Jefrey Asteraceae LC S discoidea, and Garcinia buchananii. Te most domi- Solanum marginatum L.f Solanaceae NA H nant shrub species included Argomuellera macrophylla, Vepris dainellii (Pic. Serm.) Whitfeldia elongata, Dracaena fragrans, Vernonia Rutaceae LC T Kokwaro amygdalina, and Phyllanthus reticulatus. Leptaspis Vernonia fligera Oliv. & Hiern Asteraceae LC H zeylanica, Aspilia mossambicensis, Asplenium Vernonia leopoldi (Sch. Bip. ex Asteraceae LC H bugoiense, Asplenium anisophyllum, and Pollia con- Walp.) Vake densata were the dominant herb layers in the com- Note. LF � life form, T �tree, S � shrub, H � herb, and L � liana; IUCN threat munity. Te common lianas of this community were categories (LC � least concern, NA � not available, and NT �near Embelia schimperi, Combretum paniculatum, Dregea threatened). schimperi, and Goupia glabra. C2: Pouteria altissima-Lecaniodiscus fraxinifolius community herb layer was dominated by Asplenium bugoiense and Aspilia mossambicensis. Tis community was distributed in the elevation range of 800 to 2310 m.a.s.l. and was represented by 77 C3: Antiaris toxicaria-Celtis toka community species. Of these, 71 species are commonly shared with Tis community was found in the elevation range of other communities, while 6 are found only in this 850–2300 m.a.s.l and was represented by 105 species. community type. Among the total species, four species Of these, 92 species are commonly shared with other were signifcant indicators with higher indicator values communities, while 13 are found only in this com- (Table 6), namely, Pouteria altissima, Lecaniodiscus munity type. Among the total species, four species are fraxinifolius, Ficus exasperata, and Ritchiea albersii. signifcant indicator species with higher indicator Moreover, the dominant tree species layers were values (Table 6), namely, Celtis zenkeri, Morus meso- Lannea welwitschii, Ritchiea albersii, Grewia molli, zygia, Vernonia hochstetteri, and Celtis toka. Moreover, Ficus sur, Ficus mucuso, Ficus ovata, Ficus exasperata, the dominant species in the tree layer were Diospyros Ficus umbellata, Allophylus macrobotrys, and Croton abyssinica, Morus mesozygia, Combretum molle, Apo- sylvaticus. Erythrococca trichogyne, Whitfeldia elon- dytes dimidiata, Olea capensis, Dombeya torrida, gata, Dracaena fragrans, Rinorea friisii, Vernonia Buddleja polystachya, and Lepisanthes senegalensis. Te amygdalina, Acalypha orbata, Pittosporum viridi- dominant species in the shrub layer were Vernonia forum, Phyllanthus reticulatus, and Oxyanthus spe- hochstetteri, Acanthopale pubescens, Erythrococca ciosus were prominent shrubs in this community. trichogyne, Plectranthus sp, and Capparis tomentosa. Jasminum dichotomum, Combretum paniculatum, Marantochloa leucantha, Solanum nigrum, Achyr- Phytolacca dodecandra, Dregea schimperi, Symphytum anthes aspera, and Asplenium aethopicum were the ofcinale, Pseuderanthemum tunicatum, Uncaria afri- dominant species in the herb layer. Te dominant li- cana, and Saba comorensis were common lianas. Te anas in this community were Peponium vogelii, International Journal of Forestry Research 7 Table 5: Synoptic cover value of plants in MFBR for species ≥1% in at least one community. Scientifc name C1 C2 C3 C4 Cluster size 36 25 44 35 Celtis zenkeri 3.450 2.120 2.659 0.000 Blighia unijugata 2.806 1.040 1.841 0.000 Apodytes dimidiata 2.444 2.000 1.364 0.000 Pouteria alnifolia 2.222 1.240 1.000 0.000 Trichilia prieuriana 1.694 1.200 0.864 0.000 Margaritaria discoidea 1.694 0.640 0.818 0.000 Argomuellera macrophylla 1.611 1.320 1.000 0.000 Diospyros abyssinica 1.556 0.800 1.955 0.000 Lannea welwitschii 1.500 1.600 1.409 0.000 Combretum molle 1.389 0.600 0.432 0.000 Pouteria altissima 2.472 3.400 2.864 0.000 Lecaniodiscus fraxinifolius 1.472 3.220 1.955 0.000 Erythrococca trichogyne 1.361 2.040 0.750 0.943 Apodytes dimidiata 2.444 2.000 1.364 0.000 Ritchiea albersii 0.944 1.840 1.227 0.000 Ficus exasperata 1.000 1.760 0.364 0.000 Antiaris toxicaria 1.778 1.640 3.532 0.000 Celtis toka 1.417 1.200 2.909 0.000 Morus mesozygia 0.556 0.480 1.477 0.000 Croton macrostachyus 0.472 0.800 1.045 1.171 Deinbollia kilimandschrica 0.250 1.040 1.045 0.000 Grewia mollis 1.222 1.400 1.159 0.000 Ficus sur 0.222 1.520 1.136 0.943 Dracaena afromontana 0.000 0.000 0.000 3.514 Cyathea manniana 0.000 0.000 0.000 3.414 Schefera abyssinica 0.000 0.000 0.000 2.800 Trilepisium madagascariense 0.000 0.000 0.000 2.686 Galiniera saxifraga 0.000 0.000 0.000 2.600 Schefera myriantha 0.000 0.000 0.000 1.914 Vernonia bipotini 0.000 0.000 0.000 1.886 Allophylus abyssinicus 0.000 0.000 0.000 1.857 Pouteria adolf-friedericii 0.000 0.000 0.000 1.829 Clausena anisata 0.000 0.000 0.000 1.829 Note. C1 � community one, C2 � community two, C3 � community three, and C4 � community four. 5.0 4.5 4.0 3.5 3.0 2.5 2.0 1.5 1.0 0.5 0.0 Plots Figure 4: Dendrogram of the cluster analysis results of species abundance found in 140 of MFBR. C1: 1, 17, 5, 21, 8, 24, 14, 30, 31, 38, 45, 61, 86, 90, 78, 98, 7, 23, 10, 26, 15, 50, 66, 43, 59, 33, 47, 63, 40, 56, 74, 94, 83, 103, 77, and 97; C2: 2, 18, 9, 25, 13, 29, 3, 19, 11, 27, 72, 92, 85, 105, 76, 96, 4, 20, 89, 12, 28, 75, 95, 82, and 102; C3: 6, 22, 35, 16, 54, 70, 36, 53, 69, 42, 58, 49, 65, 71, 91, 79, 99, 87, 80, 100, 84, 104, 88, 32, 39, 55, 46, 62, 48, 64, 52, 68, 73, 93, 81, 101, 34, 37, 44, 60, 41, 57, 51, and 67; C4: 106, 126, 110, 130, 119, 139, 112, 132, 116, 136, 123, 109, 129, 120, 140, 117, 137, 114, 134, 125, 107, 127, 118, 138, 111, 131, 121, 115, 135, 124, 108, 128, 113, 133, and 122. Dissimilarity 8 International Journal of Forestry Research Table 6: Higher indicator values with signifcance of species in the 3.2.1. Species Richness, Evenness, and Diversity among Plant clusters. Community Types. Te overall Shannon diversity index and evenness in MFBR were 3.64 and 0.94, respectively (Table 7). Te Species Community R. Ab R. Fr Indval highest Shannon diversity was shown in community 1 (3.8), value while the least was exhibited in community 4 (3.5). Te max- Blighia unijugata C1 0.493 0.778 0.384 0.001 imum Shannon evenness was indicated in community type four Pouteria alnifolia C1 0.498 0.667 0.332 0.001 (0.9), followed by community types three, two, and one (each Margaritaria C1 0.537 0.583 0.314 0.001 with 0.7 evenness) (Table 8). Te highest species richness was discoidea Garcinia buchananii C1 1.000 0.306 0.306 0.001 exhibited in community type one (102 species), whereas the Pouteria altissima C2 0.283 0.722 0.389 0.001 lowest plant species richness was revealed in community type Lecaniodiscus four (70) (Table 7). C2 0.225 0.278 0.343 0.001 fraxinifolius Similarity coefcients of all communities range from 0.61 Ficus exasperata C2 0.320 0.361 0.293 0.001 to 0.91. Te highest Sorensen similarity was exhibited be- Ritchiea albersii C2 0.235 0.306 0.275 0.001 tween communities 1 and 2 (0.91) followed by communities Celtis zenkeri C3 0.841 0.365 0.307 0.005 2 and 3 (0.90). Te lowest Sorensen similarity was revealed Morus mesozygia C3 0.523 0.588 0.307 0.001 between communities 1 and 4 (0.61) and communities 2 and Vernonia hochstetteri C3 0.341 0.799 0.272 0.001 4 (0.63) of the total species in MFBR (Table 8). Celtis toka C3 0.636 0.422 0.268 0.005 Sorenson similarity coefcients and beta diversity were Dracaena C4 1.000 1.000 1.000 0.001 afromontana inversely related to each other. Terefore, communities with Cyathea manniana C4 1.000 1.000 1.000 0.001 the lowest similarity coefcients had the highest beta di- Vernonia bipotini C4 1.000 0.914 0.914 0.001 versity (0.45) in communities 1 and 4, whereas communities Schefera abyssinica C4 1.000 0.886 0.886 0.001 with the highest similarity coefcients had the lowest beta Note. R.Ab � relative abundance, R.Fr � relative frequency, diversity (0.102) (Table 8). Indval � indicator values in the class, and indicator species in the respective community types at P < 0.05. 3.2.2. Species Richness, Evenness, and Diversity along Envi- ronmental and Disturbance Factor. Species diversity varied Ampelocissus bombycina, Sericostachys scandens, from 2.33 to 3.64; the higher value was recorded at the higher Tacazzea apiculata, Paullinia pinnata, Lagenaria elevation gradient (>1800 m.a.s.l), and the lower value was siceraria, and Cayratia gracilis. recorded at the lower elevation gradient (<1200 m.a.s.l). Species richness and Shannon diversity exhibited a signifcant positive C4: Dracaena afromontana-Cyathea manniana correlation with elevations (r� 0.302, p � 0.0002) and community (r� 0.493, p � 0.00005), respectively, while evenness showed Te community is distributed in the elevation range of a signifcant negative correlation with the elevation gradient 1820–2359 m.a.s.l and was represented by 70 species in (r� −0.385, p � 0.00003) (Table 9). Similarly, the slope showed 35 plots. Of these, 35 species are commonly shared with a signifcant positive correlation with species richness (r � 0.262, other communities, while 35 are found only in this p � 0.00002) and species diversity (r� 0.373, p � 0.00001), community type. Among the total species, four species while the slope showed a signifcant negative correlation with are signifcant indicator species with higher indicator species evenness (r� −0.181, p � 0.0005) (Table 9). values (Table 6), namely, Dracaena afromontana, Te highest species diversity was shown on the in- Cyathea manniana, Vernonia bipotini, Schefera termediate slope (10–20) gradient, while the lowest diversity abyssinica, and Pouteria adolf-friedericii. Te domi- recorded the lower slope (<10). Te herbaceous cover nant tree species in the community were Allophylus showed a signifcant positive correlation with species rich- abyssinicus, Pouteria adolf-friedericii, Triumfetta ness and Shannon diversity (r � 0.137, p � 0.01) and tomentosa, Pycnostachys eminii, Dracaena refexa, (r � 0.171, p � 0.004), respectively. Species richness and Ekebergia capensis, Ilex mitis, Elaeodendron buchana- Shannon diversity showed a signifcant negative correlation nii, Albizia schimperiana, Dracaena afromontana, with the harvesting index (r � −0.243, p � 0.003) and Cyathea manniana, Schefera abyssinica, Trilepisium (r � −0.376, p � 0.00004), respectively. Similarly, soil pH and madagascariense, Galiniera saxifrage, and Schefera silt showed a signifcant negative relationship with species myriantha. Justicia schimperiana, Acanthopale aethio- richness (r � −0.321, p � 0.0001) and (r � −0.504, germanica, Clausena anisata, Vernonia auriculifera, p � 0.0004) and Shannon diversity (r � −0.302, p � 0.0002) Maesa lanceolata, Maytenus gracilipes, Clematis long- and (r � -0.458, p � 0.00001), respectively. However, soil icauda, and Vernonia bipotini were the dominant pH and silt showed a signifcant positive relationship with species in the shrub layer. Te dominant species in herb species evenness (r � 0.365, p � 0.0009) and (r � 0.353, layer were Alchemilla fscheri, Bothriocline schimperi, p � 0.00001), respectively (Table 9). Justicia sp, Asplenium friesiorum, Setaria megaphylla, Asplenium sandersonii, and Physalis angulata. Te dominant lianas in this community were Clematis 3.3. Relationship between Community Types with Environ- mental and Disturbance Factors. Te length of the frst axis simensis, Culcasia falcifolia, Hippocratea pallens, and Jasminum abyssinicum. of DCA indicates species heterogeneity in vegetation as International Journal of Forestry Research 9 Table 7: Species richness, evenness, and diversity indices of plant (r � 0.0445, p � 0.001), elevation (r � 0.0906, p � 0.001), community types. slope (r � 0.0143, p � 0.014), pH (r � 0.0127, p � 0.032), and silt (r � 0.0114, p � 0.0047) exhibited a signifcant Community type Vegetation parameters diference among environmental and disturbance factors C1 C2 C3 C4 (Table 10). Tus, signifcant environmental and distur- Species richness 102 77 93 70 bance factors infuence the species distribution pattern in H′ 3.8 3.6 3.7 3.5 all community types. Shannon evenness 0.7 0.7 0.7 0.9 2 −1 CCA ordination was performed using data on cover Basal area (m )/ha 58.55 48.82 64.64 76.3 −1 abundance values of plant species in the plots and six sta- Density (ha ) 1532 1018 1478 1309 −1 tistically signifcant environmental disturbance factors. In Seedling density (ha ) 3349 3087 4431 3804 −1 the canonical correspondence analysis (CCA), the eigen- Sapling density (ha ) 1629 1457 1789 1509 values obtained in the frst and second axes were 54.4 and Note. H′ � Shannon diversity index, C1 � community 1, C2 � community 2, C3 � community 3, and C4 � community 4. 4.4, respectively. Te correlation results of environmental disturbance factors and the CCA axis showed both a positive and negative correlation from the frst to the last axis Table 8: Pairwise comparison of Sorensen’s similarity coefcient (Table 11). Te elevation and slope showed a positive cor- and beta diversity among four plant communities. relation (r � 0.868, p < 0.05) and (r � 0.638, p < 0.05), re- spectively, in the frst axis. Among signifcant environmental Communities C1 C2 C3 C4 disturbance factors, the elevation was an extremely im- C1 1 portant constraining factor followed by the slope and har- C2 0.91 (0.102) 1 C3 0.87 (0.13) 0.90 (0.104) 1 vesting index (HI) on plant species distribution patterns in C4 0.61 (0.45) 0.63 (0.43) 0.64 (0.38) 1 MFBR. On the other hand, pH and silt showed a strong Note. Values outside the bracket indicate Sorensen’s coefcient, while those negative correlation (r � −0.907, p < 0.05) and (r � 0.792, inside the bracket indicate the beta diversity index. C1 � community 1, p < 0.05), respectively, in the frst axis. CCA showed that the C2 � community 2, C3 � community 3, and C4 � community 4. cumulative proportion and proportion explained in the frst axis accounted 76.3%, respectively, which is explained as species and environmental disturbance factor variations (Table 11). Table 9: Pearson correlation coefcients between diversity indices Ordination of the study plots of MFBR formed four and environmental and disturbance factors in MFBR. communities based on the species composition. Tese Environmental disturbance Diversity indices four community types were separated by following the site SR H′ J arrows of the environmental variables. Red, green, blue, factors and purple represent sample plots of community types 1, ns ∗ ∗ Hca 0.137 0.171 −0.030 ns 2, 3, and 4, respectively (Figure 5). Community four ∗∗∗ ∗∗∗ HI −0.243 −0.376 −0.040 ∗∗∗ ∗∗∗ ∗∗∗ generally occurs at a higher elevation, while species in Ele 0.302 0.493 −0.385 ∗∗∗ ∗∗∗ ∗ communities one, two, and three are distributed at the Slope 0.262 0.373 −0.181 ∗∗∗ ∗∗∗ ∗∗∗ middle and lower elevations. Te distribution of species in Soil pH −0.321 −0.504 0.365 ∗∗∗ ∗∗∗ ∗∗∗ communities four was infuenced by elevation, slope, Silt −0.302 −0.458 0.353 harvesting index, and herbaceous cover factors, whereas Note. Hca � herbaceous cover, HI � harvesting index, Ele � elevation, SR � species richness, H′ � species diversity index, and J � species evenness. the distribution of species in communities one, two, and Positive signs indicate a positive correlation, and negative signs indicate three was strongly infuenced by soil pH and silt factors ∗ ∗∗ ∗∗∗ inverse relations. p < 0.05, p < 0.01, p < 0.001, and ns � no (Figure 5). Te length and direction of the vectors in the signifcance. plots and environment and disturbance biplot represent the degree of correlation between the plots and envi- a result of environmental variables (Table 3). Te frst axis of ronmental disturbance factors. Communities’ types one, DCA was> 3 (4.88), which indicates a wider length of the two, and three are more closely related to one another, gradient and the presence of higher β- diversity or het- whereas community type four is diferent from the other erogeneous vegetation data due to the unimodal relationship three community types (Figure 5). between species and environmental factors (Table 3). Te correlation results between environmental and Terefore, CCA was selected to show the efect of envi- disturbance factors exhibited both positive and negative ronmental and disturbance factors on the patterns of plant relationships in MFBR (Table 12). Te elevation showed community distribution. a signifcant positive correlation with slopes (r � 0.44), MC Furthermore, signifcant environmental and distur- (r � 0.58), sand (r � 0.33), and TN (r � 0.43), while it showed bance factors (P < 0.05) were selected using permuta- a signifcant negative relationship with pH (r � −0.76), silt tional multivariate analysis of variance (function adonis (r � −0.90), and clay (r � −0.91). Similarly, the slope showed test). Out of 14 environmental and disturbance factors, a signifcant negative relationship with MC (r � −0.38), six factors were signifcant in explaining patterns of plant pH (r � -0.46), silt (r � −0.42), TN (r � −0.47), and clay community distribution. More specifcally, herbaceous (r � −0.46), while it showed a signifcant positive correlation cover (r � 0.0409, p � 0.001), harvesting index with sand (r � 0.45). Soil pH showed a signifcant positive 10 International Journal of Forestry Research Table 10: Result of the function Adonis test of signifcant envi- ronmental and disturbance factors in MFBR. Sums of Mean Variables Df F model R Pr(>F) squares squares ∗∗∗ Hca 1 1.8900 1.8898 6.9218 0.0409 0.001 CaOp 1 0.1770 0.1766 0.6470 0.0038 0.767 ∗∗∗ HI` 1 2.0610 2.0608 7.5481 0.0445 0.001 ∗∗∗ Ele 1 4.1910 4.1912 15.3514 0.0906 0.001 Slope 1 0.6600 0.6600 2.4173 0.0143 0.014 Aspect 1 0.4640 0.4640 1.6994 0.0100 0.08 -3 MC 1 0.2000 0.1998 0.7316 0.0043 0.703 pH 1 0.5860 0.5863 2.1473 0.0127 0.032 Sand 1 0.2280 0.2284 0.8367 0.0049 0.559 -6 Silt 1 0.5290 0.5293 1.9388 0.0114 0.047 Clay 1 0.3360 0.3365 1.5142 0.0631 0.441 -1 012 TN 1 0.4140 0.4135 1.5146 0.0089 0.137 CCA1 OM 1 0.2900 0.2898 1.0614 0.0063 0.369 Communities P 1 0.1750 0.1748 0.6403 0.0038 0.814 a Community 1 a Community 3 Residuals 126 34.4000 0.2730 0.7436 Total 139 46.2640 1.0000 a Community 2 a Community 4 Note. Hca � herbaceous cover, CaOp � canopy openness, HI � harvesting Figure 5: Canonical correspondence analysis (CCA) ordination index, Ele � elevation, MC � moisture content of the soil, TN � total ni- diagram of 140 plots and 6 signifcant environmental variables and trogen, OM � organic matter, and P � available phosphorus. plot number distribution in four community types. forests of Ethiopia may be due to ecological and geographical Table 11: Scores of constraining variables and correlations between environmental and disturbance factors with CCA axes in MFBR. separations [84]. Based on the plant species endemic to Ethiopia, Rinorea friisii and Crotalaria rosenii were listed as Environmental variables CCA1 CCA2 CCA3 CCA4 near threatened, while other eight were registered as least Hca 0.429 −0.128 0.160 −0.541 concerned (Table 4) [85]. Tis suggests that special attention HI 0.522 0.028 0.207 0.335 needs to be given to the forest stand containing species Ele 0.868 0.066 0.100 −0.014 which are threatened and least concerned. Slope 0.638 −0.238 0.135 −0.280 Te overall diversity investigation of MFBR showed the pH −0.907 0.047 −0.029 −0.020 higher Shannon diversity index (3.64) and evenness (0.94). Silt −0.792 0.060 −0.187 −0.243 Te value of species diversity and evenness in MFBR is Eigenvalue 0.544 0.044 0.040 0.033 Proportion explained 0.763 0.062 0.057 0.046 greater than that of Gesha-Sayilem forest, Kibate forest, Cumulative proportion 0.763 0.825 0.882 0.928 Gerba Dima forest, Wurg forest, and Agama forest, while it Note. Hca � herbaceous cover, HI � harvesting index, Ele � elevation, and is lower than that of Belete forest (Table 13). Tis diference MFBR � Majang Forest Biosphere Reserve. may be due to links with geographical locations, climatic conditions, and elevation factors. relationship with silt (r � 0.91), TN (r � 0.93), and clay 4.2. Plant Community Types. In this study, a total of four (r � 0.91), while it showed a signifcant negative relationship plant communities were identifed, which are rich in species with sand (r � −0.93) (Table 12). composition. Te major distinguishing features of the identifed plant communities were the diference in domi- 4. Discussion nant and indicator plant species. However, communities 1–3 4.1. Endemic Plant Species and Diversity. Te Majang Forest share more species (>70 species) than community 4 (35 Biosphere Reserve had shown high endemism as proposed species), which could be directly related to environmental by the World Conservation and Monitoring Centre (com- and disturbance factors that make the plant communities have their own distinct or characteristic species [42]. Almost prises high or 9.5% endemic species) (Table 4) [79]. Tus, endemism is higher than reported in other moist Afro- all communities identifed more than four indicator species with a signifcant value. Indicator plant species are plants montane forests of southwestern Ethiopia. For instance, Bonga forest recorded 13 endemic species (5.35%) [80] and that are easily monitored and predict the condition of the Yayu forest with only 3 endemic species (1.36%) [81]. On the environment where they originated. Indicator species can other hand, the number of endemic species recorded in this also be a sign of a distinctive set of environmental qualities or study is lower than in some dry Afromontane forests of characteristics found in a specifc place [91]. Ethiopia. For example, Wof Washa forest has 29 species Blighia unijugata and Pouteria alnifolia were indicator (12%) [82]; Chilimo forest has 18 endemic species (8.45%) species in community 1. Blighia unijugata is the dominant [83]. Te lower number of endemics in the moist Afro- and indicator species and is among the indicator species of montane forests of the southwestern and dry Afromontane moist Afromontane forests in the middle canopy [92, 93]. CCA2 International Journal of Forestry Research 11 Table 12: Pearson’s correlation coefcient matrix for environmental and disturbance factors (N � 14) in MFBR. Clay Hca CaOp HI Elevation Slope Aspect MC pH Sand Silt TN OM P (ns) Hca CaOp 0.27 ns ns HI 0.13 0.05 ns ∗∗ ∗∗∗ Elevation 0.23 0.10 0.33 ns ns ∗ ∗∗∗ Slope 0.15 −0.07 −0.28 0.44 ns ns ns ns ns Aspect −0.08 0.08 −0.03 −0.09 −0.11 ns ns ∗ ∗ ∗∗∗ ∗∗∗ MC 0.18 −0.05 −0.14 0.58 −0.38 −0.09 ∗∗ ns ∗∗∗ ∗∗∗ ∗∗∗ ns ∗∗∗ pH −0.28 −0.08 −0.32 −0.76 −0.46 0.09 −0.57 ns ns ∗∗ ∗∗∗ ∗∗∗ ∗∗∗ ∗∗∗ ∗∗∗ Sand −0.27 0.08 0.31 0.33 0.45 −0.06 −0.47 −0.93 ∗∗ ns ∗∗ ∗∗∗ ∗∗∗ ns ∗∗∗ ∗∗∗ ∗∗∗ Silt 0.25 −0.09 −0.29 −0.90 −0.42 0.06 0.45 0.91 −0.98 ns ns ∗∗ ∗∗ ∗∗∗ ∗∗∗ ∗∗∗ ∗∗∗ ∗∗∗ ∗∗∗ TN −0.24 −0.11 −0.27 0.43 −0.47 0.02 −0.57 0.93 −0.87 0.85 ∗∗∗ ns ∗∗∗ ∗∗∗ ∗∗∗ ns ∗∗∗ ∗∗∗ ∗∗∗ ∗∗ ∗∗∗ Clay −0.28 −0.06 −0.32 −0.91 −0.46 0.06 0.67 0.91 −0.98 −0.52 0.85 ns ns ns ns ns ns ns ns ns ns OM 0.03 0.01 −0.07 0.32 −0.11 −0.03 0.21 −0.12 −0.13 0.12 0.41 0.16 ns ns ns ns ns ns ns ns ns ns ns P 0.03 0.10 −0.03 0.09 0.01 0.03 0.08 −0.09 0.11 0.10 0.01 0.01 0.18 ∗ ∗∗ ∗∗∗ Note. Te magnitude indicates the degree of correlation, positive signs indicate positive correlations, and negative signs indicate inverse relations. p < 0.05, p < 0.01, p < 0.001, and ns � no signifcance. N � number of variables, MFBR � Majang Forest Biosphere Reserve, Hca � herbaceous cover, CaOp � canopy openness, HI � harvesting index, MC � moisture content of the soil, TN � total nitrogen, OM � organic matter, and P � available phosphorus. 12 International Journal of Forestry Research Table 13: Comparisons of the Shannon diversity index and Te communities with the highest similarity coefcients evenness of MFBR with other moist Afromontane forests in had the least beta diversity, while communities with the least southwestern Ethiopia. similarity coefcients had the highest beta diversity. Te higher Sorensen similarity between communities 1 and 2 Forest area Elevation (m) H′ J Sources might be due to the narrow topographical distance between Gesha-Sayilem forest 1734–2803 3.56 0.84 [86] the two communities where most of the plots forming these MFBR 800–2400 3.64 0.94 Present study communities may have relatively similar environmental Gerba Dima forest 1677–2240 3.45 0.93 [87] factors [52, 102–105]. Wurg forest 900–2500 3.38 0.90 [88] Agama forest 1800–2370 3.25 0.78 [89] Belete forest 1800–2300 3.79 0.95 [90] 4.3. Species Richness, Evenness, and Diversity Indices along Note. H′ � Shannon diversity index, J � evenness, and MFBR � Majang Environmental Gradients. We found high species richness Forest Biosphere Reserve. and diversity at higher elevations than at lower elevations (Table 9). Higher species richness and diversity at higher elevations may be due to variations in climatic and edaphic Celtis zenkeri is the dominant species and signifcant in- factors [106]. Te maximum species diversity and richness dicator species in this community among fve species. Tis were found on intermediate sloping terrain, followed by on community comprises an important fodder tree for honey lower sloping terrain (Table 9). Te slope has been identifed production, which contributes to the means of livelihood for as one of the topographic factors that infuence species local communities in the area [94]. diversity and richness [95, 96, 107]. Tis result agrees with P. altissima was the dominant and signifcant in- the fndings of Wondie et al. [108], who stated that forests dicator species in community 2 (Table 6) and is among grow favourably on intermediate sloping terrain. Te in- the indicator species in the upper canopy of the moist crease in species richness and diversity at intermediate Afromontane forest [92, 93], while L. fraxinifolius was sloping terrain may be due to soil moisture availability and also the dominant species and signifcant indicator edaphic factors (soil nutrients and low soil erosion) species in the middle canopy. [109, 110]. A. toxicaria is the dominant species and the distinctive species of moist Afromontane forests in the middle canopy, whereas Celtis toka was the other dominant and signifcant 4.4. Relationship between Community Types with Environ- indicator species in community 3 and is among the indicator mental and Disturbance Factors. Studies focusing on the species of moist Afromontane forests in the middle canopy relationship between plant communities with environ- [92, 93]. Te community had the highest number of plant mental and disturbance factors have become increasingly species compared to other communities. Tis could be due important in understanding the ecology of forest commu- to their location away from human disturbances since they nities [111–115]. Plant species distribution patterns and plant community formation are highly infuenced by mi- are located in difcult terrain such as sloppy and deep gorges [95, 96]. croclimate, edaphic, and anthropogenic factors [41, 93, 116–119]. Plants form a community when a plant Dracaena afromontana and Cyathea manniana were the dominant and indicator species with a signifcant indicator species repeatedly appears in a similar environment. Simi- value in this community, and they are among the distinctive larly, it is the classifcation of plant communities, which species in the middle canopy of the moist Afromontane temporally and spatially determines the boundaries of forest [92, 93]. Te D. afromontana-C. manniana com- a plant community [120–122]. Each community has its own munity type comprises diferent species as compared to distribution area with a specifc combination or preference other plant communities, which are found at higher ele- of environmental variables. vations. Te species composition diference in the com- Relatively, the elevation was the most signifcant envi- munity may be due to an environmental and anthropogenic ronmental factor infuencing and describing the community type distribution [42, 81, 123–129]. Accordingly, the vari- factor [95, 96]. Te Shannon diversity index was extended from 3.5 to ation of plant communities was also closely related to other environmental, disturbance, and site factors, including 3.8 in four plant community types, which indicates the existence of high diversity in MFBR. Te plant communities herbaceous cover, harvesting index, slope, pH, and silt showed a minor variation in their species richness, diversity, (Table 10). and evenness. Somewhat, community types such as Disturbance (illegal logging) afects the distribution of B. unijugata-P. alnifolia and P. altissima-L. fraxinifolius were plant communities by hindering seedling establishment and higher in species richness, while community types regeneration in tropical forests [130–133]. It can also assist in A. toxicaria-C. toka and D. afromontana-C. manniana had the ground layer plant species growth by facilitating the availability of light [134, 135]. the lowest species richness. Tis variation in species richness among communities could be due to the variations of to- Te correlation among environmental, disturbance, and site factors governs the pattern of species distribution in the pography, anthropogenic impact, climate, and edaphic factors [97, 98]. Each community has its own distribution plant communities of MFBR (Table 12). Tere was a positive correlation between the elevation and organic matter. Te area with a specifc combination of environmental variables [27, 99–101]. high amount of organic matter at higher elevations can be International Journal of Forestry Research 13 edaphic gradients,” Journal of Ecology, vol. 100, no. 1, attributed to a low rate of decomposition due to relatively pp. 253–263, 2012. low temperatures [113]. High organic matter content in soils [3] S. Harrison, M. J. Spasojevic, and D. Li, “Climate and plant of higher elevations was also reported in other Afromontane community diversity in space and time,” Proceedings of the forests of Ethiopia [136]. National Academy of Sciences, vol. 117, no. 9, pp. 4464–4470, 5. Conclusion [4] C. A. Baldeck, K. E. Harms, J. B. Yavitt et al., “Soil resources and topography shape local tree community structure in Tis study was aimed to identify the type of plant com- tropical forests,” Proceedings of the Royal Society B: Biological munity in MFBR and determine plant community distri- Sciences, vol. 280, no. 1753, Article ID 20122532, 2013. bution patterns in response to environmental disturbance [5] Z. H. Zhang, G. Hu, and J. Ni, “Efects of topographical and factors (elevation, slope, disturbance, and physical and edaphic factors on the distribution of plant communities in chemical composition of soil) in MFBR. MFBR is rich in two subtropical karst forests, southwestern China,” Journal endemic plant species. Our study identifed four plant of Mountain Science, vol. 10, no. 1, pp. 95–104, 2013. community types and indicated the elevation as a major [6] M. V. Odgaard, P. K. Bøcher, T. Dalgaard, J. E. Moeslund, environmental factor for determining the community type and J.-C. 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International Journal of Forestry Research – Hindawi Publishing Corporation

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Effects of Environmental and Disturbance Factors on Plant Community Distribution in Tropical Moist Afromontane Forests, South-West Ethiopia

Effects of Environmental and Disturbance Factors on Plant Community Distribution in Tropical Moist Afromontane Forests, South-West Ethiopia

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Effects of Environmental and Disturbance Factors on Plant Community Distribution in Tropical Moist Afromontane Forests, South-West Ethiopia

Effects of Environmental and Disturbance Factors on Plant Community Distribution in Tropical Moist Afromontane Forests, South-West Ethiopia

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