Floristic composition along the agro-climatic zone
Tables 1 and 2 presented descriptive statistics and mean differences in species diversity across the agro-climatic zones of the BMNP. Overall species richness did not show significant differences (p < 0.05) among the agro-climatic zones, with the highest mean value of 94 species recorded in the cool moist mid highland zone. The sub-moist mid highland zone followed with 85 species, while the cold humid afro-alpine zone had the lowest species richness of 72 species. However, the mean (± SE) species richness per 400 m² plot was significantly higher in the cool moist mid highland compared to the sub-moist mid highland (Tables 1). The species richness in the cold humid afro-alpine zone did not significantly differ from the other two agro-climatic zones.
Herbaceous species richness was significantly higher (p < 0.05) in both the cool moist mid highland (60 species) and the cold humid afro-alpine zone (43 species) compared to the sub-moist mid highland (18 species). Conversely, woody species richness was significantly higher in the sub-moist mid highland (76 species) than in the cool moist mid highland (31 species) and the cold humid afro-alpine zone (29 species) at p < 0.05. The number of endemic species was significantly higher (p < 0.05) in the cool moist mid highland (14 species) compared to the cold humid afro-alpine zone (11 species) and the sub-moist mid highland (2 species).
Asteraceae was the most species-rich family in the cool moist mid highland and cold humid afro-alpine zones, with 21 and 16 species respectively. In the sub-moist mid highland, Celasteraceae and Rutaceae were the richest families, each with 7 species. Helichrysum was the most species-rich genus in the cool moist mid highland and cold humid afro-alpine zones, with 6 and 5 species respectively. In the sub-moist mid highland, Hippocratea and Olea were the richest genera, each with 3 species.
Table 1
Descriptive statistics on species richness in agro-climatic zones of the BMNP
Statistics | ACZ 1 | ACZ 2 | ACZ 3 |
Min | 5 | 6 | 7 |
Max | 21 | 23 | 21 |
Mean | 10.35 | 12.40 | 11.29 |
Standard deviation | 4.04 | 4.19 | 3.79 |
Standard error | 0.58 | 0.60 | 0.55 |
df | 143 | | |
MS | 50.13 | | |
F | 3.12 | | |
P-value | 0.047 | | |
Note: Min: minimum and Max: maximum, df: degrees of freedom, MS: Mean squares, ACZ 1: Sub-moist mid highland, ACZ 2: Cool moist mid highland, ACZ 3: Cold humid afro-alpine zone. |
The mean (± SE) Shannon diversity and Pielou’s evenness indices did not show significant differences (p < 0.05) among the agro-climatic zones. However, both indices were highest in the cool moist mid highland, followed by the sub-moist mid highland and the cold humid afro-alpine zone (Table 2). The computed Sorensen’s similarity index indicated that the sub-moist mid highland shared 31% floristic similarity with the cool moist mid highland and 4% similarity with the cold humid afro-alpine zone. In comparison, the cool moist mid highland and the cold humid afro-alpine zone shared 7% floristic similarity. Overall, the floristic similarity among the agro-climatic zones was weak, indicating distinct plant communities within each zone.
Table 2
Mean differences of species richness, diversity, and evenness
Agro-climatic zones | S | H’ | J |
ACZ 1 | 10.35 ± 0.58b | 2.71 ± 0.05 | 0.77 ± 0.02 |
ACZ 2 | 12.40 ± 0.60a | 3.02 ± 0.03 | 0.86 ± 0.01 |
ACZ 3 | 11.29 ± 0.55ab | 2.69 ± 0.09 | 0.81 ± 0.05 |
Mean | 11.35 ± 0.57 | 2.81 ± 0.06 | 0.81 ± 0.02 |
F | 3.12 | 0.75 | 1.65 |
P-value | < 0.05 | 0.051 | 0.072 |
Note: S: Species richness, H’: Shannon diversity index, J: Pielou’s evenness index, ACZ 1: Sub-moist mid highland, ACZ 2: Cool moist mid highland, ACZ 3: Cold humid afro-alpine zone. |
Vegetation structure along the agro-climatic zone
The structural characteristics of woody species were analyzed in the sub-moist mid highland and cool moist mid highland zones, as woody species with a DBH > 2 cm were absent in the cold humid afro-alpine zone. The mean woody species density did not show a significant difference (p < 0.05) between the sub-moist mid highland and the cool moist mid highland (Table 3). However, the overall density of woody species was higher in the sub-moist mid highland, with 914 stems per hectare, compared to 653 stems per hectare in the cool moist mid highland.
Table 3
Mean differences of vegetation structure parameters
Agro-climatic zones | Density (stems ha− 1) | BA (m2 ha− 1) | DBH (cm) | Height (m) |
ACZ 1 | 65.28 ± 11.90 | 6.50 ± 1.76 | 49.95 ± 1.34a | 23.02 ± 0.53 |
ACZ 2 | 46.65 ± 12.15 | 5.70 ± 0.67 | 44.50 ± 1.42b | 21.94 ± 0.61 |
Mean | 55.97 ± 12.02 | 6.10 ± 1.21 | 47.23 ± 1.38 | 22.48 ± 0.57 |
F | 1.20 | 2.31 | 7.67 | 1.74 |
P-value | < 0.056 | 0.082 | < 0.05 | 0.063 |
Note: BA: Basal area, DBH: Diameter at breast height, ACZ 1: Sub-moist mid highland, ACZ 2: Cool moist mid highland. |
The mean basal area of woody species did not show significant differences (p < 0.05) between the two agro-climatic zones (Table 3). Nonetheless, the overall basal area was higher in the sub-moist mid highland, with 91.16 m² ha⁻¹ compared to 79.10 m² ha⁻¹ in the cool moist mid highland. Conversely, the mean DBH of woody species in the sub-moist mid highland was significantly higher (p < 0.01) than in the cool moist mid highland (Table 3). The mean height of woody species did not significantly differ (p < 0.05) between the two agro-climatic zones.
Woody species with a DBH > 2 cm across the agro-climatic zones were classified into 14 DBH classes. The distribution pattern of woody species across these classes exhibited an inverted J-shape beyond the second DBH class. Most individuals were concentrated in the third DBH class (21–30 cm), with a sharp decline in density towards higher DBH classes (Fig. 4a and b). The sub-moist mid highland recorded the highest density of woody species, with approximately 12.38% of the species having a DBH > 100 cm, contributing to the higher basal area in this zone. The largest recorded DBH was 187.58 cm for Juniperus procera in the cool moist mid highland and 129.62 cm for Syzygium guineense in the sub-moist mid highland.
The height distribution of woody species in both agro-climatic zones was categorized into 12 classes. Consistent with the DBH distribution, the height distribution of woody species followed an inverted J-shape pattern (Fig. 5a and b). The tallest recorded tree species was Olea capensis, reaching 52 meters in the sub-moist mid highland, while Juniperus procera reached 45 meters in the cool moist mid highland.
Croton macrostachyus was the most frequently occurring woody species in the sub-moist mid highland, present in 81% of the sample plots, followed by Podocarpus falcatus at 63% and Syzygium guineense at 48% frequency. In contrast, Juniperus procera was the most frequently occurring species in the cool moist mid highland, found in 79% of the sample plots, followed by Hypericum revolutum at 71% and Hagenia abyssinica at 60% frequency. The dominance of woody species, as indicated by the Importance Value Index (IVI), ranged from 0.77 to 38.45 in the sub-moist mid highland, and from 1.34 to 96.47 in the cool moist mid highland (Table 4). The most dominant species in the sub-moist mid highland was Croton macrostachyus with an IVI of 38.45, followed by Syzygium guineense at 27.22, and Podocarpus falcatus at 25.16. In the cool moist mid highland, Juniperus procera was the most dominant species with an IVI of 96.47, followed by Hypericum revolutum at 58.47, and Hagenia abyssinica at 52.18.
Table 4
IVI of top 10 woody species in ACZ 1 and ACZ 2 with their corresponding relative values
Species | Rde (%) | Rfr (%) | Rdo (%) | IVI |
| ACZ 1 | ACZ 2 | ACZ 1 | ACZ 2 | ACZ 1 | ACZ 2 | ACZ 1 | ACZ 2 |
Juniperus procera | 0.18 | 24.12 | 0.63 | 23.60 | 1.05 | 48.75 | 1.86 | 96.47 |
Olea europae | 0.91 | 3.53 | 2.52 | 6.21 | 1.84 | 11.68 | 5.26 | 21.42 |
Myrsine melanophoeos | 22.14 | 5.03 | 5.49 | 2.99 | 0.82 | 0.31 | 28.45 | 8.33 |
Hypericum revolutum | - | 32.29 | - | 21.12 | - | 5.06 | - | 58.47 |
Hagenia abyssinica | - | 8.83 | - | 18.01 | - | 25.34 | - | 52.18 |
Rapanea melanophloeos | - | 7.71 | - | 13.66 | - | 5.50 | - | 26.94 |
Croton macrostachyus | 21.56 | - | 12.26 | - | 4.63 | - | 38.45 | - |
Syzygium guineense | 5.98 | - | 7.23 | - | 14.01 | - | 27.22 | - |
Podocarpus falcatus | 11.78 | - | 9.43 | - | 3.95 | - | 25.16 | - |
Ehretia cymosa | 7.79 | - | 5.35 | - | 8.09 | - | 21.22 | - |
Note: Rde: Relative density, Rfr: Relative frequency, and Rdo: relative dominance, ACZ 1: Sub-moist mid highland, ACZ 2: Cool moist mid highland. |
Woody species regeneration status along the agro-climatic zone
The mean densities of seedlings, saplings, and mature trees were significantly different between the sub-moist mid highland and cool moist mid highland at p < 0.05 (Table 5). In the sub-moist mid highland, the total densities were 5363 seedlings ha− 1, 2930 saplings ha− 1, and 940 mature trees ha− 1. In contrast, the cool moist mid highland had total densities of 3668 seedlings ha− 1, 1921 saplings ha− 1, and 627 mature trees ha− 1. The highest seedling and sapling densities in the sub-moist mid highland were recorded for Croton macrostachyus, with 967 seedlings ha− 1 and 629 saplings ha− 1. In the cool moist mid highland, the highest densities were for Juniperus procera (190 seedlings ha− 1) and Hypericum revolutum (302 saplings ha− 1).
Table 5
Mean differences in the seedling, sapling, and mature tree
Agro-climatic zones | Seedling | Sapling | Mature tree |
ACZ 1 | 1592.40 ± 147.46a | 884.84 ± 81.94a | 297.01 ± 27.50a |
ACZ 2 | 1089.11 ± 142.59b | 580.13 ± 75.95b | 198.11 ± 25.94b |
Mean | 1340.75 ± 145.02 | 732.48 ± 78.94 | 247.56 ± 26.72 |
F | 8.43 | 7.25 | 4.25 |
P-value | < 0.01 | < 0.05 | < 0.05 |
Driving factors for plant composition and structure change
Changes in plant species composition and structure across the designated agro-climatic zones of the BMNP are driven by both direct and indirect factors. Over the past 40 years, direct drivers such as overgrazing, illegal logging, agricultural development, and settlement expansion have been prominent, particularly affecting the cool moist mid highland and sub-moist mid highland zones. The Gaysay grassland in the cool moist mid highland, which is rich in endemic and medicinal plant species, faces critical challenges from unrestricted livestock grazing. In the cold humid afro-alpine zone, recurrent fires and grazing are significant stresses. Indirect factors, including rapid population growth and poverty, exacerbate these impacts. Field observations and interviews identified seven key drivers of landscape change, with farmland and settlement expansions as the leading proximate factors, while population growth and climate issues were significant underlying causes. These pressures have led to extensive human activity, notably illegal settlements from the park's northern sides, intensifying farming and overgrazing, particularly in the Gaysay grassland in the cool moist mid highland.
Table 6
Driving factors of landscape change in the BMNP
Drivers | Percent (%) | Rank |
Farmland expansion | 30.46 | 1 |
Settlement expansion | 19.41 | 2 |
Population growth | 14.56 | 3 |
Overgrazing | 12.43 | 4 |
Forest fire | 11.34 | 5 |
Illegal logging | 7.62 | 6 |
Climate related problems | 4.18 | 7 |